Dicotyledons: Rosids (Illustrated Handbook of Succulent Plants) [2 ed.] 3030934918, 9783030934910

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Illustrated Handbook of Succulent Plants

Urs Eggli · Reto Nyffeler Editors

Dicotyledons: Rosids Second Edition

Illustrated Handbook of Succulent Plants Series Editors Urs Eggli, Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland Heidrun E. K. Hartmann, Hamburg, Germany

During evolution, many plant species have adapted to life in seasonally dry and arid environments by developing the ability to store water in their stems, leaves, and/or underground organs – this phenomenon is called “succulence.” Succulent plants can be found in more than 70 plant families. The Illustrated Handbook of Succulent Plants (excluding cacti and orchids) was first published in six volumes in 2001–2003. A second revised edition, of which the first volume (treating the family Aizoaceae) was published in 2017, and the second (treating the monocotyledons) in 2020, provides a taxonomical treatment of all estimated 11,000 taxa of succulents (excluding orchids). In addition to the volumes on Monocotyledons and Dicotyledons, separate volumes are devoted to those families with an especially great diversity of succulent species, namely, Aizoaceae, Apocynaceae, Cactaceae, and Crassulaceae. Each volume is organized by alphabet of families, genera, and species, with comprehensive synonymies added at all levels. Detailed descriptions are given for all accepted taxa, together with data on the distribution and typification, and references to the most important literature. Where necessary, information on ecology, ethnobotany, history, etc. is added, and in many places, proposed relationships are critically discussed. Over 2000 superb color photographs complete this inventory of succulent plants.

Urs Eggli • Reto Nyffeler Editors

Dicotyledons: Rosids Second Edition

With 526 Figures and 16 Tables

Editors Urs Eggli Sukkulenten-Sammlung Zürich Grün Stadt Zürich Zürich, Switzerland

Reto Nyffeler Institut für Systematische und Evolutionäre Botanik (ISEB) Universität Zürich Zürich, Switzerland

Illustrated Handbook of Succulent Plants ISBN 978-3-030-93491-0 ISBN 978-3-030-93492-7 (eBook) https://doi.org/10.1007/978-3-030-93492-7 1st edition: © Springer-Verlag Berlin Heidelberg New York 2002 © Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are solely and exclusively licensed 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, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Preface to the Second Edition

Succulents are an attractive group of plants that have for centuries attracted the interest of botanists and gardeners alike, both professionals and hobbyists. During this time, a vast body of specialized books and especially journal papers has accumulated that contains our knowledge of these fascinating plants. Over time, there were several efforts to collate the existing knowledge and information into lexica devoted to succulent plants, and thus allow an ordered presentation of the diversity of succulents. The publication of the first edition of the present series of Illustrated Handbooks of Succulent Plants in the years 2001–2003 in six volumes marked a milestone in this field, even though coverage remained incomplete since the families Bromeliaceae and Cactaceae were not included. The past 20 years have seen a continued, and in many fields, revived interest in all aspects of succulent plant biology, and the handbook series soon became a widely accepted taxonomic reference for ordering the biodiversity of succulents. Twenty years is a long stretch of time for any lexicon, and much new data has been generated by recent research in all fields of botany. During this time, especially molecular studies have led to a much improved understanding of the living diversity of plants, and a firm phylogenetic backbone is now in place for almost all major groups. The past more than 20 years have also seen an important change in the way taxonomical studies are conducted, that is, an increasingly common replacement (or at least support) of intuition-based interpretations with data-derived conclusions. We are now well aware that many of the characters traditionally used to define taxonomical units are homoplasious (i.e., have evolved repeatedly in parallel), and thus are of limited or no value to understand evolutionary pathways. Such increase in knowledge has led to important changes in the circumscription or affiliation of many groups. Such changes often meet with considerable antipathy from many user groups, but when they are based on firm data derived from adequate sampling depth, they will become the accepted standards of future times. For this handbook series, a phylogenetic approach, based on the most recent available standard phylogenetic framework, has been selected. In cases where the available data is inconclusive, or contradictory, established classifications are usually preferably followed, with the necessary cautious comments.

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For this second edition, it was decided to split the original one-volume treatment of the Dicotyledons into two separate sets of volumes, one (the present set of volumes) covering the families of the Rosids, and the other (publication estimated for 2025) covering all the remaining families of the Dicotyledons plus those of the Magnoliids. The existing entries from the first edition have been thoroughly checked and updated, including revised classifications and new taxa. Especially the discussions of classifications have been expanded considerably within a phylogenetic context, but also new data, for example, on ecology or ethnobotany, have been added. For most groups, the original authors have been able to provide these updated treatments, and we are most grateful to our colleagues for their continued support of the project. For some groups, new contributors had to be sought as the original authors have died, or stepped back, or could not be contacted. We are delighted about the willingness of the new authors to adopt these “orphaned” groups. The overall layout of the volume has changed considerably in comparison with the original edition, while the sequence of information presented for each taxon has largely remained the same. We hope that the new layout, which places the illustrations near the text to which they belong, will prove helpful for the users. Our sincere thanks go to all the many contributors, both of texts and illustrations, for timely delivery of materials, and for the patience needed to see the project come to fruition. The coronavirus pandemic placed a considerable burden on many of us, and the more we would like to express our gratitude for all the help we received. Words of thanks moreover go to the entire staff at the Sukkulenten-Sammlung Zürich for providing support and time and to the team at the library of the Botanical Garden of the University Zürich for their continued help. Sukkulenten-Sammlung Zürich, Grün Stadt Zürich Zürich, Switzerland Institut für Systematische und Evolutionäre Botanik (ISEB) Universität Zürich Zürich, Switzerland February 2023

Urs Eggli

Reto Nyffeler

From the Preface to the First Edition

Handbooks devoted to succulent plants (including cacti) have a long-standing tradition, and the demand for updated editions is a good indication of the high degree of interest that exists in this fascinating group of plants. While first handbooks devoted to the family Cactaceae were already published in the 19th century, the first handbook dealing with the so-called ‘other’ succulents was authored by Hermann Jacobsen and published in 1954–1955, then called “Handbuch der sukkulenten Pflanzen”. A revised and enlarged English edition was made available in 1959 and was repeatedly reprinted in the following years. A major step towards a compact handbook including short descriptions, full synonymy and hundreds of illustrations was the publication of the first edition of Hermann Jacobsen’s celebrated “Sukkulentenlexikon” in 1970, followed by the English edition in 1975, and a revised German edition in 1981, finished by Klaus Hesselbarth following Hermann Jacobsen’s death in August 1978. The demand for updated compact information on succulent plants has not diminished since then, and contrary to Hermann Jacobsen’s opinion (preface to the German edition 1970) that the number of papers dealing with succulent plants was on the decline, the interest in succulent plants was growing, both among hobbyists and among botanists. The results are numerous new taxa in many families, and many monographs of previously little-known groups have been published over the years. Accordingly, a need for a “New Lexicon” was beginning to be felt, both for ‘other’ succulents and for cacti. After some informal initial discussions held among various botanists and interested specialist collectors, and faced with the desire of Gustav Fischer Verlag Jena, the publisher of both Backeberg’s “Kakteenlexikon” and Jacobsen’s “Sukkulentenlexikon”, to prepare revised editions, a meeting was called for during the 1990-congress of the International Organization for Succulent Plant Study (IOS) in Zürich. There was a general feeling that the situation presented a unique occasion to produce a set of authoritative volumes embracing all succulent plants (except orchids), and produced according to a rigid set of common standards adopted to suit all families involved. It was planned that the project, informally termed the “New IOS Succulent Plant Lexicon”, should consist of three volumes, one devoted to the Cactaceae, one to the Aizoaceae, and one to the remaining succulent taxa from various families.

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From the Preface to the First Edition

For various reasons, the Lexicon project took much longer to be completed than anticipated. The Cactaceae part had not become available in time for inclusion in the first handbook edition. In the case of the Aizoaceae, time permitted a much more complete treatment of the family, which was published in two volumes. Moreover, it soon became apparent that a single volume will not be sufficient to cover the vast array of the “remaining succulents”. Accordingly, they were split into four volumes (Monocotyledons, Dicotyledons, Asclepiadaceae (¼ Apocynaceae in the revised edition) and Crassulaceae).

General Introduction to the Series

Setup of the Handbook Series The present series of handbooks is devoted to a lexicographic treatment of the known diversity of succulent plants (excluding pteridophytes and orchids, see below). The APG IV classification (APG 2016) is used as the phylogenetic framework at the rank of order and supraordinal taxa, while the slightly modified APG III-classification (APG 2009) as defined by Nyffeler & Eggli (2010a) is used at the rank of family.

What is a Succulent? It is a continuing challenge to define what constitutes a succulent plant – at least in view of the several competing definitions (see Eggli & Nyffeler 2009 for a review and a modern definition). For the purpose of this handbook, a pragmatic approach has been selected. Apart from the multitude of unambiguous succulents, borderline cases are included as well, especially when they occur sympatrically with undisputed succulents in more or less semi-arid regions, and consequently show some degree of xerophytic adaptation, and more so when the species in question are encountered in cultivation together with other succulents. This, then, includes many of the so-called caudex and pachycaul plants popular in cultivation in public and private collections alike. Other borderline cases included are a number of bulbous and rhizomatous groups, where examples from several genera are covered, as well as some weakly developed leaf succulents from e.g. the Begoniaceae and Melastomataceae (e.g. Medinilla), and several other families. On the other hand, purely halophytic succulents (such as Salicornia) are omitted from this series of Handbooks since they are as a whole neither adapted to climatically arid or semi-arid conditions (see Ogburn & Edwards 2010 for an explanation) nor encountered in collections devoted to succulent plants. Finally, the family Orchidaceae is excluded (apart from a short general overview), despite the uncontested claim to succulence of many taxa. A preliminary count (Nyffeler & Eggli 2010a) showed some 2200 succulent species for this family, and a more detailed census (see the family treatment in the Monocot volume published 2020) arrives at a minimum count of 4,500 ix

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General Introduction to the Series

succulent species – this makes the Orchidaceae the most important family containing succulents in term of species number.

Succulence Amongst Flowering Plants Succulence, without any doubt, has evolved repeatedly and independently in many clades of the Angiosperms. Succulence thus illustrates in an exemplary manner how adaptations to a given set of ecological conditions appear times and again throughout the evolution of the Plant Kingdom (and likely has disappeared again in numerous cases, but as no macrofossils of succulents are known, this will be difficult to prove). Figure 1 illustrates the occurrence of succulence throughout the clades of the Angiosperms. Succulent plants are known to occur in 31 out of a total of 64 orders accepted. While some orders only count with a small number of succulent representatives, succulence is a more prevailing character in others. A notable concentration of succulents is to be found in the order Caryophyllales, and its suborder Portulacineae embraces almost exclusively succulent taxa (Nyffeler & Eggli 2010b).

Division of the Handbook into Individual Volumes The handbook is divided into individual volumes that either cover an entire individual family, or a coherent group of families. The planned break-down into volumes is as follows (see also Fig. 1): • Monocotyledons: Eggli, U. & Nyffeler, R. (eds.): Illustrated Handbook of Succulent Plants. Monocotyledons. Published 2020. • Dicotyledons A: Eggli, U. (ed.): Illustrated Handbook of Succulent Plants. Magnoliids and Dicotyledons (excluding Rosids). • Dicotyledons B: Eggli, U. (ed.): Illustrated Handbook of Succulent Plants: Dicotyledons: Rosids. This volume. • Aizoaceae: Hartmann, H. E. K. (ed.): Illustrated Handbook of Succulent Plants: Aizoaceae. Volumes 1 and 2. Published 2017. • Apocynaceae: Meve, U. & Eggli, U. (eds.) Illustrated Handbook of Succulent Plants: Apocynaceae. • Cactaceae: NN. (ed.): Illustrated Handbook of Succulent Plants: Cactaceae. • Crassulaceae: U. Eggli (ed.): Illustrated Handbook of Succulent Plants: Crassulaceae. For ease of reference, an alphabetical list of families, together with the orders to which they belong, and the volume of the series which covers them, is also supplied (Table 1).

General Introduction to the Series

xi Dicot B Dicot A Monocot

MAG LILIANAE Fabidae

Rosidae

Malvidae Lamiidae Campanulidae

Asteridae

ASTERANAE

EUDICOTYLEDONEAE

ROSANAE

Fig. 1 Summary phylogeny of the Angiosperms, based on APG IV (2016), Moore & al. (2010a) and Soltis & al. (2011). The estimated number of succulent species (modified from Nyffeler & Eggli 2010a) is indicated to the right of the order names. Supraordinal names follow Soltis & al. (2011) and Chase & Reveal (2009; their

Amborellales Nymphaeales Austrobaileyales Chloranthales Canellales Piperales 100 Laurales Magnoliales Acorales Alismatales 1 Petrosaviales Dioscoreales 7 Pandanales Liliales Asparagales 5973 Arecales Poales 500 Commelinales 25 Zingiberales Ceratophyllales Ranunculales 10 Proteales Trochodendrales Buxales Gunnerales Saxifragales 1440 Vitales 68 Zygophyllales 6 Celastrales Malpighiales 1014 Oxalidales 41 Fabales 26 Rosales 32 Cucurbitales 147 Fagales Geraniales 196 Myrtales 34 Crossosomatales Picramnales Sapindales 37 Huerteales Malvales 41 Brassicales 34 Dilleniales Berberopsidales Santalales 3 Caryophyllales3991 Cornales 1 Ericales 25 Icacinales 2 Metteniusales Garryales Boraginales Vahliales Solanales 90 Gentianales 1210 Lamiales 220 Aquifoliales Escalloniales Asterales 140 Bruniales Apiales 8 Paracryphiales Dipsacales

Crassulaceae

Aizoaceae Cactaceae

Apocynaceae

Lilianae conform to the Monocotyledoneae of Soltis & al., their Rosanae to the Superrosidae, and their Asteranae to the Superasteridae). MAG Magnolianae. The division of the Handbook series into individual volumes is shown on the right of the cladogram. Dicot B ¼ this volume. (Diagram by Urs Eggli)

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Table 1 Alphabetical list of families with their assignments to orders, and their assignments to the individual volumes of this Handbook series Family Agavaceae Aizoaceae Amaranthaceae Amaryllidaceae Anacampserotaceae Anacardiaceae Anthericaceae Apiaceae Apocynaceae Araceae Araliaceae Asparagaceae Asphodelaceae Asteraceae Balsaminaceae Basellaceae Begoniaceae Bixaceae Brassicaceae Bromeliaceae Burseraceae Cactaceae Campanulaceae Capparaceae Caricaceae Clusiaceae Commelinaceae Convolvulaceae Crassulaceae Cucurbitaceae Cyperaceae Didiereaceae Dioscoreaceae Doryanthaceae Ericaceae Eriospermaceae Euphorbiaceae Fabaceae Fouquieriaceae Francoaceae Geraniaceae Gesneriaceae Halophytaceae Hyacinthaceae Icacinaceae Lamiaceae

Order Asparagales Caryophyllales Caryophyllales Asparagales Caryophyllales Sapindales Asparagales Apiales Gentianales Alismatales Apiales Asparagales Asparagales Asterales Ericales Caryophyllales Cucurbitales Malvales Brassicales Poales Sapindales Caryophyllales Asterales Brassicales Brassicales Malpighiales Commelinales Solanales Saxifragales Cucurbitales Poales Caryophyllales Dioscoreales Asparagales Ericales Asparagales Malpighiales Fabales Ericales Geraniales Geraniales Lamiales Caryophyllales Asparagales Icacinales Lamiales

Monocot (published 2020) 

Dicot A

Dicot B (Rosids) (this volume)

Separate volume 

                                            (continued)

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Table 1 (continued) Family Lentibulariaceae Limeaceae Loasaceae Loranthaceae Malvaceae Melastomataceae Meliaceae Menispermaceae Montiaceae Moraceae Nyctaginaceae Orchidaceae Oxalidaceae Passifloraceae Pedaliaceae Phyllanthaceae Phytolaccaceae Piperaceae Plantaginaceae Poaceae Portulacaceae Rubiaceae Ruscaceae Santalaceae Sapindaceae Saxifragaceae Solanaceae Talinaceae Tropaeolaceae Urticaceae Vitaceae Zygophyllaceae

Order Lamiales Caryophyllales Cornales Santalales Malvales Myrtales Sapindales Ranunculales Caryophyllales Rosales Caryophyllales Asparagales Oxalidales Malpighiales Lamiales Malpighiales Caryophyllales Piperales Lamiales Poales Caryophyllales Gentianales Asparagales Santalales Sapindales Saxifragales Solanales Caryophyllales Brassicales Rosales Vitales Zygophyllales

Monocot (published 2020)

Dicot A    

Dicot B (Rosids) (this volume)

Separate volume

                           

References APG [Angiosperm Phylogeny Group] (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Bot. J. Linn. Soc. 161(2): 105–21. https://doi.org/10.1111/ j.1095-8339.2009.00996.x. APG [Angiosperm Phylogeny Group] (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Bot. J. Linn. Soc. 181(1): 1–20. https://doi.org/10.1111/ boj.12385.

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Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions – succulence as an adaptive strategy. Bradleya 27: 13–36. https://doi.org/10. 25223/brad.n27.2009.a10. Nyffeler, R. & Eggli, U. (2010a) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/10. 25223/brad.n28.2010.a1. Nyffeler, R. & Eggli, U. (2010b) Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59(1): 227–240. https://doi.org/ 10.1002/tax.591021. Ogburn, R. M. & Edwards, E. J. (2010) The ecological water-use strategies of succulent plants. Advances Bot. Res. 55: 179–225. https://doi.org/10.1016/ S0065-2296(10)55004-3.

How to Use This Handbook

Sequence of Information All information is presented in strictly alphabetical sequence of families, genera and species within each volume. It is therefore easy to directly find the entry for a given species as long as its family placement is known. An alternative way is to use the taxonomic index supplied at the end of each volume. This index contains all the names covered in the volume and indicates the page where a treatment can be found or where the name is mentioned, or in the case of synonyms gives the page reference where the accepted name is treated. For each family the handbook supplies keys to the genera with succulent representatives. Please note that these keys are designed to work for the succulent taxa treated, and do not account for the total variation encountered in a genus. If the family is not known when trying to identify a specimen, readers are referred to general botanical books that include keys to plant families. Rowley (1980) and Eggli (2008) provided keys for flowering and non-flowering succulents, and Geesink & al. (1981) produced a well-known book of keys to all flowering plants worldwide.

Scope of Information Presented Families The family names adopted are always conforming to the standard form (ending in -aceae); alternative names (such as Cruciferae for Brassicaceae) are not used. Synonyms at family rank are indicated as completely as possible, and at least cover those synonymous names that include succulents. The order to which the family belongs is also given. Within each family, the genera are treated in alphabetical sequence. Some genera of minimal importance or with borderline succulence are only mentioned or at the most described, but no individual species are treated. The family description characterizes the family as a whole, which often includes much more variation than that observed amongst its succulent representatives. This is supplemented by notes on the distribution, phylogeny,

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classification and economic importance of the family, and the occurrence of succulence if this is not a general feature of the family as a whole. Also, a key to genera with succulents is included, and special terminology used for genera and species descriptions is discussed where necessary. The family concept adopted more or less follows APG III and APG IV, as outlined by Nyffeler & Eggli (2010).

Genera and Species The entries for genera and species are structured in the following way: Names of authors are given in full, with initials added where necessary according to Brummitt & Powell (1992) and IPNI. The literature reference of the original description or combination is followed by information on typification (where available, see below). In the case of genera, important taxonomic literature is then cited. This is followed by information on geographical distribution (including notes on ecology for species where available) and an explanation of the etymology for generic names. For genera of larger families, their placement within the family is also indicated. The main part of the entry is made up by the diagnostic description of the taxon, followed by a discussion of its variability, circumscription and/or application where necessary. It should be noted that the descriptions reflect major variability only, but do not usually include all the reported minor variations. For larger genera, an outline of the accepted formal or informal classification is also given, with individual taxa or groups numbered in sequence. These sequence numbers are subsequently given in square brackets at the start of each species description to indicate its placement within the genus. If recent conflicting classifications are available for a given group, this is shortly discussed and the classification adopted is indicated. Further data, e.g. on population biology, ecology, pollination and seed dispersal, or ethnobotany, is also included. Minor spelling variants of epithets are not indicated; instead, the ‘corrected’ spelling is used throughout for accepted names and synonyms. Major spelling variants are treated as synonyms and are listed accordingly.

Infraspecific Taxa Infraspecific taxa are given in strict alphabetic order of rank and name (i.e. ranks in the sequence cultivar, forma, subspecies, variety). The typical infraspecific taxon (i.e. the one repeating the species name) is thus not treated first as in many handbooks, but in its appropriate alphabetical sequence.

Cultivars, Hybrids Cultivars (rank indicated by placing the epithet in simple quotes) are not included on an exhaustive base; only cultivars of exceptional horticultural merit or botanical interest are covered. Cultivars not associated with a species

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are enumerated immediately following the entry for the genus. Cultivars associated with a species name are included under that species, either as an entry of their own (and in the same form as subspecies etc.), or, in the case of cultivars of minor importance, in the form of a short mention in the species discussion. Cultivar nomenclature follows the guidelines of the current edition of the ICN. Formally named hybrid genera (nothogenera) are covered as genera of their own in the proper alphabetical sequence. The same applies to formally named hybrid species (nothospecies) and infraspecific nothotaxa. Hybrids only known with their hybrid formula are either discussed in the generic entry, or mentioned under one or the other of their parent species. Formally named naturally occurring hybrids are usually covered in full including a description. No attempt has been made, however, to include all the numerous named artificial hybrids that originated in cultivation.

Phylogenetic Diagrams Phylogenetic summary cladograms are provided for all major families, as well as for larger genera with a well-established infrageneric phylogeny. The width of the triangles in these cladograms is always proportional to the number of species of the clade, and the proportion of succulent taxa per clade is indicated with a darker shade. The length of the branches leading to each terminal clade has no phylogenetic meaning, i.e. branch length does not indicate “phylogenetic distance” and thus is not indicative of the degree of relationships between clades. An asterisk (*) accompanying a clade name indicates that the entity is paraphyletic.

Descriptions The descriptions are as compact, concise and diagnostic as possible. Characters that do not vary for the group concerned are usually not repeated from the family or genus descriptions. In the case of genera further subdivided, information already presented in the group definitions is also not normally repeated in the descriptions of individual taxa.

Measurements All measurements are given in metric units. Measurements without further qualification always refer to the long axis of the organ described (i.e. length, height etc.); two measurements united with the -sign stand for length  width.

Terminology Special terms used in descriptions are explained when first used; other botanical terminology is not further explained, and the readers are referred to the

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numerous botanical glossaries, of which Stearn (1992) is cited by way of a most important and useful example. A glossary specifically compiled for succulent plant descriptions is that of Eggli (1993).

Typification An attempt is made to include typification data (including lecto- and neotypifications) for all accepted names. The type citations include the country and major administrative unit where the type was collected, the collector(s) and collection number, and the herbaria where material is said or known to be deposited. The herbarium acronyms conform to Index Herbariorum, Ed. 8 (Holmgren & al. 1990) and its current later electronic version. Where more than one herbarium acronym is given, the first relates to the holotype, the others to isotypes (and similarly to lectotype or neotype, and isolectotypes or isoneotypes). Additional information on typification is sometimes added, especially in the case of lecto- or neotypes, but epitypes are usually not cited. If a taxon is based on several syntype collections, this is also indicated, but due to technical shortcomings of the database system used, only one syntype collection is shown.

Nomenclatural Status of Names For all taxa treated, every attempt has been made to use only valid and legitimate names, but this was not achievable in a small number of cases. In the synonym lists, the nomenclatural status (invalid, illegitimate, rejected, incorrect) is indicated by citing the ICN articles violated. An attempt has been made to exclusively use the numbering in the “Melbourne Code” (McNeill & al. 2012). Spelling variants are considered as invalidly published according to ICBN/ICN Art. 61. While invalid names are, strictly speaking, not considered as names in the ICN (see Art. 6.3), they are nonetheless listed (and flagged accordingly with the violated articles of the ICN) in the synonymies in this handbook to draw attention to the fact that they are invalid.

Synonymies The synonymies given for genera and especially species are as complete as possible and include all names recognized as synonyms. The first synonym(s) – if applicable – is/are the basionym and/or later combination(s) for the accepted name of the entry. All combinations of the same basionym are given in sequence of publication year and are united with the -sign to indicate that these are homotypic (nomenclatural) synonyms. Please note that the -sign is only used for combinations based on the same basionym and does not indicate other

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homotypic synonyms (e.g. nomina nova, which are by definition based on the type of the replaced name). All other synonyms are headed with ‘incl.’ to indicate that they are, with the exception of nomina nova as just discussed, taxonomic synonyms (¼ heterotypic synonyms). They are listed in chronological order of their basionyms, and all combinations based on the same basionym are then given in chronological order. For a small number of taxa, the numbers of synonyms are exaggeratingly large, and here, only the more important synonyms are listed, with an appropriate remark in the discussion of the genus or species concerned. Omission of synonyms usually concerns taxa that have suffered from severe oversplitting in past times, e.g. the genus Begonia.

Geographical Names Country names are listed roughly in a North to South and West to East sequence. Every attempt has been made to standardize geographical names (of countries, administrative units, regions, etc.) as far as possible, but there is a surprising amount of change relating to such names. This is specifically the case for the names of the provinces of the Republic of South Africa, which have changed considerably in the 1990ies, especially affecting the former Cape Province, which has been split up into four units (North-West Province, Northern Cape, Western Cape, Eastern Cape). We have tried as best as we could to provide the modern names in the distribution information included in the handbook, but it has been impossible to adjust all the data for type localities, where the name “Cape Prov.” is still used in a small number of cases. Some difficulties were also encountered in a few cases where countries have been amalgamated (as in the case of the former North Yemen and South Yemen) or divided (e.g. Eritrea, formerly part of Ethiopia, Sudan and South Sudan, or the nations that formerly made up Yugoslavia). Full consistency in all these cases was impossible to achieve. In order to save space, geographical directions such as North, South, etc., are always abbreviated (N, S, etc.). Please note that SW Africa indicates ‘southwestern Africa’ and not the former Southwest-Africa (now Namibia). Similarly, S Africa indicates ‘southern Africa’ and not the Republic of South Africa, for which the abbreviation RSA is always used.

Literature References Literature references are given for all accepted names. Normally, the publication is cited with a full abbreviation according to the standards defined in BPH2 (Bridson & al. 2004) for periodicals (supplemented with those of Eggli (1985) and Eggli (1998) for specialist succulent plant periodicals), while TL2 of Stafleu & Cowan (1976–1988) and supplements

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(Stafleu & Mennega 1992–2000; Dorr & Nicolson 2008–2009) are followed for book abbreviations (in both cases with some minor exceptions to conserve uniformity). In the running text, literature is cited in the usual way (author(s) and year, sometimes supplemented by a page reference), and full details can be found in the list of references at the end of each family or genus.

Illustrations An attempt has been made to cite at least one readily accessible illustration for each species or infraspecific taxon when no illustrations are included in the literature reference for the accepted name. If the name used in the cited publication differs from the accepted name in the handbook, it is indicated (genus name abbreviated to first letter if identical, specific or infraspecific epithet omitted if identical to the accepted name).

Indication of Authorships of the Handbook Texts For each family and genus, the authorship of the handbook text is indicated at the start of the entry. If more than one author has contributed species entries for a genus, each species entry has its own indication of authorship when this differs from the authorship of the genus as a whole.

References Bridson, G. D. R. [& al. 2004], Townsend, S. T., Polen, E. A. & Smith, E. R. (2004) BPH-2. Periodicals with botanical contents. Constituting a second edition of Botanico-Periodicum Huntianum. Pittsburgh (US): Hunt Institute for Botanical Documentation, Carnegie Mellon University. 2 vols., 1470 pp. Brummitt, R. K. & Powell, C. E. (eds.) (1992) Authors of plant names. A list of authors of scientific names of plants, with recommended standard forms of their names, including abbreviations. Richmond (GB): The Board of Trustees of The Royal Botanic Gardens, Kew. 732 pp. Dorr, L. J. & Nicolson, D. H. (2008–2009) Taxonomic literature. Ruggell (FL): A. R. G. Gantner-Verlag KG. Supplements VII-VIII to Ed. 2; 2 vols. https://doi.org/10.5962/bhl.title.48631. Eggli, U. (1985) A bibliography of succulent plant periodicals. Bradleya 3: 103–119. https://doi.org/10.25223/brad.n3.1985.a6. Eggli, U. (1993) Glossary of botanical terms with special reference to succulent plants. Richmond (GB): British Cactus & Succulent Society. 109 pp. Eggli, U. (1998 [‘1995’]) Bibliography of succulent plant periodicals. Bibliografie casopisu o sukulentních rostlinách. Friciana 60: 139 pp. Eggli, U. (2008) Sukkulenten. Stuttgart (DE): Eugen Ulmer. Ed. 2, 392 pp., ills., keys.

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Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions – succulence as an adaptive strategy. Bradleya 27: 13–36. https://doi.org/10. 25223/brad.n27.2009.a10 Geesink, R. [& al. 1981], Leeuwenberg, A. J. M., Ridsdale, C. E. & Veldkamp, J. F. (1981) Thonner’s analytical key to the families of flowering plants. Den Haag (NL): Leiden University Press. 231 pp. Holmgren, P. K. [& al. 1990], Holmgren, N. H. & Barnett, L. C. (1990) Index Herbariorum. Part I: The herbaria of the world. New York (US): New York Botanical Garden. Ed. 8; 693 pp. Current electronic version: http:// sweetgum.nybg.org/science/ih/ IPNI: The International Plant Name Index, accessible at: http://www.ipni.org/ McNeill, J. [& al. 2012], 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. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Königstein (DE): Koeltz Scientific Books. Nyffeler, R. & Eggli, U. (2010) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/ 10.25223/brad.n28.2010.a1 Rowley, G. D. (1980) Name that succulent. Keys to the families and genera of succulent plants in cultivation. Cheltenham (GB): Stanley Thornes (Publishers) Ltd. 268 pp., ill., keys. Stafleu, F. A. & Cowan, R. S. (1976–1988) Taxonomic literature. Utrecht (NL): Bohn, Scheltema & Holkema, etc. Ed. 2; 7 vols. https://doi.org/10. 5962/bhl.title.48631. Stafleu, F. A. & Mennega, E. A. (1992–2000) Taxonomic literature. Königstein (DE): Koeltz Scientific Books. Supplements I-VI to Ed. 2; 6 vols. https://doi.org/10.5962/bhl.title.48631. Stearn, W. T. (1992) Botanical Latin. Newton Abbot (GB): David & Charles Publishers. Ed. 4; 560 pp.

Contents

Volume 1 Introduction to the Diversity of Succulent Rosids . . . . . . . . . . . . . U. Eggli and Reto Nyffeler

1

Part I

Anacardiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

9

Anacardiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

11

........................

15

Pachycormus ANACARDIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

19

Part II

Begoniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

23

Begoniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

25

................................

27

Part III Bixaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

51

Bixaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

53

Amoreuxia BIXACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

57

Cochlospermum BIXACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

61

Part IV

Brassicaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

63

Brassicaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

65

Operculicarya ANACARDIACEAE U. Eggli

Begonia BEGONIACEAE U. Eggli

xxiii

xxiv

Contents

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71

Heliophila BRASSICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. J. Van Jaarsveld

73

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79

Burseraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

85

Burseraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

87

Beiselia BURSERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

93

Boswellia BURSERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. Eggli

95

................................

97

Aethionema BRASSICACEAE U. Eggli

Lepidium BRASSICACEAE P. I. Forster and U. Eggli Part V

Bursera BURSERACEAE U. Eggli

Commiphora BURSERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 E. J. Van Jaarsveld Part VI

Capparaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

Capparaceae U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

Maerua CAPPARACEAE U. Eggli Part VII

Caricaceae

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

Caricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 U. Eggli Carica CARICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 U. Eggli Jacaratia CARICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 U. Eggli Jarilla CARICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 U. Eggli Vasconcellea CARICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 U. Eggli

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Part VIII

Clusiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

Clusiaceae U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Clusia CLUSIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 U. Eggli Part IX

Cucurbitaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Cucurbitaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 L. E. Newton and U. Eggli Ampelosicyos CUCURBITACEAE L. E. Newton

. . . . . . . . . . . . . . . . . . . . . . . . . 173

Anisosperma CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 177 L. E. Newton and U. Eggli Apodanthera CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 179 L. E. Newton and G. N. Njoroge Brandegea CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183 L. E. Newton Cephalopentandra CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . 185 L. E. Newton Ceratosanthes CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 187 L. E. Newton Citrullus CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191 L. E. Newton Coccinia CUCURBITACEAE L. E. Newton

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195

Corallocarpus CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 201 L. E. Newton and G. N. Njoroge Cucumis CUCURBITACEAE L. E. Newton

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205

Cucurbita CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209 L. E. Newton Cyclantheropsis CUCURBITACEAE L. E. Newton

. . . . . . . . . . . . . . . . . . . . . . . 213

Dendrosicyos CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 215 L. E. Newton Doyerea CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217 L. E. Newton

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Eureiandra CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 L. E. Newton and G. N. Njoroge Gerrardanthus CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . 223 L. E. Newton Halosicyos CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227 L. E. Newton Ibervillea CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 L. E. Newton Kedrostis CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237 L. E. Newton and G. N. Njoroge Linnaeosicyos CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 243 L. E. Newton Marah CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 L. E. Newton Momordica CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 L. E. Newton Neoalsomitra CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 259 L. E. Newton and G. N. Njoroge Parasicyos CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263 L. E. Newton Seyrigia CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 L. E. Newton Telfairia CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269 L. E. Newton and G. N. Njoroge Trochomeria CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 271 L. E. Newton Trochomeriopsis CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . 275 L. E. Newton Xerosicyos CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277 L. E. Newton and G. N. Njoroge Zehneria CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 L. E. Newton and G. N. Njoroge Part X

Euphorbiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285

Euphorbiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287 S. Carter

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Euphorbia EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 S. Carter Jatropha EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 617 S. Carter

Volume 2 Part XI

Fabaceae

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643

Fabaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 645 U. Eggli and R. P. Clark Delonix FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 655 G. Germishuizen and R. P. Clark Dolichos FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 661 G. Germishuizen and R. P. Clark Elephantorrhiza FABACEAE R. P. Clark

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 665

Erythrina FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 669 G. Germishuizen and R. P. Clark Neorautanenia FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 673 G. Germishuizen and R. P. Clark Oberholzeria FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 677 R. P. Clark Pachyrhizus FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 679 G. Germishuizen and R. P. Clark Tylosema FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 683 G. Germishuizen and R. P. Clark Part XII

Francoaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 687

Francoaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 689 U. Eggli Tetilla FRANCOACEAE U. Eggli Part XIII

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691

Geraniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693

Geraniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 U. Meve and F. Albers Monsonia GERANIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 701 U. Meve and F. Albers

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Contents

Pelargonium GERANIACEAE U. Meve and F. Albers Part XIV

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 713

Malvaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 799

Malvaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 801 C. C. Walker Adansonia MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 807 C. C. Walker and U. Eggli Brachychiton MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 821 P. I. Forster, G. P. Guymer, and C. C. Walker Cavanillesia MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 827 C. C. Walker Ceiba MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 835 C. C. Walker Megistostegium MALVACEAE C. C. Walker

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 843

Pseudobombax MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 847 C. C. Walker Sterculia MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 853 P. I. Forster, G. P. Guymer, E. J. Van Jaarsveld, and C. C. Walker Part XV

Melastomataceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 859

Melastomataceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 861 U. Eggli and F. Almeda Fritzschia MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 869 M. J. R. Rocha and D. N. Silva Gravesia MELASTOMATACEAE F. Almeda

. . . . . . . . . . . . . . . . . . . . . . . . . 871

Medinilla MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 875 U. Eggli Merianthera MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . 883 U. Eggli, M. J. R. Rocha, and D. N. Silva Miconia MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 887 U. Eggli, M. J. R. Rocha, and D. N. Silva Pachycentria MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . 895 G. Kadereit and U. Eggli Sonerila MELASTOMATACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 899 U. Eggli

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Part XVI Meliaceae

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 901

Meliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903 U. Eggli Entandrophragma MELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 907 U. Eggli Part XVII

Moraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 911

Moraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 913 C. C. Berg and U. Eggli Dorstenia MORACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 919 M. E. E. Hijman, C. C. Berg, U. Eggli, and L. E. Newton Ficus MORACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 939 C. C. Berg and U. Eggli Part XVIII

Moringaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 947

Moringaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 949 U. Eggli Moringa MORINGACEAE U. Eggli Part XIX

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 953

Oxalidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 961

Oxalidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 963 J. M. H. Shaw Oxalis OXALIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 967 J. M. H. Shaw Part XX

Passifloraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1001

Passifloraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1003 W. J. de Wilde and U. Eggli Adenia PASSIFLORACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1007 W. J. de Wilde and U. Eggli Part XXI

Phyllanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041

Phyllanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1043 S. Carter Phyllanthus PHYLLANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 1047 S. Carter

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Part XXII Sapindaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1051 Sapindaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1053 U. Eggli Erythrophysa SAPINDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1057 U. Eggli Part XXIII

Tropaeolaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1059

Tropaeolaceae U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1061

Tropaeolum TROPAEOLACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 1065 U. Eggli Part XXIV

Urticaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1069

Urticaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1071 C. C. Walker and U. Eggli Laportea URTICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1075 C. C. Walker Obetia URTICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1077 C. C. Walker Pilea URTICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1079 C. C. Walker Part XXV

Vitaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1085

Vitaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1087 B. Descoings, U. Eggli, and L. E. Newton Cissus VITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1093 B. Descoings and L. E. Newton Cyphostemma VITACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1107 B. Descoings and L. E. Newton Part XXVI

Zygophyllaceae

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

Zygophyllaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1127 G. F. Smith and E. Figueiredo Augea ZYGOPHYLLACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1133 G. F. Smith and E. Figueiredo Seetzenia ZYGOPHYLLACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1137 G. F. Smith and E. Figueiredo

Contents

xxxi

Tetraena ZYGOPHYLLACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1139 G. F. Smith and E. Figueiredo Zygophyllum ZYGOPHYLLACEAE G. F. Smith and E. Figueiredo

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

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1145

List of Abbreviations and Symbols

Symbols  † ∅ 

Mathematical “equal,” used for homotypic synonyms Dagger symbol, used for lost type specimens, and to indicate death dates Diameter symbol, used in descriptions Mathematical multiplication symbol, used to connect measurements of length  width, and used to indicate hybrid status  Plus-minus symbol, used to indicate approximate values * (Beneath a taxon name) indication of a paraphyletic taxon

General Abbreviations Art. BCE BG C CE clono

cons. cult. cv. Distr Distr. E e.g. epi esp. Etym excl. fa. fig., figs. Fl.

Article of ICBN/ICN Before Common Era Botanical Garden central Common Era clonotype, i.e., a herbarium specimen derived from the type gathering after the publication of the protologue, and not a “true” type in the sense of botanical nomenclature “conservandum,” to be conserved (taxonomical concept as nom. cons. ¼ conserved name) cultivated cultivar distribution District East, eastern for example epitype (nomenclatural type) especially etymology excluding forma (botanical rank) figure, figures Flora xxxiii

xxxiv

fl. Gr. Herb. holo Hort., hort. I ICBN ICN

ill., ills. illeg. in sched. incl. ING inval. IPNI KG, KGW l.c. lecto Lit ms. Mt., Mts. my mybp N NBG NE neo nom. NW p., pp. p.a. p.p. pers. comm. pl. pp. Prov. publ. q.v. RSA S s.a. s.l. s.n.

List of Abbreviations and Symbols

“floruit,” i.e., lived Greek Herbarium holotype (nomenclatural type) Garden, of gardens Illustration(s) International Code of Botanical Nomenclature (“Vienna Code,” 2006) International Code of Nomenclature for Algae, Fungi, and Plants (“Melbourne Code,” 2012; and “Shenzhen Code,” 2018) illustration, illustrations; illustrated illegitimate (nomenclatural concept as nom. illeg. ¼ illegitimate name) “in schedis,” i.e., in the herbarium including Index Nominorum Genericorum invalid (nomenclatural concept as nom. inval. ¼ invalid name) International Plant Name Index, https://www.ipni.org/ Karroo Garden, Worcester, RSA “loco citato,” i.e., at the place cited lectotype (nomenclatural type) literature manuscript mountain, mountains million years million years before present North, northern National Botanical Garden, RSA North-East, north-eastern neotype (nomenclatural type) “nomen,” i.e., botanical name North-West, north-western page, pages “per annum,” i.e., per year “pro parte,” i.e., partly personal communication plate pages Province published “quod vide,” i.e., which see Republic of South Africa South, southern “sine anno,” i.e., without year “sensu lato,” i.e., in a wide sense “sine numero,” i.e., without (collection) number

List of Abbreviations and Symbols

xxxv

s.s. SE Sect. SEM Ser. spp. ssp. Subgen. Subsect. Subser. SUG SW syn t., tt. unpubl. USA var. W WCSP

ybp

“sensu stricto,” i.e., in the strict/narrow sense South-East, south-eastern section (botanical rank) Scanning Electron Microscope series (botanical rank) several species subspecies (botanical rank) subgenus (botanical rank) subsection (botanical rank) subseries (botanical rank) Stellenbosch University Botanical Garden South-West, south-western syntype (nomenclatural type) “tabula,” i.e., illustration plate/plates unpublished United States of America variety (botanical rank) West, western World Checklist of Selected Plant Families. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet; http:// apps.kew.org/wcsp/ years before present

Abbreviations in Plant Descriptions Anth Ar Ax Bo Br Bra Bri Ca Cal Cap Ci Cn Cs Cy Fil Fl Fr Gl Gy Ha Inf

Anther, anthers Areole, areoles Axil, axils Body, i.e., the plant body of a cactus, etc. Branch, branches Bract, bracts Bristle, bristles Carpel, carpels Calyx Capitulum (of Asteraceae) Interstaminal corona (of Apocynaceae) Corona Staminal corona (of Apocynaceae) Cyathium (of Euphorbiaceae) Filament, filaments Flower, flowers Fruit, fruits Gland, glands Gynostegium (of Apocynaceae) Hair, hairs Inflorescence, inflorescences

xxxvi

Int ITep L Nec NGl NSc OTep Ov Pc Ped Per Pet Phy Poll R Rec Ri Ros Sc Se Sep Sp SpS St Sti Sty Tep Tu

List of Abbreviations and Symbols

Internode, internodes Inner tepal, inner tepals Leaf, leaves Nectary, nectaries Nectary gland, nectary glands Nectary scale, nectary scales Outer tepal, outer tepals Ovary, ovaries Pericarpel Pedicel, pedicels Perianth Petal, petals Phyllaries (of Asteraceae capitula) Pollinia Root, roots Receptacle Rib, ribs Rosette, rosettes Scale, scales Seed, seeds Sepal, sepals Spine, spines Spine shield (in Euphorbiaceae) Stamen, stamens Stigma Style, styles Tepal, tepals Tubercle, tubercles (of cactus bodies, etc.)

About the Editors

Dr. Urs Eggli studied botany at the University of Zürich and produced a monographic study of the genus Rosularia (Crassulaceae) for his Ph.D. Subsequently, he became a scientific assistant at the Sukkulenten-Sammlung Zürich, Switzerland, where he continues to curate the herbarium and is responsible for public education, special exhibits, and all aspects of information dissemination. His main focus of interest is the diverse biological aspects of succulent plants. Dr. Eggli is an expert in the classification of succulents, especially of the families Crassulaceae and Cactaceae, and a specialist of nomenclature. He traveled widely to study succulent plants in habitat, especially in Latin America. His numerous publications cover a diversity of aspects of the biology of succulent plants. Since 1995 he is editor of the annual Repertorium Plantarum Succulentarum. During 2001–2003 he edited four volumes of the first edition of the present series, and in 2005 he translated and revised E. F. Anderson’s 2001 monograph of the Cactaceae. Dr. Eggli was awarded the Cactus d’Or by the International Organization for Succulent Plant Study in 2015. Dr. Reto Nyffeler studied biology at the University of Zürich. In 1997 he received a doctor’s degree with a Ph.D. thesis on the systematics of the tribe Notocacteae (Cactaceae) from South America. Subsequently, he spent 4 years as a postdoctoral student at Harvard University and Stanford University and conducted during that stay the first comprehensive molecular phylogenetic study of Cactaceae. Since 2002 he is curator of the vascular plant collections at the United

xxxvii

xxxviii

About the Editors

Herbaria of the University and ETH Zurich, and he is a lecturer for plant systematics at the Department of Systematic and Evolutionary Botany, University of Zürich. His research focuses on the systematics of succulent plants and on the history of botany. During the past two decades he published several papers on succulents from different plant families in collaboration with Dr. Urs Eggli.

Contributors

F. Albers Formerly at Wilhelms-Universität, Münster, Germany F. Almeda California Academy of Sciences, Institute for Biodiversity Sciences and Sustainability, San Francisco, CA, USA C. C. Berg Formerly of The Norwegian Arboretum, Hjeltestad, Norway S. Carter Godalming, UK R. P. Clark Royal Botanic Gardens, Kew, Richmond, UK B. Descoings Rosieres, France W. J. de Wilde Leiden, The Netherlands U. Eggli Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland E. Figueiredo Ria Olivier Herbarium, Department of Botany, Nelson Mandela University, Gqeberha, South Africa P. I. Forster Department of Environment and Science, Brisbane Botanic Gardens, Queensland Herbarium, Toowong, QLD, Australia G. Germishuizen Formerly National Botanical Institute, The National Herbarium, Pretoria, Republic of South Africa G. P. Guymer Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Queensland Herbarium, Toowong, QLD, Australia M. E. E. Hijman Formerly of Utrecht University, Utrecht, The Netherlands G. Kadereit Botanischer Garten München-Nymphenburg and Botanische Staatssammlung München, Ludwig-Maximilians-Universität München, Munich, Germany U. Meve Lehrstuhl für Pflanzensystematik, Universität Bayreuth, Bayreuth, Germany

F. Albers: deceased. C. C. Berg: deceased. B. Descoings: deceased. xxxix

xl

L. E. Newton Herbarium, Royal Botanic Gardens, Kew, UK G. N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya Reto Nyffeler Institut für Systematische und Evolutionäre Botanik (ISEB), Universität Zürich, Zürich, Switzerland M. J. R. Rocha Departamento de Ciências Biológicas, Universidade do Estado de Minas Gerais – Unidade Ibirité, Ibirité, Brazil J. M. H. Shaw Horticultural Taxonomy, Science and Collections, Royal Horticultural Society, Woking, UK D. N. Silva Programa de Pós-graduação em Botânica, Escola Nacional de Botânica Tropical, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro, Brazil G. F. Smith Ria Olivier Herbarium, Department of Botany, Nelson Mandela University, Gqeberha, South Africa E. J. Van Jaarsveld Babylonstoren Farm, Simondium, South Africa C. C. Walker School of Environment, Earth and Ecosystem Sciences, Open University, Milton Keynes, England

Contributors

Introduction to the Diversity of Succulent Rosids U. Eggli and Reto Nyffeler

General Remarks Rosids (rosid clade, also more formally called Rosidae or Rosanae) comprise some 70,000 species, roughly one-fourth of all angiosperms, that are classified into about 140 families and 17 orders (APG 2016, but also Magallón & al. 1999; Christenhusz & al. 2017; Li & al. 2021). Rosids exhibit a great many different habits and growth forms, habitat preferences, and life strategies; the rosid clade includes ephemeral to perennial and often deciduous herbs, shrubs, trees, vines, aquatics, epiphytes, parasites, as well as succulents, from a wide variety of biomes. Remarkably, this clade includes most lineages of extant temperate and tropical forest tree species (Wang & al. 2009). This said, succulence is comparatively poorly developed in the rosid clade: overall, close to 1,500 species, making up slightly more than 2% percent of the total species diversity of Rosids, are recognized as succulents (vs.  4% succulents in the Angiosperms overall [Eggli & Nyffeler 2009: 33],  10% succulents in the Monocots [excluding orchids, Nyffeler & Eggli

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland R. Nyffeler Institut für Systematische und Evolutionäre Botanik (ISEB), Universität Zürich, Zürich, Switzerland

2020: 1],  36% succulents in the order Caryophyllales [4200 succulents out of 11,200 in the entire order; Nyffeler & Eggli 2010: 132 and Stevens 2001+, accessed November 2021]). Succulence is known to occur in 12 of the 17 orders of Rosidae and in 26 out of 140 or so families, in most cases, however, just at low percentages (Fig. 1). No clear structural synapomorphies are present to characterize the rosid clade, but there are distinct characteristic trends, such as the presence of stipules, calyx and corolla with free parts, and two whorls of stamens. Furthermore, Rosids have bitegmic, crassinucellate ovules, distinguishing them from Asterids within Eudicots that have unitegmic and tenuinucellate ovules. Rosids include many important crops, including cucumbers and pumpkins (Cucurbitaceae), legumes (Fabaceae), a great diversity of fruit crops (Rosaceae), and timber trees (Sapindaceae). Furthermore, the rosid clade contains lineages with prominent biochemical pathways for defense, such as the production of glucosinolate (Brassicales) and cyanogenic glycosides (Rosaceae and others), or non-protein amino acids (Fabaceae, some Sapindaceae, and others), and with mechanisms for the symbiosis with nitrogen-fixing bacteria (some Betulaceae, Fabaceae) (Wang & al. 2009; Mabberley 2017). Rosids consist of two major subclades, Fabids (also known as Eurosids I) and Malvids (also known as Eurosids II), each comprising eight orders, and include the order Vitales (single family

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_121

1

2

U. Eggli and R. Nyffeler

Saxifragales Vitales

(2476 / 1440) (850 / 68)

Zygophyllales (345 / 6) Celastrales (1355 / −) Fabids

Malpighiales

(16065 / 1014)

Oxalidales

(1845 / 41)

Fabales

(20410 / 26)

Rosales

(8010 / 32)

Cucurbitales

(2990 / 147)

Fagales (1175 / −) Geraniales (897 / 195)

Malvids

Myrtales

(13005 / 34)

Crossosomatales (66 / −) Picramnales (50 / −) Sapindales

(6570 / 37)

Huerteales (29 / −) Malvales

(6005 / 41)

Brassicales

(5035 / 34)

Fig. 1 Summary phylogeny of the Rosid clade with orders as terminals and proportion of total species (gray) and succulent species (black) diversity indicated by triangle widths (based on APG (2009), APG (2016), and Li & al. (2021)). Estimated total numbers of taxa per order

(left-hand figure) are based on Stevens (2001+, accessed November 2021); total numbers of succulent species (right-hand figure) are based on the individual chapters of the present handbook. (Copyright: U. Eggli)

Vitaceae) as sister-group and in a cladistically ancestral position to all other members of the clade (Fig. 1). The internal relationships of both

major subclades Fabids and Malvids are not yet fully resolved. We here follow the topology offered by APG IV (2016) and Christenhusz &

Introduction to the Diversity of Succulent Rosids

al. (2017), recently further confirmed by studies of large datasets of plastid DNA (Li & al. 2021).

Diversity of Succulents in Rosids At the rank of families, only Bixaceae, Caricaceae, Cucurbitaceae, Euphorbiaceae, Geraniaceae, and Moringaceae show proportions of succulent species higher than 10%, but only in the Cucurbitaceae, Euphorbiaceae, and Geraniaceae, more than 100 species are recognized as succulent (Table 1; i.e., 103 succulents for Cucurbitaceae, 944 for Euphorbiaceae, 195 for Geraniaceae). These three families are well-recognized for the morphological diversity among their succulent representatives. However, for most of the families with succulent representatives in Rosids, succulents make up only a very low percentage of the total species diversity. From a statistical viewpoint, exceptions are some small families, where succulents make up a significant fraction or even the majority of the total diversity, such as Bixaceae (4 out of 22 species succulent), Caricaceae (9 out of 34), and Moringaceae (11 out of 13). In the most species-rich families of the rosid clade, such as Fabaceae (26 succulents out of 19,500 species), Melastomataceae (35 out of 5750), Malvaceae (37 out of 4200), and Brassicaceae (13 out of 3700), succulence is an exceedingly rare phenomenon.

Types of Succulence and Life Strategies in Rosids The structural diversity of succulence can be classified into four distinct types: (1) leaf succulents, (2) stem succulents with cactus habit and predominantly stem-based photosynthesis, (3) stem succulents with pachycaul or caudiciform stems and no or only minimal stem-based photosynthesis, and (4) geophytic succulents with water storage in tubers, bulbs, expanded roots, or underground stems (Table 2). The first three types have the common characteristic of above-ground succulence and differ in that respect from the fourth succulence type.

3

In comparison with the remaining Eudicots, the exceedingly diverse types of stem succulence among Rosids are eye-catching: while cactoid stem succulence (i.e., with stem-based photosynthesis due to a long-lived epidermis and chlorophyllous cortex) is common in species of Euphorbia, it is almost completely absent from all other families of the rosid clade, with the exception of some rare cases in Cucurbitaceae (Seyrigia), Geraniaceae (Pelargonium tetragonum), and Vitaceae (some species of Cissus). The predominant form of stem succulence in Rosids is pachycauly (Hearn & al. 2013: 1051; woody plants with greatly enlarged trunklike stems, “sarcocaulescents” of Bobich & North 2009: 9), i.e., overbuilt disproportionally massive and thick stems relative to the branched crown. This growth form is diagnostic for the succulents of Malvaceae and Moringaceae, but also occurs scatteredly in other families (e.g., Burseraceae, Vitaceae, isolated cases such as Euphorbia pachysantha in the Euphorbiaceae). These species store water in their parenchymatic wood, which is lightweight in structure, with comparatively few fibrous components and limited lignification, and turgor pressure being an important component to provide stability. The monotypic genus Carica (Caricaceae) is notable for the overall stem construction, where lignified phloem fibers in a honeycomb-like mesh pattern act as flexible exoskeleton that encloses strongly parenchymatic wood with almost no reinforcing structures – a unique stem architecture among succulents as far as known. Overall, pachycaulous stem succulence with non-photosynthesizing stems occurs in 19 out of 26 families (73%), which is much higher than in the remaining Eudicots (19 out of 40 families; 48%) or Monocots (4 out of 15 families; 26%). A special characteristic of pachycaulous stem succulence are the abbreviated and sometimes almost tuber-like stems of the so-called caudiciform or caudex plants (Eggli & Nyffeler 2009): there is no clear-cut difference between pachycauls and caudiciforms, but rather a continuum, and the tuber-like succulent organs can be anything from true stems (e.g., Cyphostemma of

Brassicales

Malvales

341 6 1 1

Brassicaceae

Caricaceae

Moringaceae

Tropaeolaceae

243

Malvaceae

Sapindaceae 4

142

Meliaceae

Bixaceae

19 51

Burseraceae

81

Anacardiaceae

177

Melastomataceae

9

Sapindales

4

Geraniaceae

Francoaceae

Cucurbitaceae

Myrtales

Geraniales

2 97

Begoniaceae

1

1

4

3

7

2

1

1

4

2

7

1

2

30

1

3

2

8

1

1

1

2

1

4

2

Gen succ

100

13

34

3700

4200

22

1900

641

860

873

5750

31

800

1000

1871

2125

1140

19'500

770

2370

1050

6500

800

280

950

Spe tot

1

11

9

13

37

4

1

1

28

7

34

1

195

103

44

8

24

26

41

1

60

947

6

6

68

Spe succ

O

OOO

OOO

o

o

OO

o

o

O

o

o

O

o

OO

o

o

O

o

O

o

O

OO

o

O

O

Spe div

XX

x

x

XXX

x

X

X

X

X

x

X

x

XXX

XX

X

Leaf succ

x

x

x

x

XX

X

Cact succ

XX

XX

XX

X

XXX

XXX

XXX

XXX

x

XX

X

X

X

X

X

X

XXX

XX

XX

XX

Pachy succ

XXX

XX

X

x

XX

X

XX

XX

X

XX

XX

XX

XX

X

X

Geophy succ

Gen tot = total genera per family; Gen succ = genera with succulents per family; Spe tot = total species per family; Spe succ. = succulent species per family Spe div = Species diversity of families with succulents, as percentage of succulent species: o = < 1 %, O = 1–10%, OO = 10–25%, OOO = > 25% Types of succulence (see Table 2 for definitions) indicated in the last four columns; relative frequency of succulence type: X X X = all; XX = common; X = occasional; X = rare The different shades of gray indicate relative proportions: light light gray = o/x, light gray = O/X , gray = OO/XX , dark gray = OOO/ XXX

Rosids-Malvids

51

Urcaceae

Cucurbitales

39

Moraceae

Rosales

798

Fabaceae

Fabales

6

59

Phyllanthaceae

Oxalidaceae

19

Passifloraceae

Oxalidales

14 218

Euphorbiaceae

Malpighiales

30

17

Gen tot

Clusiaceae

Zygophyllaceae

Zygophyllales

Rosids-Fabids

Vitaceae

Vitales

Rosids

Table 1 Classification of succulent Rosids: orders, families, species diversity, type of succulence, and life strategies (figures compiled from the individual chapters of this handbook, and Mabberley (2017) and Stevens (2001+, accessed Nov. 2021))

4 U. Eggli and R. Nyffeler

Introduction to the Diversity of Succulent Rosids

5

Table 2 Structural and ecological characteristics of four types of succulence in Rosids, based on the location of the water storing plant part, place of photosynthesis, and size and duration of the leaves Succulence Water storage Photosynthesis Leaf size

Above-ground Leaves Leaf-based Large to small

Leaf longevity Type

Persistent Leaf succulents

Stems Stem-based Small to tiny, or fully reduced Absent or caducous Cactoid stem succulents

the Vitaceae) with highly parenchymatous wood to hypocotyl tubers (e.g., Gerrardanthus of the Cucurbitaceae) that sometimes merge invisibly into the thickened taproot. A special feature of many of these caudiciform stem succulents is the phenomenon of periodic shoot reduction, where much of the non-succulent aerial growth shrivels and dries out at the onset of the dry season, and is eventually shed completely, to be replaced with a completely new set of shoots and leaves in the next rain period. Among the succulents of the Rosid clade, this growth form is particularly prevalent among the climbing Cucurbitaceae and Vitaceae, but is also known from Adenia (Passifloraceae). Periodical shoot reduction is otherwise best known from geophytes and hemicryptophytes, but among succulents this phenomenon also occurs in chamaephytes and nanophanerophytes with above-ground water storage stems (“reductive chamaephytes,” Jürgens 1990: 324–325). In some genera and families of the Rosid clade, two or more architectures and types of succulence have evolved side by side among their species. A well-studied example is the genus Adenia (Passifloraceae), where a complete continuum from pachycauls to caudiciforms and geophytes exists, often associated with anomalous secondary growth, and including hypocotyl tubers and true moniliform succulent roots. Hearn (2006, 2009a, b) stresses the plasticity involved. Similar diversity is present in Moringa (Moringaceae), which was studied by Olson & Carlquist (2001) and Olson (2007). Hearn & al. (2013) found that there is a general “apparent lability of

Stems Leaf-based Large to small Caducous Pachycaul stem succulents

Below-ground (Geophytes) Underground parts Leaf-based Large to small Caducous or persistent Geophytic succulents (tubers, bulbs, roots, underground stems)

evolutionary change among succulent forms, esp. in this continuum from stem to root succulence.” Furthermore, some special developments can be observed among the succulents of Rosids: Species of Clusia (Clusiaceae) are unique in that they combine an “ordinary” woody shrub or tree architecture with distinctly succulent large leaves. Some species of Cissus (Vitaceae) have evolved a similar combination of non-succulent stems and succulent leaves, but these species are trailers or climbers with primarily herbaceous and fibrous stems. Finally, the monotypic genus Tetilla (Francoaceae) is unique in its combination of water storage in an underground tuber and in the petioles of its deciduous leaves. “Petiole succulents” are otherwise only known from the monotypic monocot genus Zamioculcas (Araceae).

Evolutionary Diversification of Succulence in Rosids From an evolutionary viewpoint, the pattern of diversification of succulent plants in Rosids makes a scenario of recurrent parallel innovation of the “succulent life strategy” very plausible. A very conservative estimate is that succulence evolved independently at least 26 times, namely once for each of the 26 families of Rosids that have at least one succulent representative. But the evolutionary pattern of succulence diversification is much more complex, since for many families, succulence has evolved independently in members of different clades – this is true for

6

Brassicaceae (3 out of 4 clades), Euphorbiaceae (2 out of 4 subfamilies), Fabaceae (3 out of 6 subfamilies), Malvaceae (3 out of 9 subfamilies), Melastomataceae (at least 4 out of 15 clades in 1 out of 2 subfamilies), Moraceae (2 out of 6 tribes), Urticaceae (2 out of 4 tribes), and Vitaceae (2 out of 5 tribes of 1 out of 2 subfamilies) (all data based on the individual chapters of this handbook). In addition, succulence is not a genus-wide characteristic in most larger genera, but again, repeated independent origins must be assumed, as exemplified by Begonia (at least 1 succulent species in 8 out of 9 major clades) or Euphorbia, where succulent species occur in variable proportions in all four subgenera: succulents are present in 3 out of 21 sections of subgen. Esula, in 6 out of 7 sections of subgen. Athymalus, in 8 out of 15 sections of subgen. Chamaesyce, and in 16 out of 21 sections of subgen. Euphorbia – summing up to probably as many as 33 independent occurrences of succulence just in this single genus, and for the Rosid clade as a whole, succulence likely has evolved independently at least 60 times. Future research with additional molecular phylogenetic studies will enable more succulent plant taxa to be included in the sampling strategy, and synthetic analyses of all studies of Rosid taxa published so far will together provide more information to tackle this fundamental research question.

References APG (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Bot. J. Linn. Soc. 161(2): 105–121. https://doi.org/10.1111/j.1095-8339.2009. 00996.x. APG (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Bot. J. Linn. Soc. 181(1): 1–20. https://doi.org/10.1111/boj.12385. Bobich, E. G. & North, G. B. (2009) Structural implications of succulence: Architecture, anatomy, and mechanics of photosynthetic stem succulents, pachycauls, and leaf succulents. In: Barrera, E. de la & Smith, W. K. (eds.): Perspectives in biophysical plant ecophysiology. A tribute to Park S. Nobel; pp. 3–37. México (MX): Universidad Nacional Autónoma de

U. Eggli and R. Nyffeler México. http://www.researchgate.net/publication/ 261065161 Christenhusz, M. J. M. [& al. 2017], Fay, M. R. & Chase, M. W. (2017) Plants of the World. An illustrated encyclopedia of vascular plants. Richmond (GB): Kew Publishing, Royal Botanic Gardens, Kew/Chicago (US): University of Chicago Press. vii + 792 pp., ills., maps. Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions – succulence as an adaptive strategy. Bradleya 27: 13–36, ills. https://doi.org/10.25223/brad. n27.2009.a10. Hearn, D. J. (2006) Adenia (Passifloraceae) and its adaptive radiation: Phylogeny and growth form diversification. Syst. Bot. 31(4): 805–821, ills. https://doi.org/10. 1600/036364406779695933. Hearn, D. J. (2009a) Developmental patterns in anatomy are shared among separate evolutionary origins of stem succulent and storage root-bearing growth habits in Adenia (Passifloraceae). Amer. J. Bot. 96(11): 1941–1956, ills. https://doi.org/10.3732/ajb.0800203. Hearn, D. J. (2009b) Descriptive anatomy and evolutionary patterns of anatomical diversification in Adenia (Passifloraceae). Aliso 27(1): 13–38. https://doi.org/ 10.5642/aliso.20092701.03. Hearn, D. J. [& al. 2013], Poulsen, T. & Spicer, R. (2013) The evolution of growth forms with expanded root and shoot parenchymatous storage is correlated across the Eudicots. Int. J. Pl. Sci. 174(7): 1049–1061, ills. https:// doi.org/10.1086/671745. Jürgens, N. (1990) A life form concept including anatomical characters, adapted for the description of succulent plants. Mitt. Inst. Allg. Bot. Hamburg 23a: 321–341. Li, H. T. [& al. 2021], Guo, Y., Gan, L., Ma, P.-F., Gao, L.M., Yang, J.-B., Cai, J., Gitzendanner, M. A., Fritsch, P. W., Zhang, T., Jin, J.-J., Zeng, C.-X., Wang, H., Yu, W.-B., Zhang, R., Bank, M. van der, Olmstead, R. G., Hollingsworth, P. M., Chase, M. W., Soltis, D. E., Soltis, P. S., Yi, T.-S. & Li, D.-Z. (2021) Plastid phylogenomic insights into relationships of all flowering plant families. BMC Biol. 19: 232, 13 pp. https://doi.org/10.1186/s12915-021-01166-2. Mabberley, D. J. (2017) Mabberley’s Plant-Book. A portable dictionary of plants, their classification and uses. Ed. 4. Cambridge (GB): Cambridge University Press. xix + 1102 pp. Magallón, S. [& al. 1999], Crane, P. R. & Herendeen, P. S. (1999) Phylogenetic pattern, diversity, and diversification of eudicots. Ann. Missouri Bot. Gard. 86(2): 297–372. http://www.jstor.org/stable/2666180 Nyffeler, R. & Eggli, U. (2010) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/10.25223/brad.n28.2010.a1. Nyffeler, R. & Eggli, U. (2020) Introduction to the classification of Monocotyledons. In: Eggli, U. & Nyffeler, R. (eds.): Illustrated Handbook of Succulent Plants: Monocotyledons, ed. 2; pp. 1–6. Berlin (DE): Springer. https://doi.org/10.1007/978-3-662-56486-8_113. Olson, M. E. (2007) Wood ontogeny as a model for studying heterochrony, with an example of paedomorphosis in Moringa (Moringaceae). Syst. Biodivers. 5(2):

Introduction to the Diversity of Succulent Rosids 145–158, ills. https://doi.org/10.1017/S14772000060 02180. Olson, M. E. & Carlquist, S. (2001) Stem and root anatomical correlations with life form diversity, ecology, and systematics in Moringa (Moringaceae). Bot. J. Linn. Soc. 135(4): 315–348, ills. https://doi.org/10. 1111/j.1095-8339.2001.tb00786.x. Stevens, P. F. (2001+) Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously

7 updated since]. http://www.mobot.org/MOBOT/ research/APweb/ Wang, H. [& al. 2009], Moore, M. J., Soltis, P. S., Bell, C. D., Brockington, S. F., Alexandre, R., Davis, C. C., Latvis, M., Manchester, S. R. & Soltis, D. E. (2009) Rosid radiation and the rapid rise of angiospermdominated forests. Proc. Nation. Acad. Sci. USA 106(10): 3853–3858. https://doi.org/10.1073/pnas. 0813376106.

Part I Anacardiaceae

Anacardiaceae U. Eggli

Anacardiaceae R. Brown (in Tuckey, Narr. Exped. Zaire, 431, 1818). Type: Anacardium Linné. Nom. cons. (ICBN 2006: 138). Order: Sapindales. Distribution: Tropics and subtropics worldwide, Mediterranean, N America. Including Blepharocaryaceae Airy Shaw (1965). Including Comocladiaceae Martynov (1820). Including Julianaceae Hemsley (1906). Including Lentiscaceae Horaninow (1843). Including Pistaciaceae Martynov (1820). Including Podoaceae Baillon ex Franchet (1889). Including Rhoaceae Sadler (1826). Including Schinaceae Rafinesque (1837). Including Spondiadaceae Martynov (1820). Including Vernicaceae Schultz-Schultestein (1832). Trees or shrubs, rarely climbers, stems sometimes conspicuously thickened, with resin ducts and often black or blackening aromatic-resinous exudate, often highly allergenic, sometimes with exfoliating bark; L spirally arranged (rarely opposite or whorled), frequently imparipinnate but rarely simple or bipinnate, leaflets opposite or

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

sometimes alternate, entire to serrate; stipules absent; Inf terminal and/or axillary, of various architectures (thyrsoid, paniculate, racemose, or spicate), rarely cauliflorous or flowers solitary; Fl small and rather inconspicuous, (3- to) 5- (to 7-) merous, actinomorphic, bisexual or apparently perfect but unisexual (and plants dioecious, monoecious, andromonoecious, polygamous, or hermaphrodite), sometimes with a hypanthium; Ped often articulated; Sep (3
) 4–5, mostly basally united; Pet (3
) 4–5 (
8), or rarely absent, free, caducous to persistent; St (1
) 5–10 or more (to >100), in 1 or 2 whorls, filaments sometimes basally united and forming a tube; Ov superior or rarely inferior, of 1 or 2–12 united (or very rarely free) carpels, 1- to 5- (to 12-) locular, ovule 1 per locule; Fr drupes or samaras (rarely utricles or baccate), often flat, often operculate with excentric style remains, or variously winged (adnate to broad bracts, with flattened peduncle of the inflorescence, or much enlarged sepals or petals). Important Literature: Pell (2004: molecular phylogeny); Mitchell & al. (2006: phylogeny, esp. Spondioideae); Bachelier & Endress (2009: flower morphology & anatomy); Pell & al. (2011: synoptical overview); Weeks & al. (2014: phylogeny & biogeography); Herrera & al. (2018: fruit morphology & anatomy Spondiadoideae). A medium-sized family with 81 genera and 800 species (Pell & al. 2011) (80 genera and 873 species according to (Stevens 2001+:

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_1

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U. Eggli

accessed Feb. 2021) with a predominantly tropical-subtropical occurrence, but also with many species in temperate N America. Classification: Anacardiaceae are closely related to the similarly sized Burseraceae, and taxa with exfoliating bark (e.g. Pachycormus) are looking remarkably similar to some species of Bursera. The 2 families, together with Kirkiaceae (6 species in 1 genus from Africa and Madagascar, no succulents) form a wellsupported branch in the Sapindales (Kubitzki 2011), confirmed by many recent molecular studies, including Sun & al. (2016: 382). Weeks & al. (2014) date the divergence between Anacardiaceae and Burseraceae to about 100 mybp, associated with a vicariance between E and W hemispheres. The crown age of Anacardiaceae is given as 97 my (83–128 my range) by Weeks & al. (2014). Anacardiaceae differ from Burseraceae in having often allergenic (vs. always non-allergenic) resin, usually imbricate (vs. induplicate-valvate) petals, and 1 apotropous ovule (vs. 2 epitropous ovules) per locule (¼ carpel) (Weeks & al. 2005: 85). The family steadily accumulated diversity starting in the late Cretaceous/Palaeocene, and multiple intercontinental colonization events are postulated, involving frequent shifts between climatic niches (Weeks & al. 2014).

Anacardioideae

Traditionally, Anacardiaceae are divided into 5 tribes or 4–5 subfamilies (Pell 2004: 13). Based on molecular studies, a division into 2 subfamilies has been found to be more adequate, however (Pell 2004, Mitchell & al. 2006, Pell & al. 2011, Sun & al. 2016) (Fig. 1): Spndiadoideae with ovaries mostly 4- to 5-locular, corresponding to the earlier tribe Spondiadoideae, and Anacardioideae with ovaries 1- or 2-locular, corresponding to the remaining traditional tribes Anacardieae, Dobineae, Rhoeae and Semecarpeae. Spondiadoideae are polyphyletic, though, and further research is necessary to gain a complete picture. Weeks & al. (2014) also found a deep split between the 2 subfamilies, and recognize 4 clades in Anacardioideae and 2 clades in Spondiadoideae. Economical Importance: Important crop plants of the family are pistachio (Pistacia vera Linné), mango (species of Mangifera) and cashew nut (Anacardium occidentale Linné); Schinus species are the origin of pink pepper. Numerous species of sumac (Rhus) contain strong allergenic poisons while others furnish tannins and lacquers. Succulence: Little-pronounced succulence is found in the form of pachycaul trunks in a few taxa from dry habitats. Tuberous but woody underground stems (xylopodia) are common in

Spondiad.

200 species

Fig. 1 Summary phylogeny of Anacardiaceae based on Stevens (2001+, accessed Feb. 2021) (873 species total). Spondiad. ¼ Spondiadoideae. (Copyright: U. Eggli)

Anacardiaceae

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species from fire-prone habitats such as African savannas of the Zambezian region, or cerrado vegetation in S America (Pell & al. 2011). Several of the pachycaul species are termed succulent in the literature, e.g. Pachycormus (“one of the most extreme cases of wood succulence among pachycauls”, Bobich & North (2009: 21)), or Spondias purpurea Linné (“water-storage plant”, Schulze & al. (1988); “tropical succulent tree”, Borchert & Rivera (2001)). For others, e.g. species of Cyrtocarpa Kunth, “succulent stems” or “succulent branchlets” are reported (Pell & al. 2011: 7, 40). Phenological and physiological data for Spondias purpura show that the taxon is physiologically active during the dry season: Stem water potential is always high throughout the year (Borchert 2000), and bud break usually occurs well before the first rains of the season (Borchert 1996). For S. mombin Linné, seedling stem dry matter content is only 27–28%, resulting in stem densities of 0.26 g/cm3 (Poorter & Markesteijn 2008: supplemental table). Seedlings of Operculicarya decaryi and O. pachpus produce a well-defined tuberous caudex in the first weeks after germination (Houston & Stead 2007). Fleshy water storage roots in the form of intermittent swellings to 20 cm ∅ have also been reported, e.g. for Spondias tuberosa Arruda (Goebel 1889: 27). Cyrtocarpa edulis Standley is reported as pachycaulous with “succulent/woody swollen trunks”, and is regarded as succulent since even cuttings of branches with 15 cm ∅ can be successfully rooted (Trager & Farrier 2021: 81, with ills.). Apart from the taxa treated below, seedling plants of species of Cyrtocarpa and Lannea A. Richard are also occasionally cultivated by succulent plant enthusiasts. The degree of succulence of species of Spondias, as mentioned above, needs further study, and the genus is at present omitted from this Handbook. Key to genera with succulents 1a 1b

Fl solitary or in small fascicles; Fr fleshy drupes Fl in many-flowered panicles; Fr dry

Operculicarya Pachycormus

References Bachelier, J. B. & Endress, P. K. (2009) Comparative floral morphology and anatomy of Anacardiaceae and Burseraceae (Sapindales), with a special focus on gynoecium structure and evolution. Bot. J. Linn. Soc. 159(4): 499–571, ills. https://doi.org/10.1111/j.10958339.2009.00959.x. Bobich, E. G. & North, G. B. (2009) Structural implications of succulence: Architecture, anatomy, and mechanics of photosynthetic stem succulents, pachycauls, and leaf succulents. In: Barrera, E. de la & Smith, W. K. (eds.): Perspectives in biophysical plant ecophysiology. A tribute to Park S. Nobel; pp. 3–37. México (MX): Universidad Nacional Autónoma de México. http://www.researchgate.net/publication/ 261065161 Borchert, R. (1996) Phenology and flowering periodicity of neotropical dry forest species: Evidence from herbarium collections. J. Trop. Ecol. 12(1): 65–80. https:// doi.org/10.1017/S0266467400009317. Borchert, R. (2000) Organismic and environmental controls of bud growth in tropical trees. In: Viemont, J. D. & Crabbé, J. (eds.): Dormancy in plants: From whole plant behaviour to cellular control; pp. 87–107. Wallingford (GB): CAB International. Borchert, R. & Rivera, G. (2001) Photoperiodic control of seasonal development and dormancy in tropical stemsucculent trees. Tree Physiol. 21(4): 213–221, ills., diag. https://doi.org/10.1093/treephys/21.4.213. Goebel, K. (1889) Pflanzenbiologische Schilderungen. Erster Teil. Marburg (DE): N. G. Elwert’sche Verlagsbuchhandlung. https://doi.org/10.5962/bhl. title.1665. Herrera, F. [& al. 2018], Mitchell, J. D., Pell, S. K., Collinson, M. E., Daly, D. C. & Manchester, S. R. (2018) Fruit morphology and anatomy of the Spondioid Anacardiaceae. Bot. Rev. (Lancaster) 84: 315–393, ills. https://doi.org/10.1007/s12229-018-9201-1. Houston, D. M. & Stead, J. M. (2007) Camelhair confidential. Techniques for sexual propagation of Operculicarya species. Cact. Succ. J. (US) 79(3): 97–108, ills. https://doi.org/10.2985/0007-9367(2007) 79[97:CC]2.0.CO;2. Kubitzki, K. (2011) Introduction to Sapindales. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 1–3. Berlin/Heidelberg (DE) etc.: Springer Verlag. https://doi.org/10.1007/978-3-64214397-7_1. Mitchell, J. D. [& al. 2006], Daly, D. C., Pell, S. K. & Randrianasolo, A. (2006) Poupartiopsis gen. nov. and its context in Anacardiaceae classification. Syst. Bot. 31(2): 337–348, ills. https://doi.org/10.1600/ 036364406777585757. Pell, S. K. (2004) Molecular systematics of the Cashew Family (Anacardiaceae). Baton Rouge (US): Louisiana State University; unpublished Ph.D. thesis. Pell, S. K. [& al. 2011], Mitchell, J. D., Miller, A. J. & Lobova, T. A. (2011) Anacardiaceae. In: Kubitzki,

14 K. (ed.): The families and genera of vascular plants, volume X; pp. 7–50, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-642-14397-7_3. Poorter, L. & Markesteijn, L. (2008) Seedling traits determine drought tolerance of tropical tree species. Biotropica 40(3): 321–331. https://doi.org/10.1111/j. 1744-7429.2007.00380.x. Schulze, E.-D. [& al. 1988], Mooney, H. A., Bullock, S. H. & Mendoza, A. (1988) Water contents of wood of tropical deciduous forest species during the dry season. Bol. Soc. Bot. México 48: 113–118. https://doi.org/10. 17129/botsci.1349. Stevens, P. F. (2001+) Angiosperm Phylogeny Website, Version 14, July 2017 [and more or less continuously updated since]. St. Louis (US): Missouri Botanical Garden. http://www.mobot.org/MOBOT/research/ APweb/ Sun, M. [& al. 2016], Naaem, R., Su, J.-X., Cao, Z.-Y., Burleigh, J. G., Soltis, P., Soltis, D. E. & Chen, Z.-D.

U. Eggli (2016) Phylogeny of the Rosidae: A dense taxon sampling analysis. J. Syst. Evol. 54(4): 363–391. https:// doi.org/10.1111/jse.12211. Trager, J. N. & Farrier, A. (2021) The 2021 offering of the International Succulent Introductions. Cact. Succ. J. (US) 93(2): 72–89, ills. https://doi.org/10.2985/015. 093.0201. Weeks, A. [& al. 2005], Daly, D. C. & Simpson, B. B. (2005) The phylogenetic history and biogeography of the Frankincense and Myrrh family (Burseraceae) based on nuclear and chloroplast sequence data. Molec. Phylogen. Evol. 35(1): 85–101. https://doi. org/10.1016/j.ympev.2004.12.021. Weeks, A. [& al. 2014], Zapata, F., Pell, S. K., Daly, D. C., Mitchell, J. D. & Fine, P. V. A. (2014) To move or to evolve: Contrasting patterns of intercontinental connectivity and climatic niche evolution in "Terebinthaceae" (Anacardiaceae and Burseraceae). Frontiers Genetics 5: 409, 20 pp. https://doi.org/10.3389/fgene.2014.00409.

Operculicarya ANACARDIACEAE U. Eggli

Operculicarya Perrier (Mém. Mus. Nation. Hist. Nat., n.s. 18(7): 248, 1944). Type: Operculicarya decaryi H. Perrier [lectotype, designated by Capuron, Adansonia, ser. 2, 2(2): 270–283, 1962]. Spondiadoideae Lit: Teichman & Hardy (1992: flower & fruit structure); Eggli (1995: synopsis); Randrianasolo & Lowry (2006: systematics, conservation); Eslamieh & Stead (2011: cultivation, hybridization); Herrera & al. (2018: fruit morphology & anatomy); Houston & Stead (2019: intermittent dioecy); Distr: Madagascar, Comoros, Aldabra. Etym: Lat. ‘operculum’ small lid; and Gr. ‘karya’, nut tree; for the operculate nut-like seeds. Gnarled dioecious (sometimes only partially so) shrubs or small trees with regularly or irregularly thickened trunks with smooth or  tubercular-bulbous bark, or stems not conspicuously thickened; young twigs slender, straight or zigzag, with long internodes and lateral short shoots (excl. O. gummifera, O. multijuga, both not succulent), drought-deciduous; L spirally arranged, dispersed or in rosettes from the short shoots, glabrous or hairy, imparipinnate with 3–20 pairs of leaflets, rachis  to strongly winged (except O. gummifera and O. multijuga); Fl usually unisexual but sometimes bisexual mixed with

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

unisxeual flowers in the same inflorescence; male Fl solitary or in small fascicles on short shoots, 5-merous; St 10; Fil basally broadened and thickened; Nec irregularly lobed to ring-like, pistillodium small to large; female Fl solitary on short shoots (in spicate many-flowered inflorescences in O. gummifera), similar to male flowers but with sterile anthers; Ov globose to oblong; Sty 5, peripheral; Fr fleshy drupes, sessile or shortly stalked with persistent calyx, red to puplish-black when ripe, stone unilocular (rarely bilocular), with 1 (rarely 2) oblong-elliptic operculum. The genus (8 species) has a wide but scattered distribution in Madagascar (excepting the moist forests of the E coast). Only 2 species are unequivocally succulent, but with the exception of O. calcicola Randrianasolo & Lowry and O. multijuga Randrianasolo & Lowry, all are  xerophytic and show thickened trunks and are treated below. Operculicarya is separated from the closely related Poupartia by endocarps with 1 operculum (vs. 2–5 opercula) and horseshoe-shaped (vs. slightly curved) embryos (Pell & al. 2011). Capuron (1975) included the non-succulent Poupartia gummifera Sprague from Madagascar, the Comoros and Aldabra in Operculicarya and was followed by Randrianasolo & Lowry (2006), while Eggli (1995) and Herrera & al. (2018) excluded it from Operculicarya. The recently described O. calcicola is very similar to O. gummifera, and O. multijuga bridges the gap,

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as it approaches P. gummifera in vegetative characters, but has solitary fruits. Species of Operculicarya are with the exception of O. decaryi and O. pachypus not frequently seen in cultivation, but present no special problems. Eslamieh & Stead (2011) have found most of them to be rapidly growing. Artificial hybrids are easily produced and some of them are vigorous growers. Houston & Stead (2019) found that cultivated specimens of O. gummifera, O. pachypus and esp. O. hyphaenoides occasionally to regularly produce perfect bisexual flowers (sometimes exhibiting spontaneous selfpollination before anthesis), often mixed with unisexual flowers within the same inflorescence. Benadom (2021) also observed a specimen of O. decaryi to produce perfect flowers with spontaneous fruit set. O. borealis Eggli (Bull. Mus. Nation. Hist. Nat., Sect. B, Adansonia 17(3–4): 152–153, ill. (p. 155), 1995). Type: Madagascar, Antsiranana (Capuron 23097SF [P]). Distr: N Madagascar (Antsiranana: Orangea and Sahafary forests); on sandy soils and sandstone, to 550 m. I: Eslamieh & Stead (2011: 245). Small trees; trunk thin in juveniles, becoming thickened with age, bark of adult plants unknown but juveniles with smooth bark; twigs straight, glabrous or slightly hairy; L 2.8–5 cm with (4–) 5–7 pairs of leaflets, rachis a little winged, with scattered bristly hairs, leaflets overlapping one another, roundish-ovate, 4–8  3–4 mm or larger, tip rounded, upper face glabrous and glossy or slightly hairy, lower face densely hairy, with 8–12 pairs of veins; Fl solitary, colour not recorded; Fr solitary, sessile,  globose, turning dark purple or black at maturity. O. capuronii Randrianasolo & Lowry (Adansonia, sér. 3, 28(2): 364, Fig. 2 (p. 365), 2006). Type: Madagascar, Toliara (Service Forestier (Capuron) 28571 [P, MO, P, TEF]). Distr: SE Madagascar (Toliara: Mt. Vohitsiandriana); granite slopes, to 300 m. I: Eslamieh & Stead (2011: 245). Small trees with  swollen trunk, 2–8 m; Br waxy, greyish to brown, with 1–3 cm long short

U. Eggli

shoots, basally glabrous, apically puberulous; L 2.5–9 cm with 3–4 pairs of (sub-) opposite leaflets, rachis winged, leaflets sessile, somewhat coriaceous, obovate, 10–30  4–18 mm, with impressed venation, glabrous but young sparsely puberulous; Fl unknown; Fr 2–3 together or solitary, 7–10  5–7 mm, 1–2 mm pedicellate, glabrous. O. decaryi H. Perrier (Mém. Mus. Nation. Hist. Nat., n.s. 18(7): 249, 1944). Type: Madagascar, Toliara (Decary 9305 [P]). Lit: Teichman & Hardy (1992: flower & fruit structure); Houston & Stead (2007: propagation, hybridization, with ills.). Distr: S and SW Madagascar (Fianarantsoa, Toliara); xerophyte-rich forests, 5–600 m. I: Rauh (1998: 27: Figs. 78, 81–82; 28); Lenain (2012: 178–179); Benadom (2021). Fig. 1. Shrubby or trees to 6 ( 15 ?) m, trunk parallelsided, bottle-shaped or conical, to 1 m ∅, bark irregularly warty-bumpy, dark grey; twigs glabrous or hairy and glabrescent,  straight; L 2.5–6 cm, with (4–) 5–7 ( 9) pairs of leaflets, glabrous or with scattered hairs, rachis  to conspicuously winged, leaflets circular to ovate, 4–7  3–4 mm, not touching each other, tip rounded, upper face dark green and glossy, lower face paler, with 3–4 pairs of veins; Fl dark red; Fr solitary, 5–6 mm pedicellate, globose to drop-shaped.

Fig. 1 Operculicarya decaryi (cult. SukkulentenSammlung Zürich; without known wild origin). (Copyright: U. Eggli)

Operculicarya ANACARDIACEAE

The taxon easily hybridizes with other cultivated species of the genus. Seedlings rapidly grow a juicy thickened succulent root (Houston & Stead 2007). Benadom (2021) reports that a cultivated plant produced clustered bisexual flowers by the thousands that set seed spontaneously, though with a low germination rate of only some 3%. O. hirsutissima Eggli (Bull. Mus. Nation. Hist. Nat., Sect. B, Adansonia 17(3–4): 154–155, ill., 1995). Type: Madagascar, Toliara (Léandri 3447 [P]). Distr: S-C Madagascar (Toliara, Fianarantsoa: area around Ihosy and Sakaraha); deciduous and xerophytic forests, on gneiss,  500–800 m. I: Eslamieh & Stead (2011: 247). Trees to 6 m; trunk to 40 cm ∅, bark of adults unknown; twigs glabrous or slightly tomentose, straight; L 5.5–7 cm, with (9–) 11–13 ( 14) pairs of leaflets, rachis hardly to broadly winged, leaflets often asymmetrically arranged, hardly or not overlapping one another, broadly ovate to elliptic, 6–9  3–4.5 mm, tip rounded to truncate, both faces conspicuously greyish-tomentose, lower face with 5–10 indistinct pairs of veins; Fl cream-coloured; Fr solitary, pendent, 8–12 mm pedicellate, broadly ovoid, without persistent calyx. O. hyphaenoides H. Perrier (Mém. Mus. Nation. Hist. Nat., n.s. 18(7): 249, 1944). Type: Madagascar, Toliara (Perrier 19169 [P]). Distr: SW Madagascar (Toliara: Lac Tsimanampetsotsa); xerophytic scrub on limestone; 0–120 m. I: Rauh (1998: 28); Eslamieh & Stead (2011: 247, 249); Houston & Stead (2019). Small trees to 1.5 m with several gnarled trunks to 40 cm ∅ from a common base, each with a densely branched crown, bark warty-bumpy; twigs glabrous, brownish-grey,  straight; L 2.5–6 cm, medium to dark greenish-olive, with (11–) 14–18 ( 20) pairs of leaflets, rachis hardly to conspicuously winged, leaflets 4  2 mm, roundish-rectangular with truncate tip, margins strongly revolute, lower face with 5–7 pairs of raised veins; Fl pale to bright red, sometimes petals with paler margins; Fr solitary,

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pendent, to 18 mm pedicellate, depressedglobose, pale red to pinkish-red. Plants can look superficially similar to Commmiphora monstruosa (H. Perrier) Capuron (Burseraceae). Houston & Stead (2019) report on plants that regularly produce bisexual flowers, sometimes mixed with unisexual flowers within the same inflorescence. Some of the bisexual flowers showed autonomous self-pollination before anthesis, and developed mature fruits. Seedlings produce thickened tuberous main roots within 2 years from germination. O. pachypus Eggli (Bull. Mus. Nation. Hist. Nat., Sect. B, Adansonia 17(3–4): 155–157, ill., 1995). Type: Madagascar, Toliara (Humbert 2614 [P]). Distr: SW Madagascar (Toliara: vicinity of Toliara); xerophytic forest on limestone, 10–260 m. I: Eggli (2008: 165); Rauh (1998: 26–27 (figs. 79–80), as O. decaryi); Houston & Stead (2019: 166, flowers). Figs. 2 and 3. Bonsai-like to 1 m, trunk conical to irregularly pyramidal, narrowing towards the tip, to 50 cm ∅ with spreading crown, bark irregularly bumpy, silver-grey; twigs glabrous, pale grey to greybrown, often  strongly zigzag, tips often spinescent; L 1.5–3.6 cm with 3–4 ( 5) pairs of leaflets, glabrous, somewhat coriaceous, rachis rather broadly winged, leaflets circular to elongate drop-shaped, completely separate from each other or touching each other, largest near the middle of the leaves, 7–8  4 mm, tip rounded or slightly

Fig. 2 Operculicarya pachypus (cult. SukkulentenSammlung Zürich; without known wild origin). (Copyright: U. Eggli)

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Fig. 3 Operculicarya pachypus (cult. SukkulentenSammlung Zürich; without known wild origin). (Copyright: U. Eggli)

emarginate, upper face dark green, glossy, lower face paler with 3–4 pairs of veins; Fl yellowishgreen; Fr solitary, 3–4 mm pedicellate,  globose, somewhat laterally compressed. Despite the conspicuously different trunk, this species was for long incorrectly identified as O. decaryi, e.g. by Teichman & Hardy (1992) or Rauh (1998). Houston & Stead (2019) observed that perfect bisexual flowers sometimes form.

References Benadom, D. (2021) Operculicarya monoecy. Cact. Succ. J. (US) 93(1): 41–43, ills. https://doi.org/10.2985/015. 093.0105. Capuron, R. (1975) Contribution a l’étude de la flore forestière de Madagascar. Sur l’identité du Poupartia gummifera Sprague (Anacardiaceae). Adansonia, n.s. [sér. 2], 14(4): 571–572. http://www.biodiversity library.org/page/59831439

U. Eggli Eggli, U. (1995) A synoptical revision of Operculicarya (Anacardiaceae). Bull. Mus. Nation. Hist. Nat., Sect. B, Adansonia 17(3–4): 149–158, ills., key. http://biodiversitylibrary.org/page/13740037 Eggli, U. (2008) Sukkulenten. Ed. 2. Stuttgart (DE): Eugen Ulmer. Eslamieh, J. & Stead, J. (2011) The heterosis of Operculicarya. Cact. Succ. J. (US) 82(6): 244–251, ills. https://doi.org/10.2985/0007-9367-82.6.244. Herrera, F. [& al. 2018], Mitchell, J. D., Pell, S. K., Collinson, M. E., Daly, D. C. & Manchester, S. R. (2018) Fruit morphology and anatomy of the Spondioid Anacardiaceae. Bot. Rev. (Lancaster) 84: 315–393, ills. https://doi.org/10.1007/s12229-018-9201-1. Houston, D. M. & Stead, J. M. (2007) Camelhair confidential. Techniques for sexual propagation of Operculicarya species. Cact. Succ. J. (US) 79(3): 97–108, ills. https://doi.org/10.2985/0007-9367(2007) 79[97:CC]2.0.CO;2. Houston, D. M. & Stead, J. (2019) A discussion of transitional dioecy in Operculicarya. Cact. Succ. J. (US) 91(3): 164–176, ills. https://doi.org/10.2985/ 015.091.0301. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer. Pell, S. K. [& al. 2011], Mitchell, J. D., Miller, A. J. & Lobova, T. A. (2011) Anacardiaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 7–50, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-642-14397-7_3. Randrianasolo, A. & Lowry, P. P. II (2006) Operculicarya (Anacardiaceae) revisited: An updated taxonomic treatment for Madagascar and the Comoro Islands, with descriptions of two new species. Adansonia, sér. 3, 28(2): 359–371, ills., maps, key. http://www. researchgate.net/publication/228646369 Rauh, W. (1998) Succulent and xerophytic plants of Madagascar. Vol. 2. Mill Valley (US): Strawberry Press. Teichmann, I. von & Hardy, D. S. (1992) Flower and fruit structure of Operculicarya decaryi H. Perrier (Anacardiaceae) from Madagascar. Bot. Bull. Acad. Sin., n.s. 33(3): 225–232, ills.

Pachycormus ANACARDIACEAE U. Eggli

Pachycormus Coville ex Standley (Contr. US Nation. Herb. 23(3): 671, 1923). Type: Schinus discolor Bentham. – Anacardioideae – Lit: Gibson (1981: anatomy); Humphrey (1991: synopsis). Distr: Mexico (Baja California peninsula). Etym: Gr. ‘pachys’, thick; and Gr. ‘kormos’, trunk; for the thick trunk and branches. Incl. Veatchia A. Gray (1884) (nom. illeg., ICN Art. 53.1). Type: Veatchia cedrosensis A. Gray. Tree-like, 3–5 ( 10) m, dioecious, droughtdeciduous; trunk to 1 m ∅, wood soft, with yellowish, reddish, brownish or milky resinous exudate, bark pale ochre, peeling in papery pieces; Br massively thickened, often gnarled and contorted, with contracted short shoots from the axils of the primary leaves; L  asymmetrically pinnate, 1.5–8  0.7–3 cm overall; Inf lax to dense terminal panicles; Sep 5; Pet 5, elongate-ovate, pale to dark pink, becoming paler during anthesis, drying and persistent; St 10; Ov unilocular; Sty 3; Sti capitate; Fr small, dry and bladder-like, shortly tomentose.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

The genus has a single species only: P. discolor (Bentham) Coville ex Standley (Contr. US Nation. Herb. 23(3): 671, 1923). Type: Mexico, Baja California (Anonymus s.n. [BM?]). – Lit: Turner & al. (2005: ecology); Bashan & al. (2006: ecology). Distr: Mexico (Baja California, Baja California Sur); rocky slopes in Sonoran Desert vegetation, 0–800 ( 1200) m. I: Eggli (2008: 164). – Figs. 1 and 2.  Schinus discolor Bentham (1844)  Veatchia discolor (Bentham) T. S. Brandegee (1889) (incorrect name, ICN Art. 11.4); incl. Rhus veatchiana Kellogg (1863)  Veatchia discolor var. veatchiana (Kellogg) I. M. Johnston (1924) (incorrect name, ICN Art. 11.4)  Pachycormus discolor var. veatchianus (Kellogg) Gentry (1949); incl. Veatchia cedrosensis A. Gray (1885) (incorrect name, ICN Art. 11.4); incl. Bursera pubescens S. Watson (1889)  Veatchia discolor var. pubescens (S. Watson) I. M. Johnston (1924) (incorrect name, ICN Art. 11.4)  Pachycormus discolor var. pubescens (S. Watson) Gentry (1949). Description as for the genus. The species is variable and is sometimes divided into 3 varieties on account of leaf and inflorescence characters, but intermediate forms are common. Vernacular names: “Copalquin”, “Torote Blanco”, “Elephant Tree”. The stems of P. discolor are photosynthetically active due to a chlorophyll-containing phelloderm overlaid by a translucent phellem – a unique

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_3

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Fig. 1 Pachycormus discolor. (Copyright: J. Trager)

consists of mostly unlignified secondary xylem cells filled with starch grains. The wood offers little structural support, and stability is almost exclusively due to turgor pressure, and perhaps also to the tightly packed starch grains that serve as matrix for water storage (Gibson 1981). Water stored in the stem is used to buffer diminishing soil water availability at the beginning of the dry season, and stored water could support wholeplant water use for 22 days (Nilsen & al. 1990). Photosynthesis is of the usual C3 pathway (Nilsen & al. 1990, Franco-Vizcaíno & al. 1990). Plants are slow-growing in cultivation. First flowering occurred on a pot-grown plant after over 30 years, when the specimen was 85 cm tall, with a 4 cm thick basal thickening (Schmied 2016). Brand (2018) reports and illustrates plants parasitized by the holoparasite Cuscuta veatchii.

References Fig. 2 Pachycormus discolor (female inflorescence, cult. Sukkulenten-Sammlung Zürich, without known wild origin). (Copyright: U. Eggli)

arrangement according to Gibson (1981). The spongy bark can be as much as 6 cm thick (Veatch 1916) and is the primary site of water storage. Water is also stored in the wood, which

Bashan, Y. [& al. 2006], Vierheilig, H., Salazar, B. G. & Bashan, L. E. de (2006) Primary colonization and breakdown of igneous rocks by endemic, succulent elephant trees (Pachycormus discolor) of the deserts in Baja California, Mexico. Naturwissenschaften 93(7): 344–347, ills. https://doi.org/10.1007/s00114-006-0111-4. Brand, T. (2018) Cuscuta veatchii – ein pflanzlicher Parasit auf Pachycormus discolor. Kakt. and. Sukk. 69(10): 297–301, ills.

Pachycormus ANACARDIACEAE Eggli, U. (2008) Sukkulenten. Ed. 2. Stuttgart (DE): Eugen Ulmer. Franco-Vizcaíno, E. [& al. 1990], Goldstein, G. & Ting, I. P. (1990) Comparative gas exchange of leaves and bark in three stem succulents of Baja California. Amer. J. Bot. 77(10): 1272–1278. http://www.jstor.org/stable/ 2444588 Gibson, A. C. (1981) Vegetative anatomy of Pachycormus (Anacardiaceae). Bot. J. Linn. Soc. 83(4): 273–284, ills. https://doi.org/10.1111/j.1095-8339.1981. tb00351.x. Humphrey, R. R. (1991) Baja’s sacred Resin Tree, Pachycormus discolor. Cact. Succ. J. (US) 63(1): 35–39, ills.

21 Nilsen, E. T. [& al. 1990], Sharifi, M. R., Rundel, P. W., Forseth, I. N. & Ehleringer, J. R. (1990) Water-relations of stem succulent trees in north-central Baja California. Oecologia 82(3): 299–303, diags. https://doi.org/10. 1007/BF00317474. Schmied, R. (2016) Pachycormus discolor. Kakt. and. Sukk. 67(2): 40–42, ills. Turner, R. M. [& al. 2005], Bowers, J. E. & Burgess, T. L. (2005) Sonoran Desert plants. An ecological atlas. Tucson (US): University of Arizona Press. Veatch, J. A. (1916) Pachycormus discolor. In: Goldman, E. A. (ed.): Plant records of an expedition to Lower California. Contr. US Nation. Herb. 16(14): 344–345. http://biodiversitylibrary.org/page/372252

Part II Begoniaceae

Begoniaceae U. Eggli

Begoniaceae Agardh (Aphor. Bot., 200, 1824). Type: Begonia Linné. Nom. cons. (ICBN 2006, 140). See also the note by Turland & Watson, Taxon 54(2): 493, 2005. Order: Cucurbitales Distribution: Almost worldwide in the tropics and subtropics, but absent from Australia. Perennial or rarely annual herbs, or sometimes woody shrubs or climbing to trailing epiphytes, or deciduous and tuberous or rhizomatous, mesophytic to somewhat succulent; stems usually jointed, often juicy-fleshy but only rarely truly succulent, glabrous or variously hairy, sometimes with pearl glands; L spirally arranged or rarely distichous, simple or palmately lobed or incised, usually distinctly asymmetrical, rarely peltate, usually distinctly petiolate and with mostly large conspicuous persistent stipules; Inf axillary thyrses with extensive cymose branches, often long-pedunculate; Fl small to medium-sized, conspicuous, strictly unisexual (but plants monoecious and usually proterandrous, very rarely dioecious, organs of the missing sex often completely absent), often white, actinomorphic

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

or (due to unevenly developed wings on the ovary) zygomorphic; Per uniseriate and consisting of petaloid tepals (Begonia) or with separate sepals and petals (Hillebrandia); Tep (2–) 3–5 (–11), usually completely free, those of different whorls sometimes different in size or shape, in male flowers mostly 4 in 2 whorls, in female flowers mostly 5 in a single whorl; St 3–116, free or basally united; Ov inferior (Begonia) or semi-inferior (Hillebrandia), mostly with (1–) 3 (–7) distinct (and sometimes unequal) wings, mostly consisting of 2–3 united carpels and (1- to) 3- (to 7-) locular, with axile or rarely parietal septal or pseudo-axile placentation and numerous ovules; Sti 3 or 6, usually elongated, contorted and yellow (mimicking anthers; deceit pollination); Fr loculicidal (sometimes also septicidal) winged capsules, rarely fleshy or berries; Se numerous, minute, without endosperm. Important Literature: Carlquist (1985: wood anatomy); Wilde (2011: synoptical overview); The family Begoniaceae embraces only 2 genera, Hillebrandia (1 species only, H. sandwicensis, from the montane rain forest zone of Hawaii, not succulent) and Begonia (1870 species). The family appears in a derived position in the order Cucurbitales (Zhang & Renner 2003, Zhang & al. 2006), as sister to Datiscaceae (1 genus only with 2 species, not succulent, but forming basal tubers according to Stevens (2001+: accessed May 2018)), in a clade of several families characterized

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by having unisexual flowers. Sun & al. (2016: 370) found a similar topology, with Begoniaceae as sister to a clade formed by Datiscaceae + Tetramelaceae (2 monotypic genera only, no succulents), though with limited support. The divergence time is estimated at 24.6 mybp by Moonlight & al. (2018), superseding the earlier estimate of 51–65 mybp by Clement & al. (2004). Either a boreotropical or South-Pacific-Malesian origin was postulated by Clement & al. (2004) and Goodall-Copestake & al. (2008: 376), with subsequent range shifts towards the tropics during speciation during the global cooling phase between mid-eocene and early oligocene. More recently, Moonlight & al. (2018) argue that the Hawaiian occurrence of Hillebrandia is best explained by a single dispersal from Africa to Hawaii. Only a handful of species of Begonia can be considered as being truly succulent to some degree, mostly in the form of somewhat succulent leaves and petioles, or as tuberous caudices of various anatomical derivation.

References Carlquist, S. (1985) Wood anatomy of Begoniaceae, with comments on raylessness, paedomorphosis, relationships, vessel diameter, and ecology. Bull. Torrey Bot. Club 112(1): 59–69, ills. http://www.jstor.org/stable/ 2996105 Clement, W. L. [& al. 2004], Tebbitt, M. C., Forrest, L. L., Blair, J. E., Bouillet, L., Eriksson, T. & Swensen, S. M. (2004) Phylogenetic position and biogeography of Hillebrandia sandwicensis (Begoniaceae): A rare Hawaiian relict. Amer. J. Bot. 91(6): 905–917. https:// doi.org/10.3732/ajb.91.6.905.

U. Eggli Goodall-Copestake, W. P. [& al. 2008], Harris, D. J. & Hollingsworth, P. M. (2008) The origin of a megadiverse genus: Dating Begonia (Begoniaceae) using alternative datasets, calibrations and relaxed clock methods. Bot. J. Linn. Soc. 159(3): 363–380. https:// doi.org/10.1111/j.1095-8339.2009.00948.x. Moonlight, P. W. [& al. 2018], Ardi, W. H., Padilla, L. A., Chung, K.-F., Fuller, D., Girmansyah, D., Hollands, R., Jara-Muñoz, A., Kiew, R., Leong, W.-C., Liu, Y., Mahardika, A., Marasinghe, L. D. K., O'Connor, M., Peng, C.-I., Pérez, A. J., Phuttai, T., Pullan, M., Rajbhandary, S., Reynel, C., Rubite, R. R., Sang, J., Scherberich, D., Shui, Y.-M., Tebbitt, M. C., Thomas, D. C., Wilson, H. P., Zaini, N. H. & Hughes, M. (2018) Dividing and conquering the fastest-growing genus: Towards a natural sectional classification of the megadiverse genus Begonia (Begoniaceae). Taxon 67(2): 267–323, ills. https://doi.org/10.12705/672.3. Stevens, P. F. (2001+) Angiosperm Phylogeny Website, Version 14, July 2017 [and more or less continuously updated since]. St. Louis (US): Missouri Botanical Garden. http://www.mobot.org/MOBOT/research/ APweb/ Sun, M. [& al. 2016], Naaem, R., Su, J.-X., Cao, Z.-Y., Burleigh, J. G., Soltis, P., Soltis, D. E. & Chen, Z.-D. (2016) Phylogeny of the Rosidae: A dense taxon sampling analysis. J. Syst. Evol. 54(4): 363–391. https:// doi.org/10.1111/jse.12211. Wilde, J. J. F. E. de (2011) Begoniaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 56–71, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_5. Zhang, L.-B. & Renner, S. S. (2003) Phylogeny of Cucurbitales inferred from seven chloroplast and mitochondrial loci. sine loco (US): Botany 2003. Zhang, L.-B. [& al. 2006], Simmons, M. P., Kocyan, A. & Renner, S. S. (2006) Phylogeny of the Cucurbitales based on DNA sequences of nine loci from three genomes: Implications for morphological and sexual system evolution. Molec. Phylogen. Evol. 39(2): 305–322. https://doi.org/10.1016/j.ympev.2005. 10.002.

Begonia BEGONIACEAE U. Eggli

Begonia Linné (Spec. Pl. [ed. 1], 1056, 1753). Type: Begonia obliqua Linné.
Lit: Irmscher (1925: outdated monograph); Smith & Wasshausen (1986: outdated key & checklist); Doorenbos & al. (1998: sectional classification); Golding & Wasshausen (2002: key & checklist); Wilde (2002: revision sect. Tetraphila); Lebrun & Stork (2003: 470–490, checklist tropical Africa); Forrest & Hollingsworth (2003: circumscription); Plana (2003: phylogeny Africa & Madadascar); Plana & al. (2004: speciation in Africa); Forrest & al. (2005: molecular phylogeny); Kiew (2005: pictorial synopsis peninsular Malaysia); GoodallCopestake & al. (2010: molecular phylogeny); Thomas & al. (2011: molecular phylogeny Asian taxa); Wilde & al. (2011: long-distance dispersal); Thomas & al. (2012: dispersal & diversification); Moonlight & al. (2015: biogeography & diversification neotropical taxa); Gregório & al. (2016: flora Bahia, Brazil); Kidner & al. (2016: evolution of woodiness); Moonlight & al. (2018: phylogeny & sectional classification); Hughes & Peng (2018: pictorial synopsis Asia). Distr: As for the family. Etym: For Michel Begon (1638–1710), French Governor of Santo Domingo and promotor of botany.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

Incl. Diploclinium Lindley (1847). Type: Begonia evansiana Andrews. Incl. Augustia Klotzsch (1854). Type: not typified. Incl. Ewaldia Klotzsch (1854). Type: Begonia lobata Schott [lectotype, designated by Barkley & Baranov, Buxtonian 1(Suppl. I): 4, 1972]. Incl. Gireoudia Klotzsch (1854). Type: Begonia involucrata Liebman [lectotype, designated by Barkley & Baranov, Buxtonian 1(Suppl. I): 4, 1972]. Incl. Gurltia Klotzsch (1854). Type: Begonia meyeri Otto & A. Dietrich [automatic type, only element included]. Incl. Mitscherlichia Klotzsch (1854) (nom. illeg., ICN Art. 53.1). Type: Begonia albococcinea Hooker. Incl. Petermannia Klotzsch (1854). Type: Petermannia cumingiana Klotzsch [lectotype, designated by Barkley & Baranov, Buxtonian 1 (Suppl. I): 6, 1972]. Incl. Pritzelia Klotzsch (1854). Type: Begonia fischeri Otto & A. Dietrich [lectotype, designated by Barkley & Baranov, Buxtonian 1 (Suppl. I): 6, 1972]. Incl. Rachia Klotzsch (1854). Type: not typified. Incl. Saueria Klotzsch (1854). Type: Begonia sulcata Scheidweiler [automatic type, only element included]. Incl. Scheidweileria Klotzsch (1854). Type: Begonia muricata Scheidweiler [lectotype, designated by Barkley & Baranov, Buxtonian 1(Suppl. I): 7, 1972].

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Incl. Trendelenburgia Klotzsch (1854). Type: Trendelenburgia fruticosa Klotzsch [automatic type, only element included]. Incl. Nephromischus Klotzsch (1855). Type: Begonia rutilans Van Houtte [automatic type, only element included]. Incl. Steineria Klotzsch (1855). Type: Steineria ferruginea Klotzsch [lectotype, designated by Barkley & Baranov, Buxtonian 1(Suppl. I): 6, 1972]. Incl. Weilbachia Klotzsch & Oersted ex Klotzsch (1855). Type: Weilbachia reptans Klotzsch [automatic type, only element included]. Incl. Begoniella Oliver (1872). Type: Begoniella whitei Oliver. Incl. Symbegonia Warburg (1894). Type: Symbegonia fulvo-villosa Warburg. Incl. Semibegoniella C. De Candolle (1908). Type: Semibegoniella sodiroi C. De Candolle [lectotype, designated by Baranov & Barkley, Sect. Genus Begonia, 1974]. Description largely as for the family; Tep normally 2 þ 2 in male and 5 in female flowers. The genus with its 1870 species (Moonlight & al. 2018) is well-known for numerous important horticultural taxa, such as the B. semperflorens hybrids and the tuberous B. tuberhybrida cultivars, all frequently seen in gardens and on balconies. Several other taxa are valued as pot plants for indoor cultivation because of the often vividly coloured leaves. Begonia is the 6.-largest genus of flowering plants, and its full diversity is only coming to light in the past years, with 341 species described in the short period 2005–2015, and many more still undescribed (Moonlight & al. 2018)
this makes it the fastest-growing genus of plants. Classification: Past attempts to subdivide Begonia into as many as 39 segregate genera (Klotzsch 1855) were not accepted by the majority of authors, and even the classification by Irmscher (1925), who in addition to Begonia accepted Semibegoniella, Begoniella and Symbegonia, was not followed. At present, there is unanimous agreement that only the genera Hillebrandia and Begonia should be recognized

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in the family (Wilde 2011: 65, Moonlight & al. 2018). Klotzsch’s several dozen segregated genera are only listed above when placed in the synonymy of any of the sections for which species are covered in the synopsis below; a full list can be found in Smith & Schubert (1958: 41–42) and in Moonlight & al. (2018). Berg (1985) proposed an African origin for the genus; this hypothesis is neither clearly supported nor refuted by the molecular data presented by Forrest & al. (2005). However, Goodall-Copestake & al. (2010) and subsequent studies such as those of Moonlight & al. (2018) found good support that Begonia in fact first diversified in Africa, with subsequent dispersal to America and Asia. The closest African relatives of the American and Asian members are African species adapted to seasonal climates. The initial migration of Begonia from Africa to Asia is dated to early or mid-eocene (Wilde & al. 2011), coincident with a warm and moist mid-miocene climatic optimum at 17–15 mybp. The diversification of the genus in Asia was further studied by Thomas & al. (2012), revealing a general trend of dispersal from W to E and “cyclic vicariance” by habitat fragmentations and amalgamations. The infrageneric classification of Begonia proved to be a challenge for a long time: Doorenbos & al. (1998) divide the genus into 63 sections, and Wilde & al. (2011) argue for accepting 65 sections. Smith & Wasshausen (1986) (with amendments and additions provided by Golding & Wasshausen (2002)) provided a now somewhat outdated checklist and an artificial key to the species of the genus. The most complete study of Begonia classification is that of Moonlight & al. (2018), who divide the genus into 70 sections. Many of the characters or suites of characters that were treated as having special value for the delimitation of infrageneric taxa have been found to be variable and homoplasious, resulting in polyphyletic taxa in past infrageneric classifications (Thomas & al. 2011). 3 major clades were distinguished already in early analyses (Forrest & Hollingsworth 2003, Plana & al. 2004, Forrest & al. 2005): A small strictly African clade, a clade

Begonia BEGONIACEAE

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embracing S African plus all Madagascan species, and a clade with S African, Socotran and all American and Asian species. Notably, the Madagascan taxa form a monophyletic group. In the most comprehensive recent phylogeny, Moonlight & al. (2018) confirmed that all clades in an early-diverging grade are African. The remainder of the genus is made up by a neotropical clade 1 plus a sister pair of an Asiatic clade and a neotropical clade 2.
Fig. 1. Interestingly, the most basal lineages in each of these 3 major clades are made up by African taxa. Goodall-Copestake & al. (2010) and Hughes & al. (2018) propose that interspecific hybridization likely played a role in the evolution of the genus. Succulence: Many taxa are commonly described as having fleshy or succulent stems, and/or leathery to somewhat fleshy leaves, but most of these taxa inhabit rainforests or other humid places. However, species from such habitats such as B. loranthoides (esp. its ssp. rhopalocarpa) predominantly grow as epiphytes on tall emergent forest trees, as well as on rocky (often limestone) hills that emerge from the

Y

F M S Neotr-1 P

Asian

tropical forest, i.e. habitats with periodical shortterm water shortage where thick succulent leaves are of adaptive value (Wilde & Arends 1979: 372). Judged from experience in cultivation, the degree of succulence esp. of the epiphytes is often under environmental control (e.g. B. polygonoides, pers. obs.). It remains to be investigated to what extent thickened fleshy leaves are truly succulent to buffer water shortage at least in the short term, and therefore, only a selection of such species are covered in the synopsis below. Sheue & al. (2012: 1070) illustrate single- or double-layered water storage tissues devoid of chloroplasts even for the hardly succulent B. chlorosticta and an unnamed cultivar, respectively. In more succulent taxa, such as B. polygonoides, leaves can be as thick as 3–4 mm, with well-visible water storage tissues. Rudall & al. (2018) consider this water storage tissue to represent a hypodermis, often present on both leaf faces (but only 5 species were investigated), while other authors (e.g. Tang & al. (2002) or Kiew (2005: 10)) call it a multiple epidermis. According to Rudall & al. (2018), the leaves of xerophytic species with well-developed water storage hypodermis possess

A

Neotropical-2

300 species

Fig. 1 Summary phylogeny of Begonia, based on Moonlight & al. (2018) (1870 species total). A ¼ Sect. Augustia (¼ SDAAB2-clade [Seasonally-Dry Adapted African Begonia]), F ¼ FFAB-clade (Fleshy-Fruited African Begonia), M ¼ MB-clade (Madagascan Begonia), Neotrop-1 ¼

Neotropical Clade 1, S ¼ SDAAB1-clade (Seasonally-Dry Adapted African Begonia), P ¼ Sect. Peltaugustia, Y ¼ YFAB-clade (Yellow-Flowering African Begonia). (Copyright: U. Eggli)

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non-contiguous stomatal clusters, confined to the abaxial leaf face, while mesophytic species without specialized hypodermis have single stomata, rather than clusters. Stem succulence is found in a few taxa such as B. dregei. Many species are occasionally described with thickened fleshy stems, and the degree of stem thickening appears to be plastic at least for some taxa. B. dipetala Graham, e.g., is usually described as a mesophytic herb, but Matulová (2007) illustrates a specimen with a distinctly thickened main stem. Without more data about the possible succulence of this and similar other species, they are not further covered in the synopsis hereafter. Tuberous bases, rhizomes and other caudexlike water storage structures can have very different morphological origins (Forrest & al. 2005): The Socotran endemic B. samhaensis has an irregular tuber and bulbils comprised of a condensed shoot with reduced, scale-like fleshy leaves, while B. socotrona has only bulbils but no tuber. B. sutherlandii produces bulbils from leaf axils, and has underground tubers probably derived from roots. In B. dregei and related species of Sect. Augustia, the initial basal caudex likely represents a hypocotyl tuber. Parallel developments and convergence has likely occurred (Forrest & al. 2005). In other species, the tubers are either hypocotylar in origin, or represent thickened portions of creeping rhizomes (Wilde 2011). Assessing the degree of succulence in the tuberous begonias is difficult. Although many grow sympatrically with cacti in S America, it is unknown to what degree they remain physiologically active during the dry season, and they are therefore omitted from this synopsis. B. albidula Brade (Arq. Jard. Bot. Rio de Janeiro 10: 137–138, t. 7, 1950). Type: Brazil, Espírito Santo (Brade 19785 [RB]).
Distr: E Brazil (S Espírito Santo); rocky outcrops, amongst Velloziaceae, 900–1200 m. [Sect. Tetrachia] Perennial erect shrublets to 1 m; stem branched, succulent, 1–1.5 cm ∅, terete, white-tomentose becoming glabrous; stipules ovate, caducous, 2.5–4.5  1.8 cm, outer face

U. Eggli

lanate; L 0.6–1 cm petiolate, lamina strongly asymmetric, transversely ovate, 8–12  4–6 cm, stiffly thickish-succulent, base cordate, tip acute, margin almost entire, upper face densely whitewoolly becoming glabrous, lower face densely white-tomentose; Inf peduncle erect, 8–10 cm and surpassing the leaves, several times dichotomously branched; Fl white; male Fl with 4 tepals, OTep 10  9 mm, outer face minutely tomentose, ITep 7  3.5 mm; female Fl with 5 tepals, 2 broader and 3 narrower, 8–9  4.5–6 mm, outer face with stellate hairs; Sty 3, bifurcate; Fr roundish, 10–12 mm,  tomentose. B. blancii M. Hughes & C.-I. Peng (Bot. Stud. (Nangang) 52: 204–205, ills., 2011). Type: Philippines, Palawan (Blanc & Rubite 09–201 [PNH, E, SING]).
Distr: Philippines (Palawan), on the sides of boulders in the shade of primary forest, 300–400 m; known only from the type locality. [Sect. Baryandra] Perennial creeping herbs; stem with internodes to 1 cm, 5–9 mm ∅, succulent, with up to 5 mm long hairs; stipules triangular,  1.5  1.2 cm, hairy, persistent, papery with age; L appressed to the substrate in plants on vertical rock faces, otherwise more erect, petiole 1–2 cm in young leaves, extending to 5–7 cm, L lamina distinctly fleshy-succulent, 6–11  4–8 cm, oblong-spatulate, asymmetrical, green mottled with darker green, solid dark green with paler veins, or uniformly blackish-green, upper face glabrous, lower face hairy on the veins, margins entire, somewhat undulate; Inf to 20 cm, cymose,  10-flowered; peduncle to 15 cm, scatteredly hairy, flower pairs consisting of a male and a female flower open at the same time; Ped 15–20 mm; male Fl with 4 tepals, OTep 2, ovate, pink on the outside, white to pale pink otherwise, 18  10 mm, ITep 2, obovate, white, 15  7 mm; female Fl similar but ITep pale pink. The species grows on the sides of almost bare rock boulders. The succulent leaves remain green throughout the dry season. Similarities exist to other Malesian species with somewhat coriaceous leaves as well, but the triangular-spatulate leaves immediately distinguish the species.

Begonia BEGONIACEAE

B. bogneri Ziesenhenne (Begonian 40: 76, ills., 1973). Type: Madagascar, Antsiranana (Bogner 262 [M, K, M, P, US]).
Lit: Sands (1986). Distr: NE Madagascar (Antsiranana); mossy granite cliffs, in shade, 50 m. I: Rauh (1998: 337). [Sect. Erminea] Perennial tufted herbs, stemless or with several stems to 3.5 cm from a somewhat tuberous base; stipules triangular, membranous, flushed pink, to 0.4  0.2 cm, marcescent; L grass-like, arching over, to 15 cm  2 (
5) mm, 2 mm thick and weakly succulent, grass-green, margin with small irregularly spaced spicules; Inf to 15 cm with 1–2 male and 1 female Fl  20 mm ∅; Tep pale pink, of subequal size, 4 in male and 6 in female Fl; St basally united; Sti 3, horse-shoe-shaped when seen from the side; Fr with 1 wing only. Closely related to the other species of Sect. Erminea, all small herbs with tubers, and from humid places in Madagascar. B. bogneri is easily recognised by its unusual grass-like leaves. In nature, the plants are probably deciduous, while in cultivation, the leaves are evergreen. The recently described B. pteridoides (Scherberich & Duruisseau 2017) from mossy granite boulders is similar with more divided but less succulent leaves. B. cavallyensis A. Chevalier (Bull. Soc. Bot. France 58 (Mém. 8d): 176, 1912). Type: Ivory Coast (Chevalier 21422 [P]).
Lit: Hagman & Wilde (1984: with ills.); Wilde (2002: 52, with ills.). Distr: Tropical W Africa (Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana); epiphyte in forest, open shade to full sun, 500–1500 m. [Sect. Tetraphila] Perennial herbaceous stem succulent epiphytes, all plant parts with whitish globose hairs; stems to 90 cm, to 1.6 cm ∅ at the base, sturdy, erect, thick, grey-green to dark red-brown, with conspicuous leaf scars, Int 0.4–3 cm (to 8.5 cm in cultivation); stipules conspicuous, boat-shaped, (1.5–) 2.7–4.3  (0.4–) 1.3–1.5 cm, narrowly oblong to triangular, red with often transparent margins, membranous, early caducous; L few, congested near the stem tips, petiole (0.8–) 2–7 (
15) cm, red, L lamina ovate, slightly obliquely asymmetrical with

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obtuse to cordate base, 4–16 (
24)  2–10 (
15) cm, thick and somewhat fleshy, with acute to obtuse tip, green with red veins and a conspicuous dark red subentire margin to 4 mm wide; Inf uni- or bisexual; male Inf (2- to) 4- to 15-flowered dichasial cymes, peduncle 0.15–4.5 (
5.5) cm; female Inf 2- to 3-flowered cymes, peduncle to 0.55 cm; all Fl with 4 tepals or the inner pair missing; OTep pink to pinkish-white, with pink veins, broadly ovate to elliptic, 3.5–10 (
14)  3–8 (
11) mm, somewhat smaller in female flowers; ITep almost white, narrowly obovate to narrowly oblong, 4–7.5  0.7–2.5 mm, somewhat smaller in female flowers. This stem-succulent species was formerly treated as synonym of B. eminii Warburg but the careful analysis by Hagman & Wilde (1984) re-established it as separate species, most closely related to the non-succulent B. fusicarpa Irmscher. B. conchifolia A. Dietrich (Allg. Gartenzeitung 19: 258, 1851). Type: Costa Rica (Warszewicz s.n. [B]).
Lit: Brennan & al. (2012: genetics). Distr: Costa Rica (Alajuela, Cartago, Guanacaste, Heredia, Puntarenas, San José), El Salvador (Santa Ana), Nicaragua (Jinotega), Panama; rock outcrops in subhumid forests, occasionally in grasslands, (300–) 600–2000 m.  Gireoudia conchifolia (A. Dietrich) Klotzsch (1854); incl. Begonia scutellata Liebmann (1853)  Gireoudia conchifolia var. scutellata (Liebmann) Klotzsch (1855); incl. Gireoudia conchifolia var. warscewicziana Klotzsch (1855); incl. Gireoudia warscewicziana Hort. Berol. ex A. De Candolle (1864); incl. Begonia pumilio Standley (1940) (nom. illeg., ICN Art. 53.1); incl. Begonia conchifolia var. rubrimacula Golding (1973)  Begonia conchifolia fa. rubrimacula (Golding) Tebbitt (2005). [Sect. Gireoudia] Perennial  stemless usually much-branched herbs with creeping rhizomes to 0.5 cm ∅; Int 1–1.5 mm; stipules 0.6–1.5  0.2–0.9 mm, asymmetrically ovate, brown, persistent, glabrous or sparsely tomentose; L petiole (2.5–) 3–10 (
28 (
33)) cm, lamina obliquely peltate, (almost) symmetrical, fleshy-succulent

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and stiff, 2–9 (
14)  1–8 (
10) cm, broadly ovate to suborbicular, tip acute to acuminate, margins irregularly shallowly denticulate and undulate, upper face glabrous or almost so, somewhat shiny, lower face esp. on the veins brownishtomentose, veins (6–) 7–8 (
9); Inf to 15–30 cm, bisexual, often  unilaterally disposed, few- to many-flowered; Fl 4–12 mm pedicellate, white; male Fl OTep 3–7  3–7 (
8) mm, suborbicular to elliptic, pink, ITep absent or rarely 1, 6  2.5 mm; female Fl similar to male flowers, OTep 2, 2.5–7  2–6 mm, ITep absent or rarely 1,  5  2 mm; Fr subglobose, 4–10 mm, wings slightly unequal. Frequently seen in the horticultural trade, and popular as houseplant. The leaves of some forms have an attractive paler venation and/or a red blotch in the centre where the petiole attaches (var./fa. rubrimacula). The leaves of most forms are distinctly succulent (see Rudall & al. (2018: 769) for a microscope illustration of a leaf crosssection, showing a voluminous succulent hypodermis on either leaf face). B. crassula Averyanov (Turczaninowia 15(2): 8–11, ills., 2012). Type: Vietnam, Quang Binh Prov. (Hiep & al. 385800 [LE, CPC]).
Distr: C Vietnam (Quang Binh Prov.); lithophytic on open karstic limestones, 200–800 m. [Sect. Diploclinium according to the protologue, unassigned in Moonlight & al. (2018)] Perennial herbaceous shrublets, entirely glabrous; stems mostly unbranched, ascending to erect, 50–150 cm, succulent and to 0.8 cm ∅, green, often with purple tint; stipules ovate, (1–) 2–5 (
6)  1–2 cm, pale, leathery, caducous; L petiole (1–) 2–7 (
9) cm, somewhat succulent, light green, with purple base and tip, L lamina strongly asymmetrical, (narrowly) ovate in outline, fleshysucculent, (1.5–) 3–6 (
7)  (4–) 6–18 (
22) cm, with broadly cuneate base and attenuate apex, uniformly glossy-green, margin entire or indistinctly denticulate; Inf apparently unisexual, axillary, on middle and upper stem parts, 5- to 20-flowered, 4–10 (
12) cm; peduncle 1–3 (
5) cm, green to purple; Fl light pink; Ped (5–) 8–12 (
15) mm, pink to purple; male Fl with 2 tepals only, Tep broadly ovate to orbicular, (6–) 8–12

U. Eggli

(
14) mm long and wide; female Fl with 4 tepals, slightly zygomorphic, OTep subopposite, broadly reniform to almost orbicular, (6–) 8–10 (
12) mm long and wide, ITep obliquely (broadly) obovate, only (4–) 5–7 (
8) mm wide. According to the protologue “a typical element of xerophytic herbaceous vegetation”. B. dregei Otto & A. Dietrich (Allg. Gartenzeitung 4: 357, 1836). Type: RSA, Eastern Cape (Drège s.n. [B]).
Lit: McLellan (2000: geographic variation leaves); Matolweni & al. (2000: genetic diversity). Distr: RSA (Eastern Cape, KwaZulu-Natal); rocky cliffs in coastal forests, 0–1220 m. I: Flow. Pl. South Afr. 17: t. 673, 1937, as B. richardsiana; Eggli (2008: 219); Lenain (2012: 71).
Fig. 2.  Augustia dregei (Otto & A. Dietrich) Klotzsch (1855); incl. Begonia parvifolia E. Meyer ex Otto & A. Dietrich (1836) (nom. inval., ICN Art. 36.1c); incl. Begonia suffruticosa Meisner (1840)  Augustia suffruticosa (Meisner) Klotzsch (1855); incl. Begonia reniformis Hort. Berol. ex Klotzsch (1854) (nom. illeg., ICN Art. 53.1); incl. Begonia natalensis Hooker (1855)  Augustia natalensis (Hooker) Klotzsch (1855); incl. Begonia rubicunda Hort. Turic. ex A. De Candolle (1864); incl. Begonia richardsiana T. Moore (1871); incl. Begonia richardsoniana Houllet (1872); incl. Begonia dregei var. macbethii L. B. Bailey (1923); incl.

Fig. 2 Begonia dregei (cult. Sukkulenten-Sammlung Zürich; without known wild origin). (Copyright: U. Eggli)

Begonia BEGONIACEAE

Begonia partita Irmscher (1961); incl. Begonia suffruticosa fa. bolusii Irmscher (1961); incl. Begonia suffruticosa fa. worsdelii Irmscher (1961). [Sect. Augustia] Glabrous perennial herbs; stems to 30 cm, softly fleshy, branched, with thickened nodes and basally forming a round to irregular tuber to 10 cm ∅ with smooth brownishgreen bark; stipules brownish, membranous, soon caducous; L with a reddish short to long petiole, lamina green, thin, asymmetrically ovate to ovatelanceolate, 5 (
8)  2 (
3.5) cm, base shallowly cordate, apex narrowly triangular-acute to acuminate, margin moderately to deeply incised; Inf from upper leaf axils, short, few-flowered; Fl to 20 mm ∅, white to pale pink; male Fl with 2 Tep, female Fl with 5 Tep; Fr 3-winged. This is a very variable taxon (esp. as to leaf shape and size), and Hilliard (1976) prefers not to differentiate formal infraspecific taxa, but recognizes 5 informal groups. According to Matolweni & al. (2000) B. dregei is a rare endemic known from small, scattered and isolated populations, often counting 4=5 of the leaf length, or almost entire and reniform (only A. malvifolia); Fl obliquely zygomophic, 5–7.5 cm ∅; Sep linear to oblong-ovate; Pet yellow, obovate with rounded apex, the upper 2 with 2 basal red streaks, the lateral 2 with 1 basal red streak, the lowermost

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

unmarked; St unequal in 2 groups; Fil and Anth of the lower group slightly larger and tinged dark red, those of the upper group uniformly yellow; Anth with 2 apical pores; Ov of 3 fused Ca, imperfectly 3-locular, pubescent becoming glabrous; Sty incurved, slightly longer than the lower group of stamens; Fr woody capsules, globose to ovate, acute-truncate, oriented downwards with incurved pedicel, glabrous, incompletely dehiscent and seeds dispersed through the irregular fissures of the endocarp of the withering fruit; Se reniform to globose, glabrous, pilose or minutely echinate. A small genus of 4 species which resemble each other strongly. Only leaf shape, segmentation and seed characters are diagnostic. The scattered occurrence in S America is enigmatic. In additon to the scattered records of A. palmatifida and A. wrightii in S America, a collection from Bolivia was described as A. unipora Tieghem, but this species is treated as synonym of Cochlospermum regium Pilger by Poppendieck (1981). Amoreuxia species can be classified as geophytic subshrubs due to their complex tuberous underground structures (Chávez & al. 2014). At least in A. wrightii, the tuber corresponds most likely to a stem-derived xylopodium plus a rootderived tuberous part that includes lateral roots as well. The tissue of the tuber is made up largely by parenchyma (Cedano Maldonado 2000: 85), and Chávez & al. (2014) described “dilatation parenchyma” that serves for starch and water storage

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_6

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between the first-formed tissue wedges
the xylopodia can thus be termed succulent at least to some degree. All plant parts are edible and esp. the roots have been an important food item for centuries in the Sonoran Desert area and elsewhere (Hodgson 2001: 91–94). This author notes that the seeds are difficult if not impossible to germinate in cultivation, but Celaya-Michel & al. (2017) report that seeds germinate easily after scarification (in their experiment by sanding). A. gonzalezii Sprague & L. Riley (Bull. Misc. Inform. Kew 1922(3): 102, 1922). Type: Mexico, Sinaloa (Ortega 897 [K, US]).
Lit: Pío-León & al. (2014: with ill.). Distr: S USA (S Arizona), NW Mexico (Sonora, Baja California Sur, Sinaloa, Jalisco); stony to rocky hillsides in open forests, 400–1400 m. I: Cedano Maldonado (2000: 207); Hodgson (2001: 93).  Cochlospermum gonzalezii (Sprague & L. Riley) Byng & Christenhusz (2018). L palmatipartite, 2.5–6  3.5–9 cm, bright green, with (5–) 7–9 spatulate to oblanceolate lobes; Fr ellipsoid, 4.5–8 cm; Se (sub-) globose, slightly pilose, hairs sometimes stiff. According to Poppendieck (1981) morphologically intermediate between A. wrightii (globose seeds) and A. palmatifida (7 to 9 spatulate leaf lobes), but not occurring sympatric with the former anywhere. Johnson-Fulton & Watson (2017) identified the species as sister of A. wrightii. A. gonzalezii is a rare species, known from fewer than 10 localities (Johnson-Fulton & Watson 2015). The occurrence in Jalisco was reported by Cedano Maldonado (1998). A. malvifolia A. Gray (Smithsonian Contr. Knowl. 3(5): 29, 1852). Type: Mexico, Chihuahua (Anonymus s.n. [K]).
Distr: N Mexico (N Mexican Plateau: Chihuahua, Durango); rocky hills to clay flats, 1000–1500 (
1900) m. I: Poppendieck (1981: 21, fig. 4); Cedano Maldonado (2000: 236).  Cochlospermum malvivolium (A. Gray) Byng & Christenhusz (2018). L reniform, subentire to at most very shallowly 5- to 7-lobed, 2.5–5  4–7.5 cm, purple grey-

U. Eggli

green, lobe sinuses at most 3–5 mm deeper than the leaf margin; Fr ovoid, 2–4 cm; Se reniform,  5 mm, minutely echinate. The subentire leaves are unique in the genus. In the molecular phylogeny of Johnson-Fulton & Watson (2017) the species is shown as sister to the geographically wide-spread A. palmatifida. A. palmatifida Moçiño & Sessé ex De Candolle (Prodr. Syst. Regni Veg. 2: 638, 1825). Type: Mexico (Herb. Pavón s.n. [BM [status?]]).
Lit: Hodgson (2001: 91–92, with ills.; ethnobotany). Distr: S USA (Arizona), Mexico (widespread from Sonora and Baja California to Chiapas), Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Colombia; desert to grassland or open spots in seasonally deciduous forest, in gravelly to loamy soils, 0–1600 m. I: Wiggins (1980: 628, fig. 604); Hodgson (2001: 91–92); Cedano Maldonado (2000: 196, 215 [as A. schiedeana]); Johnson-Fulton & Watson (2015: 375).  Cochlospermum palmatifidum (Moçiño & Sessé ex De Candolle) Byng & Christenhusz (2018); incl. Euryanthe schiedeana Chamisso & Schlechtendal (1830)  Amoreuxia schiedeana (Chamisso & Schlechtendal) Planchon (1847); incl. Amoreuxia colombiana Sprague (1922). L palmatipartite, 2–6  3.2–9 cm, bright green, with 7–9 spatulate to linear lobes to 1 cm broad; Fr ovoid, 2–4.2 cm; Se reniform, subglabrous to densely pilose or minutely echinate. This is a very variable species (see Poppendieck (1981: 23) for the geographical distribution of leaf shapes) which has its centre of diversity in NW Mexico. Based on minor differences in leaf and seed characters, Cedano Maldonado (2000: 193–194) treats A. palmatifida in a narrow sense, limited to NW Mexico, and accepts A. schiedeana as separate, much more widespread and largely allopatric species in C and S Mexico. A. wrightii A. Gray (Smithsonian Contr. Knowl. 3(5): 26, 1853). Type: USA, Texas (Wright 79 [GH, BM, US]).
Distr: S USA (Texas), NE to S Mexico (widespread but scattered, to Yucatán in the S), Cuba, Curaçao, Venezuela (Coro), Peru (Amazonas), Ecuador?; dry rocky or gravelly hills and plains to seasonally

Amoreuxia BIXACEAE

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enigmatic; Poppendieck (1981) records a single collection only, while Molau (1983) states that it was collected twice in N-most Peru, and is likely to occur in neighbouring Ecuador as well. In the USA as well as in Mexico, the species is considered to be endangered. Soto-Mata & al. (2018) studied the ecological preferences of populations in Nuevo León and found that A. wrightii prefers open disturbed sites with pioneer vegetation.

References

Fig. 1 Amoreuxia wrightii (cult. SukkulentenSammlung Zürich; without known wild origin). (Copyright: U. Eggli)

dry marshland, 0–400 m. I: Calderón de Rzedowski (1994: 3); Cedano Maldonado (2000: 227).
Fig. 1.  Cochlospermum wrightii (A. Gray) Byng & Christenhusz (2018); incl. Amoreuxia potentilloides Pilger (1936)  Cochlospermum potentilloides (Pilger) J. F. Macbride (1940). L palmatipartite, 3–5  4–7 cm, bright green, with 5 (
7) subrhomboidal to obovate lobes 1.8–3 cm wide; Fr oblong-obovoid, 3–6 cm; Se (sub-) globose, 3.5–5 mm ∅, glabrous. Poppendieck (1981: 7) argued that this species might have repeatedly arisen from the “basal stock” of A. palmatifida, but the molecular phylogeny of Johnson-Fulton & Watson (2017) clearly found A. wrightii to be the sister species of the rare and restricted A. gonzalezii. The occurrence on Cuba was reported by Günther (1986), and the occurrence in Venezuela is based on Chávez & al. (2014). The occurrence in Peru is

Calderón de Rzedowski, G. (1994) Família Cochlospermaceae. In: Rzedowski, J. & Calderón de Rzedowski, G. (eds.): Flora del Bajio y de regiones adyacentes, Fasc. 28. Pátzcuaro (MX): Instituto de Ecología A.C., Centro Regional del Bajío. Cedano Maldonado, M. (1998) Primer informe de Amoreuxia gonzalezii Sprague et Riley (Cochlospermaceae) en Jalisco. Bol. Inst. Bot. Univ. Guadalajara 5(1–3): 217–222, ills. Cedano Maldonado, M. (2000) Revisión de la familia Cochlospermaceae para México. Guadalajara (MX): Universidad de Guadalajara, Centro Universitario de Ciencias Biológicas y Agropecuarias, M.Sc. thesis. Celaya-Michel, H. [& al. 2017], Ochoa-Meza, A., LópezElías, J. & Barrera-Silva, M. A. (2017) Germinación y crecimiento en vivero y en campo de Zaya (Amoreuxia palmatifida DC.), una especie nativa amenazada en México. Eur. Sci. J. 13(24): 66–78, ills. https://doi. org/10.19044/esj.2017.v13n24p66. Chávez, J. [& al. 2014], Jáuregui, D., Lapp, M. & Torrecilla, P. (2014) Anatomía foliar y del sistema subterráneo de Amoreuxia wrightii A. Gray (Bixaceae), especie en peligro crítico en Venezuela. Ernstia, ser. 2, 23(1): 47–65, ills. http://saber.ucv.ve/ojs/index.php/ rev_erns/article/view/5175 Günther, K. F. (1986) A new record for Amoreuxia wrightii (Cochlospermaceae) for Cuba. Feddes Repert. 97(1–2): 73–78. Hodgson, W. C. (2001) Food plants of the Sonoran Desert. Tucson (US): University of Arizona Press. Johnson-Fulton, S. B. & Watson, L. E. (2015) Cochlospermaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 376–379. New York (US)/Oxford (GB): Oxford University Press. Johnson-Fulton, S. B. & Watson, L. E. (2017) Phylogenetic systematics of Cochlospermaceae (Malvales) based on molecular and morphological evidence. Syst. Bot. 42(2): 271–282, ills., maps. https://doi.org/ 10.1600/036364417X695457. Molau, U. (1983) Cochlospermaceae. In: Harling, G. & Sparre, B. (eds.): Flora of Ecuador No. 20: 11–15, ills., keys. Stockholm (SE): Publishing House of the Swedish Research Council.

60 Pío-León, J. F. [& al. 2014], León-De la Luz, J. L. & Ortega-Rubio, A. (2014) Nuevo registro de Amoreuxia gonzalezii (Bixaceae) para la península de Baja California, México. Revista Mex. Biodivers. 85(4): 1269–1272, ills. https://doi.org/10.7550/rmb.44212. Poppendieck, H.-H. (1980) A monograph of the Cochlospermaceae. Bot. Jahrb. Syst. 101(2): 191–265, ills., keys. Poppendieck, H.-H. (1981) Cochlospermaceae. Flora Neotropica, Monograph Number 27. New York (US): Organization for Flora Neotropica/New York Botanical Garden. https://www.jstor.org/stable/4393745

U. Eggli Soto-Mata, B. [& al. 2018], Alanís-Rodríguez, E., Jurado, E., Pando-Moreno, M., Molina-Guerra, V., AlcaláRojas, A. & Marroquín-Castillo, J. J. (2018) Caracterización del hábitat de Amoreuxia wrightii (Bixaceae), una especie en peligro de extinción en el noreste de México. Acta Bot. Mex. 122: 21–31, map. https://doi. org/10.21829/abm122.2018.1196. Sprague, T. A. (1921) A revision of Amoreuxia. Bull. Misc. Inform. Kew 1922: 97–105, t. 1. https://www. biodiversitylibrary.org/item/42567#page/102 Wiggins, I. L. (1980) Flora of Baja California. Stanford (US): Stanford University Press.

Cochlospermum BIXACEAE U. Eggli

Cochlospermum Kunth (in Humboldt & al., Nov. Gen. Sp. 5: 297 [ed. quarto], 5: 231 [ed. folio], 1822). Type: Bombax gossypium Linné [nom. illeg., ¼ Bombax religiosum Linné (ICBN 2006: 318)].  Cochlospermeae  Lit: Poppendieck (1980: monograph); Poppendieck (1981: Flora Neotropica). Distr: Pantropical incl. Australia; seasonally dry deciduous forests or grasslands (rarely lowland rain-forests). Etym: Gr. ‘kochlos’, snail; and Gr. ‘sperma’, seed; for the seeds of some species that are curved like a snail shell Incl. Maximilianea Martius (1819) (nomen rejiciendum, ICN Art. 56.1). Type: Maximilianea regia Martius. Incl. Wittelsbachia Martius & Zuccarini (1824). Type: Maximilianea regia Martius. Incl. Azeredia Arruda ex Allemão (1846). Type: Azeredia pernambucana Allemão [nom. illeg.,  Maximilanea regia Martius)]. Deciduous often xeromorphic trees to 20 m or shrubs or subshrubs with woody underground trunk (xylopodium ¼ lignotuber), with secretory cells with red or orange-yellow sap; L subentire to palmatilobed or palmatisect; stipules narrow; Fl

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

actinomorphic (but slightly zygomorphic in bud), 5–15 cm ∅; Sep equal or unequal; Pet yellow, with fine red marks or stripes, tip emarginate; St all equal, free or basally slightly connate; Ov consisting of 3–5 fused Ca; Fr completely dehiscent capsules; Se reniform to cochleate, with long hairs for wind dispersal. A genus of 12 species, that is divided into 2 subgenera: • Subgen. Cochlospermum: Pantropical; Anth with 1 apical pore  10 species. This taxon conforms to the “core Cochlospermum” of Johnson-Fulton & Watson (2017). • Subgen. Diporandra Planchon: S America; Anth with 2 apical pores  2 species, placed outside core Cochlospermum by JohnsonFulton & Watson (2017) and appearing in a position basal to Amoreuxia in some of their analyses. Young plants grow rapidly and some species can flower within 1 year from seed. Flowers are usually produced during the dry season when the plants are leafless (see e.g. C. tetraporum Hallier fil., illustrated by Richtr 2016). Succulence: The species Cochlospermum vitifolium Sprengel is characterized as tropical deciduous succulent tree by Borchert & Rivera (2001) due to is large somewhat swollen stems and low-density water-storing wood (Chapotin & al. 2003). The soft wood of C. religiosum (Linné)

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Alston is formed by consecutive rings of parenchymatous tissue interspersed with thin-walled fibres and vascular tissue with large lumina (Tieghem 1900: 43, as C. gossypium De Candolle). Wood water content is 0.75–0.8 g/cm3, falling to 3
as long as wide) or short (silicula, 5 mm long Pet free Pet united in the lower ½ Hypanthium shallow Hypanthium
campanulate Pet with basal ventral scales or sometimes fimbriate Pet without basal ventral scales, not fimbriate Pet free Pet united

9b 10a 10b 11a 11b 12a 12b 13a 13b 14a 14b 15a 15b 16a 16b 17a 17b 18a 18b 19a 19b 20a 20b 21a 21b 22a 22b 23a 23b 24a 24b 25a 25b 26a

26b 27a 27b

Gynostemma (no succulents) Doyerea

Key to genera with succulents (continued) 28a 28b 29a 29b

10 Apodanthera Corallocarpus 12 33 13 20 14 17 Halosicyos 15 Brandegea 16 Marah Ceratosanthes 18 19 Parasicyos Halosicyos Odosicyos Cucumis Cephalopentandra 21 Apodanthera 22 23 25 Cucumella 24 Momordica Citrullus 26 28

30a 30b 31a 31b 32a 32b 33a 33b 34a 34b 35a 35b 36a

36b 37a 37b 38a 38b 39a 39b 40a 40b 41a 41b 42a 42b

St with hairs St without hairs L lobes with secondary lobes L lobes without secondary lobes Pet hairy Pet glabrous Fr cylindrical, tapering at both ends Fr ovoid to oblong or subglobose Hypanthium campanulate Hypanthium cylindrical Sty 2 - 3 Sty united into a single column Fr with 1 Se Fr with > 1 Se Fr dry capsules Fr fleshy Pet with basal ventral scales or sometimes fimbriate Pet without basal ventral scales, not fimbriate Pet with basal ventral scales, not fringed Pet without basal ventral scales, fringed Hypanthium short and shallow Hypanthium campanulate or cylindrical Male Fl solitary Male Fl in panicles or racemes Stems with trichomes Stems lacking trichomes Male Fl in panicles Male Fl solitary or in racemes Pet united at the base Pet separate

Zehneria 29 30 31 Ibervillea Tumamoca Trochomeriopsis 32 Kedrostis Trochomeria 34 37 Cyclantheropsis 35 Gerrardanthus 36 37

38 Momordica Telfairia Coccinia 39 40 41 Cucurbita Citrullus Parasicyos 42 Kedrostis Marah

Momordica

27

References

Eureiandra Coccinia (continued)

Ali, M. A. & Al-Hemaid, F. M. A. (2011) Taxonomic significance of trichomes micromorphology in cucurbits. Saudi J. Biol. Sci. 18(1): 87–92, ills. https://doi. org/10.1016/j.sjbs.2010.10.003.

170 Bates, D. M. [& al. 1990], Robinson, R. W. & Jeffrey, C. (1990) Biology and utilization of the Cucurbitaceae. Ithaca (US) / London (GB): Comstock Publishing Associates / Cornell University Press. http://www. jstor.org/stable/10.7591/j.ctvr7f7q4 Bobich, E. G. & North, G. B. (2009) Structural implications of succulence: Architecture, anatomy, and mechanics of photosynthetic stem succulents, pachycauls, and leaf succulents. In: Barrera, E. de la & Smith, W. K. (eds.): Perspectives in biophysical plant ecophysiology. A tribute to Park S. Nobel; pp. 3–37. México (MX): Universidad Nacional Autónoma de México. http://www.researchgate.net/publication/ 261065161 Condon, M. A. & Gilbert, L. E. (1988) Sex expression of Gurania and Psiguria (Cucurbitaceae): Neotropical vines that change sex. Amer. J. Bot. 75(6): 875–884. http://www.jstor.org/stable/2444007 Condon, M. A. & Gilbert, L. E. (1990) Reproductive biology and natural history of the neotropical vines Gurania and Psiguria. In: Bates, D. M. & al. (eds.): Biology and utilization of the Cucurbitaceae; pp. 150–166. Ithaca (US) / London (GB): Comstock Publishing Associates / Cornell University Press. http:// www.jstor.org/stable/10.7591/j.ctvr7f7q4.17 Crowley, B. E. & Godfrey, L. R. (2013) Why all those spines? Anachronistic defences in the Didiereoideae against now extinct lemurs. South Afr. J. Sci. 109(1–2): 1346, 7 pp., map. https://doi.org/10.1590/ sajs.2013/1346. Crowley, B. E. [& al. 2011], Godfrey, L. R. & Irwin, M. T. (2011) A glance to the past: Subfossils, stable isotopes, seed dispersal, and lemur species loss in southern Madagascar. Amer. J. Primatol. 73(1): 25–37. https://doi. org/10.1002/ajp.20817. Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions  succulence as an adaptive strategy. Bradleya 27: 13–36, ills. https://doi.org/10.25223/brad. n27.2009.a10. Forest, F. [& al. 2007], Grenyer, R., Rouget, M., Davies, T. J., Cowling, R. M., Faith, D. P., Balmford, A., Manning, J. C., Proches, S., Bank, M. van der, Reeves, G., Hedderson, T. A. J. & Savolainen, V. (2007) Preserving the evolutionary potential of floras in biodiversity hotspots. Nature (London) 445(7129): 757–760, ill., maps. https://doi.org/10.1038/nature05587. Godfrey, L. R. [& al. 2016], Crowley, B. E., Muldoon, K. M., Kelley, E. A., King, S. J., Best, A. W. & Berthaume, M. A. (2016) What did Hadropithecus eat, and why should paleoanthropologists care? Amer. J. Primatol. 78(10): 1098–1112. https://doi.org/10. 1002/ajp.22506. Grumet, R. [& al. 2016], Katzir, N. & Garcia-Mas, J. (2016) Genetics and genomics of Cucurbitaceae. Cham (CH): Springer International Publishing. https:// doi.org/10.1007/978-3-319-49332-9. Holstein, N. (2015) Monograph of Coccinia (Cucurbitaceae). PhytoKeys 54: 1–166, key, maps, ills. https://doi.org/10.3897/phytokeys.54.3285. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa.

L. E. Newton and U. Eggli London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. (1978) Cucurbitaceae. In: Launert, E. (ed.): Flora Zambesiaca, 4: 414–499, ills., keys. London (GB): Flora Zambesiaca Managing Committee. Jeffrey, C. (1980) A review of the Cucurbitaceae. Bot. J. Linn. Soc. 81(3): 233–247. https://doi.org/10.1111/ j.1095-8339.1980.tb01676.x. Jeffrey, C. (1990) An outline classification of the Cucurbitaceae. In: Bates, D. M. & al. (eds.): Biology and utilization of the Cucurbitaceae; pp. 449–463. Ithaca (US) / London (GB): Comstock Publishing Associates / Cornell University Press. http://www.jstor.org/stable/ 10.7591/j.ctvr7f7q4.5 Jeffrey, C. (1996) Cucurbitaceae. In: Edwards, S. & al. (eds.): Flora of Ethiopia and Eritrea, 2(2): 17–59, ills., keys. Addis Ababa (ET): National Herbarium / Uppsala (SE): Department of Systematic Botany. Jeffrey, C. (2005) A new system of Cucurbitaceae. Bot. Zhurn. (Moscow & Leningrad) 90(3): 332–335. http:// en.arch.botjournal.ru/?t¼issues&id¼20050303 Jeffrey, C. & Thulin, M. (1993) Cucurbitaceae. In: Thulin, M. (ed.): Flora of Somalia, 1: 216–240, ills., keys. Kew (GB): Royal Botanic Gardens. Kartusch, B. & Kartusch, R. (2008) Stem anatomy of Acanthosicyos horridus (Cucurbitaceae). South Afr. J. Bot. 74(4): 647-650. https://doi.org/10.1016/j.sajb. 2008.04.001. Keraudren, M. (1966) Cucurbitacées. 185e famille. In: Humbert, H. (ed.): Flore de Madagascar et des Comores. Paris (FR): Muséum National d'Histoire Naturelle, Laboratoire de Phanérogamie. http://www. biodiversitylibrary.org/page/9186087 Keraudren, M. (1968) Recherche sur les Cucurbitacées de Madagascar. Mém. Mus. Nation. Hist. Nat., Sér. B, Bot. 16(2): 127–330, 13 ills. Kocyan, A. [& al. 2007], Zhang, L.-B., Schaefer, H. & Renner, S. S. (2007) A multi-locus chloroplast phylogeny for the Cucurbitaceae and its implications for character evolution and classification. Molec. Phylogen. Evol. 44(2): 553–577. https://doi.org/10. 1016/j.ympev.2006.12.022. Kubitzki, K. (2011) Introduction to Cucurbitales. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 4–6. Berlin / Heidelberg (DE) etc.: Springer Verlag. https://doi.org/10.1007/ 978-3-642-14397-7_2. Luca, P. de [& al. 1977], Alfani, A. & Virzo de Santo, A. (1977) CAM, transpiration, and adaptive mechanisms to xeric environments in the succulent Cucurbitaceae. Bot. Gaz. 138(4): 474–478, ills. http:// www.jstor.org/stable/2473883 MacDougal, D. T. (1912) The water-balance of succulent plants. Ann. Bot. (Oxford) 26(101): 71–93, tt. 6-10. http://www.jstor.org/stable/43236763 MacDougal, D. T. & Spalding, E. S. (1910) The waterbalance of succulent plants. Washington D.C. (US): Carnegie Institution. http://www.biodiversitylibrary. org/page/38518997 Matthews, M. L. & Endress, P. K. (2004) Comparative floral structure and systematics in Cucurbitales

Cucurbitaceae (Corynocarpaceae, Coriariaceae, Tetramelaceae, Datiscaceae, Begoniaceae, Cucurbitaceae, Anisophyllaceae). Bot. J. Linn. Soc. 145(2): 129–185, ills. https://doi.org/10.1111/j.1095-8339.2003.00281.x. McCloud, E. S. [& al. 1995], Tallamy, D. W. & Halaweish, E. T. (1995) Squash beetle trenching behavior: Avoidance of cucurbitacin induction or mucilaginous plant sap? Ecol. Entomol. 20(1): 51–59. https://doi.org/10. 1111/j.1365-2311.1995.tb00428.x. McVaugh, R. (2001) Flora Novo-Galiciana. A descriptive account of the vascular plants of Western Mexico. Volume 3: Ochnaceae to Loasaceae. Ann Arbor (US): University of Michigan Herbarium. Meeuse, A. D. J. (1962) The Cucurbitaceae of southern Africa. Bothalia 8(1): 1–111, keys. https://doi.org/10. 4102/abc.v8i1.1611. Mooney, H. A. [& al. 1992], Chu, C., Bullock, S. H. & Robichaux, R. (1992) Carbohydrate, water and nitrogen storage in vines of a tropical deciduous forest. Biotropica 24(2a): 134–139. https://doi.org/10.2307/2388666. Nee, M. [& al. 2009], Schaefer, H. & Renner, S. S. (2009) The relationship between Anisosperma and Fevillea (Cucurbitaceae), and a new species of Fevillea from Bolivia. Syst. Bot. 34(4): 704–708, ills., key. https:// doi.org/10.1600/036364409790139718. Nesom, G. L. (2015) Cucurbitaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 3–58, ills., keys. New York (US) / Oxford (GB): Oxford University Press. Njoroge, G. N. & Newton, L. E. (2002) Ethnobotany and distribution of wild genetic resources of the family Cucurbitaceae in the central highlands of Kenya. Pl. Genet. Resources Newslett. 132: 10–16. Olson, M. E. (2003) Stem and leaf anatomy of the arborescent Cucurbitaceae Dendrosicyos socotrana [sic!] with comments on the evolution of pachycauls from lianas. Pl. Syst. Evol. 239(3–4): 199–214, ills. https://doi.org/ 10.1007/s00606-003-0006-1. Rayder, L. & Ting, I. P. (1983) Shifts in the carbon metabolism of Xerosicyos danguyi H. Humb. (Cucurbitaceae) brought about by water stress. Pl. Physiol. (Lancaster) 72(3): 606–610. https://doi.org/10.1104/pp.72.3.606. Renner, S. S. & Pandey, A. K. (2013) The Cucurbitaceae of India: Accepted names, synonyms, geographic distribution, and information on images and DNA sequences. PhytoKeys 20: 53-118. https://doi.org/10. 3897/phytokeys.20.3948. Renner, S. S. & Schaefer, H. (2016) Phylogeny and evolution of the Cucurbitaceae. In: Grumet, R. & al. (eds.): Genetics and genomics of Cucurbitaceae; pp. 13–23. Cham (CH): Springer International Publishing. https:// doi.org/10.1007/7397_2016_14. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector's miscellany. Mill Valley (US): Strawberry Press. Sayed, O. H. (2001) Crassulacean Acid Metabolism 19752000, a check list. Photosynthetica 39(3): 339-352. https://doi.org/10.1023/A:1020292623960.

171 Schaefer, H. & Renner, S. S. (2010) A three-genome phylogeny of Momordica (Cucurbitaceae) suggests seven returns from dioecy to monoecy and recent longdistance dispersal to Asia. Molec. Phylogen. Evol. 54(2): 553-560. https://doi.org/10.1016/j.ympev.2009. 08.006. Schaefer, H. & Renner, S. S. (2011a) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi. org/10.1002/tax.601011. Schaefer, H. & Renner, S. S. (2011b) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Schaefer, H. [& al. 2009], Heibl, C. & Renner, S. S. (2009) Gourds afloat: A dated phylogeny reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous oversea dispersal events. Proc. Roy. Soc., Biol. Sci., Ser. B, 276(1658): 843–851, maps. https://doi. org/10.1098/rspb.2008.1447. Sun, M. [& al. 2016], Naaem, R., Su, J.-X., Cao, Z.-Y., Burleigh, J. G., Soltis, P., Soltis, D. E. & Chen, Z.-D. (2016) Phylogeny of the Rosidae: A dense taxon sampling analysis. J. Syst. Evol. 54(4): 363–391. https:// doi.org/10.1111/jse.12211. Wang, J. [& al. 2018], Sun, P., Li, X., Liu, Y., Yang, N., Yu, J., Ma, X., Sun, S., Xia, R., Liu, X., Ge, S., Liu, Y., Kong, Y., Cui, X., Lei, T., Wang, L., Wang, Z., Ge, W., Zhang, L., Song, X., Yuan, M., Guo, D., Jin, D., Chen, W., Pan, Y., Liu, T., Yang, G., Xiao, Y., Sun, J., Zhang, C., Li, Z., Xu, H., Duan, X., Shen, S., Zhang, Z., Huang, S. & Wang, X. (2018) An overlooked paleotetraploidization in Cucurbitaceae. Molec. Phylogen. Evol. 35(1): 16–26. https://doi.org/10. 1093/molbev/msx242. Wikström, N. [& al. 2001], Savolainen, V. & Chase, M. W. (2001) Evolution of the Angiosperms: Calibrating the family tree. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 268(1482): 2211–2220. https://doi.org/10.1098/rspb. 2001.1782. Wilkins, M. H. (1994) Good gourds and succulent squash: A cucurbit review. Cact. Succ. J. (US) 66(2): 65–71, ills. Winter, K. (1979) (delta)13C values of some succulent plants from Madagascar. Oecologia 40(1): 103–112, ills. https://doi.org/10.1007/BF00388814. Zhang, L.-B. [& al. 2006], Simmons, M. P., Kocyan, A. & Renner, S. S. (2006) Phylogeny of the Cucurbitales based on DNA sequences of nine loci from three genomes: Implications for morphological and sexual system evolution. Molec. Phylogen. Evol. 39(2): 305–322. https://doi.org/10.1016/j.ympev.2005. 10.002. Zimmermann, A. (1922) Die Cucurbitaceen. Beiträge zur Anatomie, Physiologie, Morphologie, Biologie, Pathologie und Systematik. Heft 1 / Heft 2. Jena (DE): Verlag von Gustav Fischer.

Ampelosicyos CUCURBITACEAE L. E. Newton

Ampelosicyos Thouars (Hist. Veg. Isles Afr. Austr., 68, t. 22, 1808). Type: Ampelosicyos scandens Thouars. Joliffieae Lit: Keraudren (1966: 64–66, 91–98, Flora Madagascar). Distr: Madagascar. Etym: Gr. ‘ampelos’, grapevine, vine; and Gr. ‘sicyos’, cucumber; for the climbing habit and the placement in this family. Incl. Delognaea Cogniaux (1884). Type: Delognaea humblotii Cogniaux. Incl. Tricyclandra Keraudren (1965). Type: Tricyclandra leandrii Keraudren. Incl. Odosicyos Keraudren (1981). Type: Odosicyos bosseri Keraudren. Perennial climbing shrubs, monoecious or dioecious; stem base and/or Rstock tuberous, to 30–120 cm ∅, often partly exposed, and extending into long thick taproots; tendrils simple; stems cylindrical or ribbed, woody, young shoots with reddish hairs; tendrils axillary, simple or bifid; L simple, petiolate, digitate or pedate, 3- to 5-lobed, leaflets entire or lobed, lower surface with reddish hairs; Inf with long peduncle; Fl unisexual, medium-sized to large, in pedunculate racemes (male) or solitary (male and female), diurnal (Ampelosicyos s.s.) or nocturnal (former Odosicyos and Tricyclandra);

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

hypanthium tube to 20 cm, narrowly funnel-shaped, only 2 mm ∅ near the base expanding to 12 mm ∅ and more above; Sep small, to 2 mm; Pet to 3 cm, white to yellowish, entire (Ampelosiycos s.s. fide Keraudren) or margins with numerous long fringes to 2 cm (former Odosicyos and Tricyclandra); St 5 (Ampelosicyos s.s.) or 3, inserted in the upper ½ or near the mouth of the tube; Fil very short but distinct; Anth all 1-thecous with triplicate to circular thecae; Ov with 3 placentae and many horizontal ovules; Sti 3, 2-lobed; Fr  pear-shaped to ellipsoid, 3  2 cm or 7–15  5–8 cm, fleshy, sometimes rostrate, indehiscent, yellow with whitish pulp, smooth; Se 5–6 or 15–25, obovate to bean-shaped, compressed,  asymmetrical, to 3  2 cm ∅. A small genus of originally 3 species, but after the inclusion of the monotypic genera Odosicyos and Tricyclandra by Schaefer & Renner (2011) it now has 5 species, all confined to Madagascar. The species of the genera now subsumed here possess succulent tubers or rootstocks, and the 2 taxa that are better known are covered in more detail below. For A. scandens Thouars (from NE and E humid forests), Keraudren (1966: 96) notes that the stem base is inflated and developped into an ovoid tuber 30  20 cm large. For A. meridionalis Keraudren (from S xeric shrubby forests) and A. humblotii (Cogniaux) Jumelle & Perrier (from E humid forests), KeraudrenAymonin (1971) notes the presence of long thickened roots from which they can regenerate. The genus name was wrongly spelled “Ampelosycios” in the protologue, whose

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Fig. 1 Ampelosicyos bosseri (Madagascar; Prov. Toliara). (Copyright: H. Schaefer)

publication year is 1808 according to TL2, but is variously also given as 1806 or 1807. A. bosseri (Keraudren) H. Schaefer & S. S. Renner (Taxon 60(1): 133, 2011). Type: Madagascar, Toliara (Keraudren 1312 [P]). Lit: Keraudren-Aymonin (1982: with ills.); Rauh (1998: 198–201, with ills.); both as Odosicyos. Distr: SW Madagascar (Toliara: Sakaraha Zombitsy); primary forest. Fig. 1.  Odosicyos bosseri Keraudren (1981). Perennial dioecious climbers to 4 m with large flattened discoid caudex up to 1.2 m ∅, barely subterranean; stems described as twining in the

protologue, but this appears unlikely, 5-ribbed, hairy, probably annual; tendrils simple; L 5–15 cm petiolate, lamina 3- to 5-lobed, to 15  10 cm overall, base cordate-truncate, margins with angled-triangular teeth, lower face with long hairs; Inf elongate racemes, severalflowered; male Fl long-pedicellate, nocturnal, pleasantly fragrant; Pet spreading,  10  5 mm, white to cream with darker veins, the somewhat truncate apical part with numerous long, often curled threads; St 3; female Fl not described; Fr obovate with subacute tip, 12–15  5–8 cm, indehiscent, fleshy, smooth; Se many.

Ampelosicyos CUCURBITACEAE

First placed in its own monotypic genus Odosicyos, but Schaefer & Renner (2011) included it in an enlarged genus Ampelosicyos. A. leandrii (Keraudren) H. Schaefer & S. S. Renner (Taxon 60(1): 133, 2011). Type: Madagascar, Mahajanga (Perrier 1016 [P]). Lit: Keraudren (1966: 64–66, with ills., as Tricyclandra). Distr: SW Madagascar (Toliara); deciduous dry forests, sandy soils or limestone outcrops.  Tricyclandra leandrii Keraudren (1965). Perennial climbers, possibly dioecious, with a globose underground caudex to 2–3 kg; stems glabrescent, with scattered hairs when young, finely streaked longitudinally, a little thickened at the nodes; tendrils simple; L petiole 1.5–2 cm, lamina palmate with 5 lobes, base cordate, 3 median lobes oblong-lanceolate, 3–5  1 cm, the 2 lateral lobes 2- or 3-lobed, touching the leaf base forming a heart-shaped sinus, glabrescent, a few hairs only on the veins, esp. when leaves are young, hairs deciduous, brownish on the upper face,  greenish on the lower face; male Inf racemes to 10 cm, finely tomentose, 5 - to 20-flowered; male Fl Ped slender, 2–3 cm, pubescent; Cl infundibuliform, 4  3 mm ∅ at the throat, finely pubescent; Sep triangular, 1  1 mm, finely pubescent;

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Pet imbricate in bud, fimbriate with numerous narrow curly divisions, basally with 3 large conspicuous veins, veins finely dividing at the tips; female Inf and Fl unknown; Fr indehiscent, stipitate on a thick pedicel to 2 cm, subellipsoid, 3  2 cm, apex obtuse; Se 5–6, leathery with smooth testa.

References Keraudren, M. (1966) Cucurbitacées. 185e famille. In: Humbert, H. (ed.): Flore de Madagascar et des Comores. Paris (FR): Muséum National d’Histoire Naturelle, Laboratoire de Phanérogamie. http://www. biodiversitylibrary.org/page/9186087 Keraudren-Aymonin, M. (1971) La survivance des Ampelosicyos (Cucurbitacées) à Madagascar. Bull. Soc. Bot. France 118(3–4): 281–285. https://doi.org/ 10.1080/00378941.1971.10838899. Keraudren-Aymonin, M. (1982) Odosicyos bosseri, Cucurbitacée énigmatique du Sud-ouest de Madagascar. Bull. Soc. Bot. France 129(2): 149–154, ills. https://doi.org/10.1080/01811797.1982.10824538. Rauh, W. (1998) Succulent and xerophytic plants of Madagascar. Vol. 2. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi.org/10. 1002/tax.601011.

Anisosperma CUCURBITACEAE L. E. Newton and U. Eggli

Anisosperma Silva Manso (Enum. Subst. Braz., 38, 1836). Type: Anisosperma passiflora Silva Manso. – Triceratieae – Lit: Nee & al. (2009: phylogeny). Distr: Brazil. Etym: Gr. ‘a-, an-’, without; Gr. ‘isos’, the same; and Gr. ‘sperma’, seed; for the very unequal seed number per fruit. Dioecous perennial climbers to several m long, with a basal caudex to 5 cm ∅; stems becoming woody with age; tendrils stout, long, apically bifid; L petiole 1–3 cm, with 2 small auriculate glands at the junction with the lamina, lamina entire, simple, 5–13  4.5–7.5 cm, narrowly ovate, membranous to slightly thickish, acuminate, both faces minutely pustulate to sparsely glandular-pubescent; male Fl in axillary fascicles or subumbelliform Inf; Ped (2–) 4–5 mm; hypanthium 4–5 (–13) mm, saucer- to cup-shaped; Sep  2.5 mm, lanceolate, fused to the petals above; Pet 5, 5–7 mm, oblong-hastate, greenishwhite to cream-coloured,  4 mm, with a median adaxial glandular ridge, the lower margin united with the sepals; St 5, in the flower centre; Fil free, 1.5–2 mm; Anth all 2-thecous, thecae straight, oriented vertically; female Fl solitary or in pairs,

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] U. Eggli Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

similar to the male flowers; Ov 6–9  4–5 mm, glabrous; Sty thickened in the lower part, upper part dilated and broadly obcordate; Fr ovoid to oblong, 8–15  5–11 cm, obscurely trigonous, fleshy, shortly apiculate, longitudinally dehiscent with several splits, smooth or somewhat irregularly verrucose; Se  15–20 per fruit, subcircular, 3.5–5.5  3–4 cm,  1.5 cm thick, yellowish-brown, striateverrucose, margin broad and wing-like. This monotypic genus was formerly included in the synonymy of Fevillea (8 species, no succulents), but the phylogeny presented by Nee & al. (2009) shows Anisosperma in a basal polytomy with all other genera of tribe Triceratieae (as Fevilleeae). The isolated position of the single species of Anisosperma within Fevillea was already recognized by Robinson & Wunderlin (2005), who placed it in a subgenus of its own. It differs from Fevillea by congested male inflorescences (vs. open panicles in Fevillea), oblonghastate petals in male flowers (vs. suborbicular), and ovoid to oblong shortly apiculate fruits (vs. subglobose with rounded apex). The seeds were traditionally used for their laxative properties. They are very rich in oil. A. passiflora (Vellozo) Silva Manso (Enum. Subst. Braz., 38, 1836). Type: Brazil, Rio de Janeiro (Vellozo s.n. [[lecto – icono]: Fl. Flumin. 10: t. 104, 1829]). – Lit: Cogniaux (1878: 120–122, t. 38); Robinson & Wunderlin (2005: 1993–1995, with ills., as Fevillea). Distr: E coastal Brazil (SE Bahia, Rio de Janeiro, E São

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_28

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Fig. 1 Anisosperma passiflora (Cultivated, São Paulo, Brazil). (Copyright: Pedro Acevedo, Smithsonian Institution)

Paulo, E Paraná, E Santa Catarina); gallery forests near sea level. – Fig. 1.  Fevillea passiflora Vellozo (1829). Description as for the genus.

References Cogniaux, A. (1878) Cucurbitaceae. In: Martius, C. (ed.): Flora Brasiliensis; 6(4): 1–126, tt. 1–38.

Leipzig (DE): F. Fleischer. https://doi.org/10.3931/ e-rara-50616. Nee, M. [& al. 2009], Schaefer, H. & Renner, S. S. (2009) The relationship between Anisosperma and Fevillea (Cucurbitaceae), and a new species of Fevillea from Bolivia. Syst. Bot. 34(4): 704–708, ills., key. https://doi.org/10.1600/0363644097 90139718. Robinson, G. L. & Wunderlin, R. P. (2005) Revision of Fevillea (Cucurbitaceae: Zanonieae). Sida 21(4): 1971–1996, key, ills. http://biostor.org/reference/ 159699

Apodanthera CUCURBITACEAE L. E. Newton and G. N. Njoroge

Apodanthera Arnott (J. Bot. (Hooker) 3: 274, 1841). Type: Apodanthera mathewsii Arnott. – Coniandreae – Lit: Jeffrey (1992: synopsis Brazil); Belgrano & Pozner (2017: synopsis with key). Distr: SW USA, Mexico, South America (Ecuador, Peru, Bolivia, Brazil, Uruguay, Argentina). Etym: Gr. ‘apodos’, without a foot; and Gr. ‘antheros’, flowering; for the sessile flowers. Incl. Guraniopsis Cogniaux (1908). Type: Guraniopsis longipedicellata Cogniaux. Climbers, sometimes with tuberous R, dioecious or monoecious; stems sometimes succulent, pubescent; tendrils simple or absent; L petiolate, simple or compound, pubescent; Fl glabrous or pubescent, hypanthium cylindrical or obconical; Cal lobes spreading in bud; Pet free or almost so, yellow; male Fl in fascicles or racemes; St 3, free; female Fl solitary or in clusters; Sti lobes various but not fimbriate; Fr ellipsoid or globose, 4–7 cm, sometimes beaked, longitudinally ridged, smooth or nearly so; Se many.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] G. N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya e-mail: [email protected]

A genus of 16 (Schaefer & Renner 2011) to 23 (Belgrano & Pozner 2017) species, of which only a few are  succulent and are treated below. As with many other cucurbits, underground plant parts are often not collected and remain undescribed, so other species might also have succulent caudices or taproots. Belgrano & Pozner (2017) present a synopsis including a key to a slightly recircumscribed Apodanthera (A. congesta and some other species are newly recognized as synomyms of Doyerea emetocathartica; see there) and recognize the three sections proposed by Cogniaux (1916), with largely allopatric distributions: [1] Sect. Apodanthera: L membranous, smooth; male Inf racemose. – 16 species from South America, no succulents. [2] Sect. Cucurbitopsis Cogniaux: L thickish, rough. – 3 species from USA and Mexico, 1 succulent. [3] Sect. Pseudoapodanthera Cogniaux: Inf densely glomerate or shortly spicate. – 3 or 4 species from NE Brazil, 3 succulent. A. hindii C. Jeffrey (Kew Bull. 47(3): 519–521, ills., 1992). Type: Brazil, Bahia (Harley & al. 26,688 [CEPEC, F, K, MBM, SPF]). – Lit: Paiva Lima (2010: with ills. pp. 17, 23, 88). Distr: NE Brazil (Bahia: Chapada Diamantina); sandstone soils and among sandstone rocks, 1000–1550 m.

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[3] Climbers, monoecious; stem to 5 m, succulent, glabrous, bright orange on new growth, becoming glaucous-green; L 1.3–6.5 cm petiolate, lamina simple, 2.3–7  3.7–9 cm, with 3–5 lobes almost to the base, puberulous-tomentose, more densely on the lower face; Fl thinly tomentose on the outside; male Fl numerous in short dense racemes, pubescent; Ped 2.5–5.5 mm; hypanthium cylindrical below, expanding above, 5.5–6.5 mm; Pet yellow, 2 mm; female Fl 2–5 in clusters coaxillary with male inflorescences; hypanthium cylindrical; Pet 2.5 mm, yellow; Fr ovoid, beaked, 1.2–2.3  0.5–1.05 cm, dark green to bright yellow-green becoming bright scarlet, with 8 Se. Very similar to A. villosa with 7–30 female flowers per inflorescence, and flowers that are more densely tomentose on the outside (Belgrano & Pozner 2017: 17). A. pedisecta Cogniaux (Mém. Cour. Acad. Roy. Sci. Belgique 28: 39, 1878). Type [lecto]: Brazil, Bahia (von Wied-Neuwied s.n. [BR]). – Lit: Jeffrey (1992: 521-522, with ill., protologue A. hatschbachii); Paiva Lima (2010: 75–76, with ill. p. 90). Distr: NE Brazil (Bahia, Minas Gerais); disturbed Caatinga vegetation in granite rock outcrop areas, 600–750 m. Incl. Anguria pedisecta Nees & Martius (1824) (nom. illeg., ICN Art. 52.1); incl. Apodanthera bradei Martinez Crovetto (1954); incl. Apodanthera hatschbachii C. Jeffrey (1992). [3] Climbers with tuberous Rstock, dioecious; stem puberulous, glabrescent, developing thin brownish bark when older; tendrils simple, thin, glabrescent; L 1.5–4 cm petiolate, lamina (3- to) 5-foliate, middle leaflet elliptic, obovate or oblanceolate, 2.2–9 (–10.5)  0.5–3.5 cm, lateral leaflets asymmetrical and 2- to 3-lobed, upper face of all leaflets glabrescent to glabrous, lower face densely puberulous; male Fl numerous in dense racemes, almost sessile; hypanthium cylindrical, 4.5–7 mm, almost glabrous; Pet oblong, 2.2–3.2 (–5) mm, yellow; female Fl numerous in clusters, sessile; hypanthium cylindrical, 2–5.5 mm, glabrous; Pet similar to male flowers; Fr ellipsoid, beaked, (7-) 11–15  5–6 cm, glabrous, scarlet.

L. E. Newton and G. N. Njoroge

Paiva Lima (2010: 75) also placed A. fasciculata Cogniaux (presence of a caudex unknown) in the synonymy here, but Belgrano & Pozner (2017: 16) disagree because of differences in inflorescence and flower architecture. A. undulata A. Gray (Pl. Wright. 2: 60, 1853). Type [lecto]: USA, Texas (Wright 1382 [GH 31797, BM, G, GH, K, MO, NY, P, PH, US]). – Distr: SW USA, Mexico (C Plateau). A. undulata var. australis McVaugh (in W. R. Anderson (ed.), Fl. Novo-Galic. 3: 493–496, ills., 2001). Type: Mexico, Jalisco (Dieterle 3550 [MICH]). – Distr: Mexico (C Plateau: Aguascalientes, Guanajuato, Hidalgo, Jalisco, Michoacán, Nayarit, Oaxaca, Querétaro, San Luis Potosí, Zacatecas); disturbed habitats, roadsides, borders of fields, 900–2200 m. [2] Differs from var. undulata: Tendrils 2-to 3-fid, usually curved around the stem at the node of origin; L lamina broadly orbicular-cordate, 7–10  8–12 cm; male Fl 1–5 in longpedunculate racemes. Belgrano & Pozner (2017: 12-13) include this in the synonymy of A. aspera Cogniaux, which is not known to have a caudex; Lira & Rodríguez Arévalo (1999: 8-10, ills.) describe the robust roots as woody, and the stems as herbaceoussucculent, and an occurrence limited to Puebla and Oaxaca. A. undulata var. undulata – Lit: Belgrano & Pozner (2017: 28-30, with ill.). Distr: SW USA (Arizona, New Mexico, Texas), N & C Mexico (Sonora, Chihuahua, Coahuila, Durango, S-wards to Zacatecas); flat plains and mesas, sometimes on limestone slopes, 450–1700 (–2100) m. I: Nesom (2015: 29) – Fig. 1. [2] Monoecious perennials; caudex large, thickened, to 1 m; stem trailing, thick, rough with appressed hairs; tendrils short, little coiled, mostly bifid; L petiole 4–7 cm, lamina reniform, 6–9  7–10 cm, shallowly lobed or with sinuatedentate margins, grey to green; male Fl few in umbels or corymbs; hypanthium 20–25 mm; Pet lobes 20–25 mm, pubescent; female Fl solitary;

Apodanthera CUCURBITACEAE

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Fig. 1 Apodanthera undulata var. undulata (USA; Arizona, Patagonia Lake Ranch Estates). (Copyright: Alan Schmierer/Wikimedia Commons)

Pet yellow; Fr ovoid,  4–5  4 cm, with longitudinal ribs, smooth.

References

A. villosa C. Jeffrey (Kew Bull. 47(3): 523–526, ills., 1992). Type: Brazil, Bahia (Harley & al. 22,753 [K, CEPEC]). – Lit: Paiva Lima (2010: 81, with ill. p. 95 & p. 92 (as A. succulenta)). Distr: Brazil (C Bahia: Serra do Tombador); scrub or low woodland on sand with sandstone outcrops (Campo Rupestre), 900–1100 m. I: Machado (2009). Incl. Apodanthera succulenta C. Jeffrey (1992). [3] Climbers, dioecious; stem to 5 m, stout, succulent, glabrous, shiny, often tinged bright orange-yellow towards the tip; tendrils simple, robust, glabrous; L 2.5–5.5 cm petiolate, lamina simple or compound, 3–6.5  4–9 cm, slightly succulent, puberulous (denser on the lower face); male Fl numerous in short dense fascicles; Ped 3–6 mm; hypanthium cylindrical, 6–10 mm, densely villous; Pet 2.5–3.5 mm, light yellow; female Fl numerous in dense fascicles; hypanthium cylindrical, 2.5–3.5 mm, densely villous, Pet 3 mm, light yellow; Fr ovoid, beaked, 1.3–1.7  0.5–0.7 cm, glabrous, scarlet; Se 2–3 per fruit, 5–5.5  2.5 mm, obovate. Similar to A. hindii (see there).

Belgrano, M. J. & Pozner, R. (2017) Sinopsis del género Apodanthera (Cucurbitaceae, Coniandreae). Darwiniana n.s. 5(1): 5–50, ills, key, maps. https://doi.org/10.14522/ darwiniana.2017.51.716. Cogniaux, A. (1916) Cucurbitaceae - Fevilleae et Melothrieae. In: Engler, A. (ed.): Das Pflanzenreich, IV. 275. I (Heft 66). Leipzig (DE): Verlag von Wilhelm Engelmann. http://bibdigital.rjb.csic.es/idurl/1/10009. Jeffrey, C. (1992) The genus Apodanthera (Cucurbitaceae) in Bahia State, Brazil. Kew Bull. 47(3): 517–528, key, ills. https://doi.org/10.2307/4110580. Lira, R. & Rodríguez Arévalo, I. (1999) Flora del Valle de Tehuacán-Cuicatlán. Fasc. 22. Cucurbitaceae A. L. Juss. México D.F. (MX): Instituto de Biologia, Universidad Nacional Autónoma de México. Machado, M. (2009) Apodanthera, caudiciform cucumbers of Bahía, Brazil. Cact. Succ. J. (US) 81(3): 147–149, ills. https://doi.org/10.2985/015.081.0309. Nesom, G. L. (2015) Cucurbitaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 3–58, ills., keys. New York (US)/Oxford (GB): Oxford University Press. Paiva Lima, L. F. (2010) Estudos taxonômicos e morfopolinicos em Cucurbitaceae brasileiras. Porto Alegre (BR): Universidade Federal do Rio Grande do Sul, Ph.D. thesis. http://www.lume.ufrgs.br/handle/ 10183/26294 Schaefer, H. & Renner, S. S. (2011) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi.org/10. 1002/tax.601011.

Brandegea CUCURBITACEAE L. E. Newton

Brandegea Cogniaux (Proc. Calif. Acad. Sci., ser. 2, 3: 58, 1890). Type: Elaterium bigelovii S. Watson. Sicyoeae Distr: SW USA, NW Mexico. Etym: For Townsend Stith Brandegee (1843–1925), US-American botanist, explorer, collector, civil engineer and topographer in California. Monoecious perennial vines with large thick R; stems climbing or trailing, slender, grooved, glabrous or almost so, sparsely scaly-pustulate; tendrils simple; L petiole 1–2 cm, lamina simple, hastate to quadrangular in outline, 1–6 cm, with 3–5 linear, triangular or ovate lobes, lateral lobes usually small, upper face pustulate-scabrous, lower face glabrous and punctate; male Fl in racemes, small, fragrant, hypanthium cup-shaped; Pet rotate, 5, lobes triangular-ovate, 2.1 mm, yellowish-white; St 3; Fil fused; female Fl solitary or 2, small, similar to male flowers; Fr obovoid, 0.4–1.2 cm, with long beak, indehiscent or irregularly dehiscent, smooth or sparsely echinate; Se 1 per fruit, 4–5 mm, subcylindricalclavate, muriculate. Some authors (e.g. Schaefer & Renner (2011b: 146) and Schaefer & Renner (2011a: 131–132)), have included this genus in Echinopepon Naudin ( 20 species from the Americas, none recorded

to be succulent). but this has not been followed by all. A single species only: B. bigelovii (S. Watson) Cogniaux (Proc. Calif. Acad. Sci., ser. 2, 3: 58, 1890). Type [syn]: USA, California (Palmer s.n. [GH, HUH]). Distr: SW USA (SE California, S Arizona), NW Mexico (Baja California, Baja California Sur, C Sonora, Jalisco?, Zacatecas?); sandy soils along stream beds, on hillsides and plains, to 900 m. I: Schlising (1993: 537); Nesom (2015: 19). Fig. 1.  Elaterium bigelovii S. Watson (1877)  Echinocystis bigelovii (S. Watson) Cogniaux (1881)  Echinopepon bigelovii (S. Watson) S. Watson (1889); incl. Echinocystis parviflora S. Watson (1882)  Brandegea parviflora (S. Watson) S. Watson ex Rose (1897); incl. Sicyos minimus S. Watson (1888)  Brandegea minima (S. Watson) Rose (1897); incl. Cyclanthera monosperma Brandegee (1889)  Brandegea monosperma (Brandegee) Cogniaux (1890); incl. Echinopepon palmeri S. Watson (1889)  Brandegea palmeri (S. Watson) Rose (1897)  Vaseyanthus palmeri (S. Watson) Gentry (1949)  Vaseyanthus insularis var. palmeri (S. Watson) Gentry (1950) (incorrect name, ICN Art. 11.4). Description as for the genus.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_30

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Fig. 1 Brandegea bigelovii (USA; California, Riverside County). (Copyright: Charles Kane)

References Nesom, G. L. (2015) Cucurbitaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 3–58, ills., keys. New York (US) / Oxford (GB): Oxford University Press. Schaefer, H. & Renner, S. S. (2011a) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi.org/10.1002/tax.601011.

Schaefer, H. & Renner, S. S. (2011b) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Schlising, R. L. (1993) Cucurbitaceae. In: Hickman, J. C. (ed.): The Jepson Manual; pp. 535–538, ills., key. Berkeley / Los Angeles (US), etc.: University of California Press. https://ucjeps.berkeley.edu/ eflora/

Cephalopentandra CUCURBITACEAE L. E. Newton

Cephalopentandra Chiovenda (Fl. Somala, 187, 1929). Type: Cephalopentandra obbiadensis Chiovenda. – Benincaseae – Lit: Jeffrey (1967: 84–85, Flora Trop. East Africa); Jeffrey & Thulin (1993: 233–234, with ills., Flora Somalia). Distr: NE Africa (Kenya, Uganda, Somalia, Ethiopia). Etym: Gr. ‘kephale’, head; Gr. ‘penta’, five; and Gr. ‘aner, andros’, male; for the cluster of stamens, erroneously stated to be 5 in the protologue. Perennials, dioecious, with tuberous exposed usually  semiglobose caudex to 20 cm, tuberculate, dark green; stems climbing or scandent, to 1 m, glabrous; tendrils simple (erroneously stated to be absent in the protologue); L subsessile, petiole 0.2–1.5 cm, lamina simple, elliptic-cordate, 3–9  4–7 cm,  pinnately 5- to 7-lobed, glaucous; Fl yellow or cream; male Fl solitary or paired; Ped 17–25 mm; hypanthium cylindrical below, expanded and campanulate above, 7–10 mm; Sep triangular-lanceolate, 2–3 mm; Pet 5, united in the lower ¼, 12–28  8–13 mm; St 3 (rather than 5 as suggested by the genus name), free; Fil distinct; Anth thecae triplicate; female Fl solitary; Ped 9–10 mm;

hypanthium cylindrical, 7 mm; Sep and Pet like those of male flowers; Fr ellipsoid, 8  4 cm ∅, red, smooth; Se many, compressed, pear-shaped, black, verrucose. A single species only, whose scientific genus and species names are both erroneous (as indicated in the description above) but cannot be changed (cf. ICN Art. 51.1): C. ecirrhosa (Cogniaux) C. Jeffrey (Kew Bull. 17(3): 477, 1964). Type: Somalia (Keller 116 [Z]). – Distr: Kenya, Uganda, Somalia, Ethiopia; deciduous Acacia woodland and bushland on sandy soils, 45–1090 m. I: Rowley (1987: 55); Lenain (2012: 84–85); Beyenbach (2018: 232); Dortort (2018: 246). – Fig. 1.  Coccinia ecirrhosa Cogniaux (1896); incl. Cephalopentandra obbiadensis Chiovenda (1929)  Coccinia obbiadensis (Chiovenda) Cufodontis (1965); incl. Coccinia quercifolia J. Hutchinson & E. A. Bruce (1941). Description as for the genus. A report about a monoecious plant (Grumbley 1994) has not been confirmed.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_31

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Fig. 1 Cephalopentandra ecirrhosa (without known wild origin, cult. Sukkulenten-Sammlung Zürich). (Copyright: U. Eggli)

References Beyenbach, J. (2018) Auf Pflanzensafari durch Kenia. Teil 3: Der Norden, von Isiolo nach Marsabit / Teil 4 und

L. E. Newton Schluss: Der Norden, Marsabit und Moyale. Avonia 36(3): 170–177, (4): 224–233, ills., map. Dortort, F. (2018) Commiphoras, Cucurbits, Adenias and more – Kenya 2017. Cact. Succ. J. (US) 90(4): 238–257, ills. https://doi.org/10.2985/015.090. 0403. Grumbley, T. (1994) Notes on propagating two rare succulents. Ballya 1(2): 35. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. & Thulin, M. (1993) Cucurbitaceae. In: Thulin, M. (ed.): Flora of Somalia, 1: 216–240, ills., keys. Kew (GB): Royal Botanic Gardens. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector's miscellany. Mill Valley (US): Strawberry Press.

Ceratosanthes CUCURBITACEAE L. E. Newton

Ceratosanthes Burman ex Adanson (Fam. Pl. 2: 139, 1763). Type: Ceratosanthes tuberosa J. F. Gmelin. – Coniandreae – Lit: Conceição (2013: monograph Brazil). Distr: Central America, Caribbean Islands, S America (to N Argentina). Etym: Gr. ‘keratos’, (forked) horn, antler; and Gr. ‘anthos’, flower; for the bifid petals. Herbaceous dioecious or rarely monoecious climbers from large tuberous Rstocks, tubers globose to napiform, 15–20 cm ∅; stem 2 cm ∅, surface rough; Br slender, angular-furrowed, glabrous; tendrils filiform, short, simple; L petiolate, ovate, pentagonal to reniform in outline, palmately 3- to 5-lobed from the slightly emarginate base; Fl small, nocturnal; male Fl several in longpedunculate racemes, with elongate cylindrically funnel-shaped tube; Sep lanceolate, to 2 mm; Pet white to whitish-green or cream-coloured, 10 mm, bifid for ½ from the tip, usually involute; St 3; Fil very short, free; Anth 3, 2 2-thecous, 1 1-thecous, thecae straight; female Fl solitary or in groups of 2–4, similar to male flowers; Fr juicy berries, ovoid-oblong, to 4  2 cm, green becoming orange to red at maturity, often white-spotted or -blotched; Se many, ovoid to subglobose, turgid, pale, smooth, margin distinct.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

A small genus of 5 species, apparently all with tuberous roots, and predominantly from semi-arid regions. C. hilariana Cogniaux (Diagn. Cucurb. 2: 31, 1877). Type [syn]: Brazil, Goiás (St. Hilaire 777bis [P]). – Distr: E to S Bazil (Goiás, Minas Gerais, Mato Grosso, São Paulo, Santa Catarina), Paraguay (Amambay, Concepción), NE Argentina (Formosa, Misiones); Cerrado and other xeric scrub formations, to 300 m. I: Conceição (2013: 6, 37). Incl. Ceratosanthes parviflora Cogniaux (1888). Monoecious climbing herbs; stem sparsely branched, cylindrical, glabrous or rarely pubescent; tendrils glabrous, delicate; L petiole 1–1.4 cm, glabrous, lamina deeply 3-lobed, or rather cordiform with 3 shallow lobes, 6.6–7.6  5.9–8 cm overall, puberulent to scabrous, base cordate, central lobe largest, obovate, margins crenate-dentate, apices acuminate to acute; male Inf 6- to 9-flowered; peduncle 7.3–10.6 cm, glabrous; male Fl  2 cm; Ped  0.5 cm, puberulent; hypanthium 1.2–1.4 cm long; Sep  1  1 mm, widely ovate to ovate, ciliate, acute; Pet  2  1.5 mm, oblong, connate at the base, externally puberulent; female Inf 3- to 13-flowered; peduncle 1.5–11.8 cm, glabrous; female Fl  1.7 cm; Ped  0.3 cm, puberulent, otherwise similar to male flowers; Ov 5–6  1 mm, narrowly ellipsoid, puberulent; Sty  5 mm; Sti bifid, plumose; Fr 1–2 per inflorescence, globose,  1 cm,

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orange-red; Se  8 per fruit, subglobose,  4 mm, brown. C. multiloba Cogniaux (Diagn. Cucurb. 2: 29, 1877). Type [syn]: Brazil (Pohl 520 [W]). – Distr: E & S Brazil (Bahia & Minas Gerais to Rio Grande do Sul), Paraguay, N Argentina (Misiones, Formosa, Tucumán), Uruguay, N Venezuela (Lara)?; semideciduous forests, xeric scrub and Monte vegetation, low elevations. – Figs. 1 and 2. Incl. Ceratosanthes hassleri Cogniaux (1903); incl. Ceratosanthes diazii Lillo (1916); incl. Ceratosanthes fiebrigii Cogniaux (1916).

Fig. 1 Ceratosanthes multiloba (Uruguay; Tacuarembó, Arroyo del Tala, Ruta 21). (Copyright: A. González)

Fig. 2 Ceratosanthes multiloba Asunción). (Copyright: U. Drechsel)

(Paraguay;

near

Monoecious climbing herbs; stem sparsely branched, cylindrical, glabrous; tendrils glabrous, delicate; L petiole 1.15–4.9 cm, glabrous, lamina 3- to 5-lobed, or simple, 6.3–10.3  5–11 cm overall, membranous, glabrous or puberulent to scabrous, margins entire or sparsely lobed, base straight or cordate, central lobe largest, oblong or (ob-) lanceolate, lateral lobes asymmetrical, apex acute, obtuse or rounded, frequently mucronate; male Inf 10- to 20-flowered; peduncle 6–22.7 cm, glabrous in the basal parts, puberulent above; male Fl 0.9–1.44 cm; Ped 0.3–0.6 cm, puberulent; hypanthium 1 cm; Sep  1.25–1.75  1.1–1.4 mm, ovate, ciliate, acute; Pet  4  2 mm, oblong, connate at the base, externally puberulent; female Inf 1- to 7-flowered; peduncle 0.6–1.2 cm, (sub-) glabrous; female Fl 0.6–1.2 cm; Ped 1.2–3.9 mm, puberulent, otherwise similar to male flowers; Ov 4–12  1–6 mm, (narrowly) ellipsoid, puberulent; Fr 1–2 per inflorescence, globose, 0.97–1.27  0.4–0.75 cm; Se  4 per fruit, globose,  4 mm, brown. C. palmata (Linné) Urban (Repert. Spec. Nov. Regni Veg. 15: 323, 1918). Type: [icono:] Burman, Pl. Amer. 1: t. 24, 1756. – Distr: Grenada, Trinidad & Tobago, Venezuela, French Guiana, E to S Brazil (Ceará and Pernambuco to Santa Catarina); xeric scrub including Caatinga and Cerrado vegetation, (semi-) deciduous forests, or disturbed lands, low elevations. I: Conceição (2013: 38).  Trichosanthes palmata Linné (1759); incl. Trichosanthes corniculata Lamarck (1783)  Ceratosanthes corniculata (Lamarck) Cogniaux (1881); incl. Ceratosanthes tuberosa J. F. Gmelin (1791); incl. Trichosanthes tuberosa Willdenow (1805); incl. Ceratosanthes gracilis Cogniaux (1877); incl. Ceratosanthes latiloba Cogniaux (1877); incl. Ceratosanthes latiloba var. acutiloba Cogniaux (1877); incl. Ceratosanthes trifoliata Cogniaux (1877); incl. Ceratosanthes angustiloba Ridley (1890); incl. Ceratosanthes cuneata Ridley (1890). Dioecious climbing herbs; stem sparsely branched, cylindrical, glabrous; tendrils glabrous, robust; L petiole 1.7–3.3 cm, glabrous, lamina 3-foliolate or deeply 3-lobed, 5.9–15 

Ceratosanthes CUCURBITACEAE

6.7–13.2 cm overall, membranous, glabrous, leaf base straight or cordate, central leaflet or lobe largest, lanceolate, lanceolate-ovate or oblanceolate, margins entire, sparsely dentate or crenate, apex acute, mucronulate, lateral leaflets or lobes asymmetrical, constricted or strongly 2-lobed; male Inf  10-flowered; peduncle to 17.5 cm, glabrous to puberulent; male Fl 2.2–2.5 cm; Ped 0.5–0.6 cm, glabrous; hypanthium 1.2–1.78 cm; Sep 1–2  1.5 mm, widely ovate, ciliate, acute; Pet  4  1.5–2 mm, oblong, connate at the base, externally puberulent; female Inf 2- to 4-flowered; peduncle 6.5 cm, glabrous to puberulent; female Fl 1.2–1.5 cm; Ped  0.5 cm, glabrous, otherwise similar to male flowers; Ov 3–4  3–5 mm, (narrowly) ellipsoid, puberulent; Fr globose to ellipsoid, 1.1–1.15  0.7 cm; Se 5–6 per fruit, globose, 4 mm, brownish. C. rupicola Ridley (J. Linn. Soc., Bot. 27: 38, 1890). Type: Brazil, Pernambuco (Ridley & al. 53 [K]). – Distr: NE Brazil (Pernambuco: Ilhas Fernando Noronha); scrambling over rocks. Dioecious climbing herbs; stem branched, cylindrical, glabrous; tendrils glabrous, robust; L petiole 1–3 cm, glabrous, lamina 3-foliolate, 5.5–7.5  4.5–13 cm overall, coriaceous, glabrous, central leaflet largest, obovate, lateral leaflets asymmetrical, oblong, all leaflets mucronate, margins entire or sparsely dentate; male Inf 5- to 18-flowered; peduncle 3.5–13.1 cm, glabrous to puberulent; male Fl 1.1–1.35 cm; Ped 0.15–0.3 cm, glabrous; hypanthium 2 cm; Sep 0.5–1  1.5 mm, widely ovate, ciliate, acute; Pet  4  2 mm, oblong, connate at the base, externally puberulent; female Inf 5- to 10-flowered; peduncle 6.5–10.7 cm, glabrous to puberulent; female Fl 1.3–1.55 cm; Ped 0.2–0.3 cm, glabrous, otherwise similar to male flowers; Ov  4  1 mm, narrowly ellipsoid, puberulent; Fr globose,

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0.5–1.1  0.5–0.8 cm; Se  4 per fruit, globose,  4  4 mm, brownish. C. tomentosa Cogniaux (Diagn. Cucurb. 2: 29, 1877). Type: Brazil, Minas Gerais (Warming s.n. [C]). – Distr: S-C Brazil (Mato Grosso, Goiás, Tocantins, Minas Gerais, São Paulo); ecology not described. I: Conceição (2013: 6, 39). Incl. Ceratosanthes tomentosa var. subnuda Cogniaux (1877); incl. Ceratosanthes warmingii Cogniaux (1877). Dioecious climbing herbs; stem branched, cylindrical, puberulent or pubescent; tendrils glabrous, robust; L petiole 0.5–3 cm, pubescent, puberulent when young or rarely strigulose when older, lamina palmately 3-lobed or -divided, or sometimes entire and sagittate, reniform or broadly ovate, 3.6–13.5  3.75–11.5 cm overall, pubescent or puberulent when juvenile, margins entire or sparsely dentate, apex acute, obtuse or rounded, mucronate, when lobed all lobes similar, (ob-) lanceolate, apex acute to rounded, mucronate; male Inf 4- to 12-flowered; peduncle 10.5–14 cm, puberulent or pubescent; male Fl 1.5–2 cm; Ped 3–5 mm; hypanthium 0.9–2.2 cm; Sep 1–1.5  1 mm, lanceolate, ciliate, acute; Pet 2–2.2  1–2 mm, oblong, connate at the base, externally puberulent; female Inf 3- to 12-flowered; peduncle 1–13 cm, puberulent or pubescent; female Fl 1–2 cm; Ped 2.5–3 mm, puberulent or pubescent, otherwise similar to male flowers; Ov 2.5–7.5  1–3 mm, narrowly ellipsoid, puberulent; Fr 2–6 per inflorescence, ellipsoid, glabrous, 1.5–2.6  1–1.5 cm; Se 2–6 per fruit, globose,  4 mm, brown, bordered.

References Conceição, T. C. (2013) The genus Ceratosanthes Adans. (Cucurbitaceae Juss.) in Brazil. Aveiro (PT): Universidade de Aveiro, Departamento de Biologia, M.Sc. thesis. http://hdl.handle.net/10773/11581

Citrullus CUCURBITACEAE L. E. Newton

Citrullus Schrader ex Ecklon & Zeyher (Enum. Pl. Afric. Austral., 279, 1836). Type: Cucurbita citrullus Linné [conserved type]. – Benincaseae – Distr: Africa, Asia. Etym: For the similarity of the fruits of some species to those of Citrus (“Orange”, “Lemon” etc.; Rubiaceae). Incl. Anguria Miller (1754) (nomen rejiciendum, ICN Art. 56.1). Type: not designated. Incl. Colocynthis Miller (1754) (nomen rejiciendum, ICN Art. 56.1). Type: not designated. Annual or tuberous-rooted perennial climbers or trailers, dioecious or monoecious, some with tuberous R to 1 m long; stems herbaceous, to 6 m; tendrils simple, 2- to 3-fid, or absent, or becoming spine-like (C. naudinianus); L petiolate, lamina simple, rounded or broadly to triangular-ovate, usually deeply
pinnately 3- to 5-lobed, segments lobulate or dissected; Fl solitary, rarely fasciculate, axillary,
2.5–5 cm, yellow; male Fl hypanthium campanulate, short; Cl rotate or broadly campanulate, medium-sized; Pet 5, united in the lower ½, yellow to white; St 3, free; Anth 2 2-thcous, 1 1-thecous; female Fl hypanthium short; staminodes present; Sti 3-lobed; Fr globose

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

or oblong, fleshy or dry, indehiscent, large, greenish, glabrous or covered with prominent spines; Se many, oblong, compressed, pale yellowish, blackish or brown, smooth, with or without distinct margin. – Cytology: x ¼ 11 (Darlington & Wylie 1955). 3 (perennial) of the 4–6 species of this genus are moderately succulent and described below. C. lanatus (Thunberg) Matsumura & Nakai (annual) is the “Water Melon” widely cultivated for its large edible fruits. C. colocynthis (Linné) Schrader (Linnaea 12: 414, 1838). Type: LINN 1152.2 [original material]. – Lit: Jeffrey (1967: 46, Flora Tropical East Africa). Distr: Tropical and subtropical NW to NE Africa, Mediterranean and to Afghanistan and Pakistan; semi-arid bushland and grassland, 0–770 (–1500) m.  Cucumis colocynthis Linné (1753); incl. Colocynthis vulgaris Schrader (1833) (incorrect name, ICN Art. 11.4); incl. Colocynthis dimidiens Gandoger (1886) (incorrect name, ICN Art. 11.4); incl. Colocynthis nubica Gandoger (1886) (incorrect name, ICN Art. 11.4). Perennial trailing herbs with woody tuberous Rstock; stems angled, shortly hairy when young, becoming scabrid; tendrils usually simple, sometimes bifid; L petiole 0.8–6 cm, lamina ovate in outline, 2.6–11  2.4–6.6 cm, palmate, with 3–5 ovate lobes, upper face smooth except on the scabrid-hairy veins, lower face scabrid-hairy; male Fl solitary; Ped 0.8–2 cm; hypanthium

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Fig. 1 Citrullus naudinianus (Namibia; Caprivi Strip, road through Bwabwata). (Copyright: B. Wursten/Online Flora of Zimbabwe)

obconical, 1.5–2.5 mm; Pet 8–9  5 mm, greenish-yellow, united below; female Fl solitary; Ped 1–4.5 cm; hypanthium short; Pet 6–10  4–6 mm; Fr subglobose, 5–12 cm ∅, smooth, green striped or mottled yellow, or uniformly yellow. This taxon has been grown since ancient times for its fruits, of which the dried pulp is used as a purgative. Vernacular names: “Colocynth”, “Bitter Apple”, “Vine of Sodom”. An anatomical study (Burrows & Shaik 2015) showed that the tuberous taproot consists mostly of secondary xylem parenchyma, with vessels of relatively small diameter and few fibres. C. ecirrhosus Cogniaux (Verh. Bot. Vereins Prov. Brandenburg 30: 151, 1888). Type: Namibia (Marloth 1192 [Z]). – Lit: Meeuse (1962: 58–59). Distr: Namibia (Omaruru, Swakopmund, Karibib, Lüderitz), NW RSA (Northern Cape); Namib desert areas, usually in sandy soils, 150–1065 m. Trailers with woody Rstock, monoecious; stems to 2 m, numerous, only little branched, puberulous to shortly hispid when young, glabrous later; tendrils absent; L petiole 0.5–2 cm, lamina ovate to suborbicular in outline, 2–6  2–6 cm, 3- to 5-lobed almost to the base, veins setose-hispid, otherwise usually glabrous; male Fl with 5–6 mm long hypanthium; Pet

7–10 mm, yellow; female Fl yellow; Fr globose or subglobose, 8–15 cm ∅, yellow. C. naudinianus (Sonder) Hooker fil. (in Oliver, Fl. Trop. Afr. 2: 549, 1871). – Distr: Angola, Namibia, RSA (Northern Cape, NorthWest Prov., Limpopo, Gauteng, Free State), Botswana, Moçambique, Zambia, Zimbabwe; deep sandy soils. – Fig. 1. Incl. Cucumis naudinianus Sonder (1862)  Colocynthis naudinianus (Sonder) Kuntze (1891) (incorrect name, ICN Art. 11.4)  Acanthosicyos naudinianus (Sonder) C. Jeffrey (1962). Perennial herbs with fusiform-cylindrical tuberous R to 1 m, to 8 cm ∅; stems prostrate, several from the crown of the root, to 3 cm ∅, often branched, angular-sulcate, shortly pubescent, glabrescent; tendrils simple, 3–5 (–7) cm, straight or slightly curved at the apex, rigid and becoming spine-like, glabrescent; L secund, petiole 0.5–4 (–7) cm, shortly setose-hispid, lamina deeply palmatifid, 3–8 (–14)  2–6 (–12) cm, scabrid above, setose below, or scabrid on both faces, lobes 5, all
irregularly and lyrately lobed, middle lobe longest, all lobes and secondary lobes entire to dentate or lobed; Fl solitary; hypanthium cup-shaped to shortly campanulate, 5–6  7–8 mm, shortly hispid; Sep linear-lanceolate, subacute or obtuse, 5–6  7–8 mm; Pet yellow,

Citrullus CUCURBITACEAE

ovate-elliptic, subacute, 15–20  8–10 mm, shortly hispid-pubescent; male Fl on Ped 0.5–2 cm long; Fil dorso-laterally flattened, 4–5 mm; female Fl on Ped 3–8 cm long; staminodes small, tongue-shaped or cylindricalsetaceous; Ov ellipsoid to ellipsoid-fusiform, 12–15 mm, covered with short conical spines; Fr ellipsoid, 6–12  4–8 cm, covered with large blunt conical spines, greenish-yellow or light greyish-green; Se 7.5–10  4–6 mm, smooth, white. The tuberous root is an important food and water resource in the Kalahari. In Botswana the fruits are eaten, raw or roasted.

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References Burrows, G. E. & Shaik, R. S. (2015) Comparative developmental anatomy of the taproot of the cucurbitaceous vines Citrullus colocynthis (perennial), Citrullus lanatus (annual) and Cucumis myriocarpus (annual). Austral. J. Bot. 62(7): 537–545, ills. https://doi.org/10. 1071/BT14124. Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Meeuse, A. D. J. (1962) The Cucurbitaceae of southern Africa. Bothalia 8(1): 1–111, keys. https://doi.org/10. 4102/abc.v8i1.1611.

Coccinia CUCURBITACEAE L. E. Newton

Coccinia Wight & Arnott (Prodr. Fl. Ind. Orient. 1: 347, 1834). Type: Coccinia indica Wight & Arnott [nom. illeg.,
Bryonia grandis Linné, cf. Holstein, PhytoKeys 54: 55, 2015].  Benincaseae  Lit: Holstein & Renner (2011: dated phylogeny); Holstein (2015: monograph). Distr: Subsaharan Africa, 1 species widespread throughout the Old World tropics and introduced into the New World, the rest in tropical Africa and S Africa. Etym: From Lat. ‘coccineus’, deep red; for the red fruits. Incl. Cephalandra Schrader ex Ecklon & Zeyher (1836). Type: Bryonia quinqueloba Thunberg. Incl. Physedra Bentham & Hooker fil. (1867). Type: Physedra heterophylla Hooker fil. [indirect lectotypifcation by Jeffrey 1962: 361 according to Holstein, PhytoKeys 54: 55, 2015]. Incl. Staphylosyce Bentham & Hooker fil. (1867). Type: Staphylosyce barteri Hooker fil. Climbers or trailers, sometimes with tuberous Rstock or R tubers, dioecious; tendrils simple or bifid near the tip; L simple, lobed or unlobed, usually very variable in shape; Fl medium to large, yellow or dull orange or brownish, rarely

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

whitish; male Fl solitary, clustered or usually in racemose Inf, to 8 cm ∅; hypanthium short and broad; Sep 5, usually small to very small; Pet 5, 1.4–5.5 cm, united forming a short tube; St usually 3; Fil usually variously united, less often free; Anth 3, all 2-thecous or rarely 2 2-thecous and 1 1-thecous, together forming a central head, thecae triplicate; female Fl solitary or rarely in racemose Inf, similar to male flowers; Sti 3-lobed; Fr globose to cylindrical, small or moderately large, baccate, indehiscent, orange to red or red mottled with green; Se many; ovate to pear-shaped, compressed, testa fibrillose, margin evident.  Cytology: x ¼ 12 (Darlington & Wylie 1955); or 2n ¼ 22 þ XY sex chromosomes in C. grandis (Linné) Voigt cited from Schaefer & Renner (2011: 164). Mostly diploid, but some triploidy reported in cultivars. Only a handful of the 25 species (Holstein 2015) of the genus are covered below, but others may have tubers as well. C. grandis has edible shoots and fruits, and C. abyssinica (Lamarck) Cogniaux is cultivated in Ethiopia for the edible tubers. Holstein & Renner (2011) and Holstein (2015) found 4 clades in the genus based on molecular data, and some indications for on-going hybridization. The genus started to diversify 6.9 mybp, and several shifts between habitat types (semiarid, woodland, forest) were found, and the occurrence in semi-arid habitats was reconstructed as ancestral condition.

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Fig. 1 Coccinia adoensis var. adoensis (Tanzania; Rukwa, Sumbawanga). (Copyright: R. von Blittersdorff/African Plants  A Photo Guide. www. africanplants. senckenberg.de)

The tubers of most species are derived from the hypocotyl, while C. adoensis and C. grandiflora Cogniaux ex Engler and probably also C. senensis Cogniaux possess true globose subterranean root tubers (Holstein 2015: 6–7). The tubers might be an adaptation to fire, and become lignified with time and appear to store primarily starch, but presence or absence of tubers remains unknown for many forest species, esp. from W and C Africa. The degree of succulence remains largely unknown. The complete monograph produced by Holstein (2015) also includes much data on use, economical potential and phytochemistry. The following name is of unresolved application but is referred to this genus: Coccinia aostae Buscalioni & Muschler (1913). C. adoensis (Hochstetter ex A. Richard) Cogniaux (in A. & C. de Candolle, Monogr. Phan. 3: 538, 1881). Type: Ethiopia, Tigray (Schimper 166 [P, BM, BR, G, G-DC, K, L, LG, M, P, TUB, W]).  Distr: Africa from Ghana to Ethiopia, S-wards to RSA.
Momordica adoensis Hochstetter ex A. Richard (1847). C. adoensis var. adoensis  Distr: Africa from Ghana to Ethiopia, S-wards to RSA; grassland and deciduous woodland, 500–2140 m. I: Keraudren (1967: 133); Jeffrey (1967: 37),

repeated by Jeffrey (1996: 53); Wilkins (1994: 67).  Fig. 1. Incl. Bryonia convolvuloides A. Richard (1847); incl. Bryonia jatrophifolia A. Richard (1847)
Coccinia jatrophifolia (A. Richard) Cogniaux (1881); incl. Cephalandra pubescens Sonder (1862)
Coccinia pubescens (Sonder) Eyles (1916); incl. Coccinia hartmanniana Schweinfurth (1868); incl. Coccinia rigida Cogniaux (1895); incl. Coccinia djurensis Schweinfurth & Gilg (1904); incl. Coccinia princeae Gilg (1904); incl. Coccinia parvifolia Cogniaux (1907); incl. Coccinia homblei Cogniaux (1916); incl. Coccinia subspicata Cogniaux (1916); incl. Coccinia roseiflora Suessenguth (1951). Climbing or trailing; R woody, also forming subglobose tubers distinct from the stem base; stems to 3 m with woody perennial base and annual aerial Br, herbaceous, ribbed, sparsely to densely pubescent; tendrils simple; L petiole 0.4–5 cm, lamina triangular-ovate or ovatecordate, 3.5–13.5  2.6–13.5 cm, unlobed or palmately 3- to 7-lobed, lobes triangular or ovate to obovate-oblong, elliptic-ovate, oblong-lanceolate or linear-lanceolate, glabrous or softly pubescent, sometimes scabrid, trichomes to 0.5 (0.8) mm; male Fl 2–22 in racemes on 0.5–7.5 cm peduncles; Ped 0.3–2.5 cm, from the same axils also solitary flowers on long pedicels; hypanthium obconical, 3.5–7 mm; Pet golden-orange to pale apricot-orange or salmon, rarely tinged purplish,

Coccinia CUCURBITACEAE

brown-veined outside, united in the lower 1=2 – 2=3, lobes 13–30 mm; St 3; female Fl solitary; hypanthium campanulate, 2–3 mm; Pet united in the lower 2=3 lobes 12–18 mm; Fr ovoid-ellipsoid, often shortly beaked, 3.6–8  1.4–3 cm, red. In Kenya, leaves are used as a vegetable and root tubers as a soap substitute (Njoroge & Newton 2002). C. adoensis var. aurantiaca (C. Jeffrey) Holstein (PhytoKeys 54: 67, 2015). Type: Tanzania, Dodoma (Polhill & Paulo 1221 [K, B, BR, EA, PRE]).  Distr: C Tanzania; dry miombo woodland, grass savanna among rocks on hills, 600–1200 m, in drier habitats than var. adoensis. I: Holstein (2015: 16).
Coccinia aurantiaca C. Jeffrey (1963). Differs from var. adoensis: Stems, L petioles and lamina, Ov and young Fr more densely covered with short trichomes to 0.5 (0.8) mm; Fr without beak; Se flat with hyaline girdle. C. adoensis var. jeffreyana Holstein (PhytoKeys 54: 70, 2015). Type: Tanzania, Iringa (Lovett 1597 [MO, EA]).  Distr: Kenya, Tanzania, Malawi; highland grassland, open woodland, 1300–2600 m. Differs from var. adoensis: Trichomes on L petiole and lamina longer, 0.8–1.2 mm; Cal lobes 1–3.5 mm. C. mackenii Naudin ex C. Huber (Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 8: 52, 1865). Type [lecto]: RSA, KwaZulu-Natal (Naudin s.n. [P, G-DC, K]).  Distr: RSA (Eastern Cape, KwaZulu-Natal, Mpumalanga, Limpopo), Swaziland; Afromontane forests, coastal forests, litoral forests, forest margins, sometimes grassland, poorly drained clay, sandstone or quartzite soils, 0–1750 (2010) m. I: Welman & Condy (2007: as C. palmata). Incl. Momordica palmata E. Meyer ex Drège (1843) (nom. inval., ICN Art. 38.1a); incl. Cephalandra palmata Sonder (1862)
Coccinia palmata (Sonder) Cogniaux (1881); incl. Cephalandra mackennii Naudin (1866); incl. Coccinia dinteri André (1900).

197

Perennial climbers or creepers, with large tuberous Rstock; stems to 9.5 m, glabrous; tendrils bifid or rarely simple; L petiole 0.7–11 cm, glabrous or with thin trichomes, lamina 3–13.5  3–15.5 cm, shallowly to profoundly 5-lobate, in the latter case often weakly lobulate, lobes triangular, lanceolate, or ovate to obovate, margins smooth, dentate, sometimes serrate to lobulate, esp. towards the tip, tip acute with final tooth, upper face glabrous with clear to white pustules, rarely with few trichomes, lower face glabrous or with thin, stiff or articulate trichomes, towards the base usually with glands; probracts to 4 mm, oblong-lanceolate; male Fl solitary or in few-flowered racemes; peduncle 5–6.5 cm; Ped in racemes to 2.5 cm, of solitary flowers 6–9 cm, glabrous, rarely with long trichomes; Cal lobes 1.5–6.5 mm, linear, subulate to narrowly triangular, first erect, later spreading to reflexed; Cl 1.3–2.7 cm ∅, cream to pale buff, glabrous; Pet triangular, 0.7–1.1 cm; St not described; female Fl solitary; Ped 0.7–5 cm, glabrous, otherwise similar to male flowers; Ov glabrous; Sty columnar; Sti bulging; Fr ellipsoid to oblong,  10  2–2.5 cm, unripe green with white mottling, ripe red-orange to red, sometimes with white mottling; Se 6–8  3–4.5  1.5 mm, flatly lenticular, slightly asymmetrical, finely rugulose, dirty white. The forest weaver (Ploceus bicolor Vieillot) was observed to feed on fruits of this species (Bleher & al. 2003). C. rehmannii Cogniaux (Bull. Herb. Boissier [sér. 1] 3: 418, 1895). Type [lecto]: RSA, Limpopo (Rehmann 5168 p.p. [Z, BR, K]).  Distr: Botswana, Namibia, Zimbabwe, Moçambique, RSA (Northern Cape, North-West Prov., Limpopo, Mpumalanga, Gauteng, Free State, KwaZulu-Natal, Eastern Cape); sandy soils in frost-free dry climates, moist coastal bushland and to 1850 m. I: Holstein (2015: 16–17). Incl. Coccinia ovifera Dinter (1912); incl. Coccinia rehmannii var. littoralis A. Meeuse (1962). Climbing or trailing; tuber not described; stems to several m, 2 cm ∅, greyish-green, pubescent usually becoming glabrous, becoming covered

198

with whitish or greyish scaly specks; tendrils simple; L petiole 0.2–4.2 cm, lamina pentagonalsuborbicular or pentagonal-cordate, palmately 3- to 5-lobed, lobes 3–6  3–6 cm, hirsute when young, soon glabrescent and scabrid; male Fl solitary or fasciculate, rarely racemose; hypanthium obconical to cup-shaped, 4–6 mm; Pet cream to pale yellow, 2.2–2.8 cm; female Fl solitary; Pet as in male flowers; Fr subglobose or ellipsoid, 3–4.5  2–3.5 cm, scarlet. C. sessilifolia (Sonder) Cogniaux (in A. & C. de Candolle, Monogr. Phan. 3: 534, 1881). Type [lecto]: RSA, Free State? (Drège 3375 [P]).  Distr: Namibia, Botswana, RSA.
Cephalandra sessilifolia Sonder (1862). C. sessilifolia var. sessilifolia  Distr: Namibia, Botswana, RSA (Northern Cape, North-West Prov., Limpopo, Gauteng, Mpumalanga, Free State); stony soil and sand, or loam on limestone or granite ground in semi-desert, grassland, bushland or open woodland, 300–1500 m. I: Rowley (1987: 59); Holstein (2015: 8, 12, 20, 22). Incl. Coccinia schinzii Cogniaux (1895); incl. Coccinia sessilifolia var. major Cogniaux (8890). Monoecious climbers with tuberous Rstock; stems to 5 m, slender, angular-sulcate; tendrils simple; L sessile to subamplexicaul (petiolate in var. variifolia), 3–12  3–12 cm, palmately 5-lobed, glaucous, smooth or minutely punctate-scabrid; male Fl solitary or racemose; hypanthium 3–4 mm, glabrous; Pet pale yellow to nearly white, sometimes pale dull orange-buff, strongly veined; female Fl with white or pale cream triangular staminodes; Fr oblongfusiform or elongated ellipsoid, 6–9  2–3 cm, red. C. sessilifolia var. variifolia (A. Meeuse) Holstein (PhytoKeys 54: 133, 2015). Type: RSA, Limpopo (Meeuse & Strey 10413 [PRE, BOL, L, SRGH]).  Distr: N RSA (Limpopo); low closed woodland on sandstone, 800–1200 m.
Coccinia variifolia A. Meeuse (1962). Differs from var. sessilifolia: L petiole 0.7–1.6 cm.

L. E. Newton

C. trilobata (Cogniaux) C. Jeffrey (Kew Bull. 15(3): 349, 1962). Type: Tanzania, Kilimanjaro (Volkens 1956 [B †, BR]).  Distr: Kenya, Tanzania; wooded grassland and deciduous bushland, 900–2170 m.
Peponia parviflora var. trilobata Cogniaux (1895) (incorrect name, Art. 11.4)
Peponia trilobata (Cogniaux) Engler (1895) (incorrect name, Art. 11.4)
Peponium trilobatum (Cogniaux) Engler (1897); incl. Coccinia kilimandjarica Cogniaux ex Harms (1923); incl. Coccinia kilimandjarica var. subintegrifolia Cogniaux ex Harms (1923). Climbing or trailing, with tuberous Rstock to 11.5 kg; stems to 4 m, herbaceous, hirsute; tendrils simple; L petiole 0.8–16.5 cm, lamina ovatecordate, 3–14.5  3.5–9 cm, palmately 3- to 5-lobed, rarely unlobed, lobes elliptic-rhombic, upper face scabrid-punctate, lower face hairy, sometimes densely so; male Fl 1–6 in clusters, sometimes on short leaf-less branches appearing racemose; Ped 0.5–6.5 cm; hypanthium funnelshaped, 3–5 mm; Pet yellow with green veins, lower 1=4 – 1=2 united, lobes 1.1–2 cm; St 3; female Fl solitary; Ped 0.6–2 cm; hypanthium 2–3.5 mm, densely glandular-hairy inside at the mouth; Pet almost free or lower 1=2 united, lobes 1.8–2 cm; Fr ellipsoid, 5.4–8.9  2.2–4.3 cm, bright red with green markings. In Kenya, leaves are used as a vegetable and also given to fatten goats, ripe fruits are eaten by children (Njoroge & Newton 2002).

References Bleher, B. [& al. 2003], Potgieter, C. J., Johnson, D. N. & Böhning-Gaese, K. (2003) The importance of figs for frugivores in a South African coastal forest. J. Trop. Ecol. 19(4): 375–386. https://doi.org/10.1017/ S0266467403003420. Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Holstein, N. (2015) Monograph of Coccinia (Cucurbitaceae). PhytoKeys 54: 1–166, key, maps, ills. https://doi.org/10.3897/phytokeys.54.3285. Holstein, N. & Renner, S. S. (2011) A dated phylogeny and collection records reveal repeated biome shifts in the African genus Coccinia (Cucurbitaceae). BMC Evol. Biol. 11: 28; 16 pp., maps. http://dx.doi.org/www. biomedcentral.com/1471-2148/11/28.

Coccinia CUCURBITACEAE Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. (1996) Cucurbitaceae. In: Edwards, S. & al. (eds.): Flora of Ethiopia and Eritrea, 2(2): 17–59, ills., keys. Addis Ababa (ET): National Herbarium/ Uppsala (SE): Department of Systematic Botany. Keraudren, M. (1967) Cucurbitacées. In: Flore du Cameroun, vol. 6. Paris (FR): Muséum National d’Histoire Naturelle. Njoroge, G. N. & Newton, L. E. (2002) Ethnobotany and distribution of wild genetic resources of the family Cucurbitaceae in the central highlands of Kenya. Pl. Genet. Resources Newslett. 132: 10–16.

199 Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Welman, W. G. & Condy, G. (2007) Coccinia palmata. Flow. Pl. Afr. 60: 118–122, map, t. 2238. Wilkins, M. H. (1994) Good gourds and succulent squash: A cucurbit review. Cact. Succ. J. (US) 66(2): 65–71, ills.

Corallocarpus CUCURBITACEAE L. E. Newton and G. N. Njoroge

Corallocarpus Welwitsch ex Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 831, 1867). Type: Rhynchocarpa welwitschii Naudin. – Coniandreae – Distr: Africa, Arabia, India, Madagascar. Etym: Lat. ‘corallum’/Gr. ‘kouralion’, coral; and Gr. ‘karpos’, fruit; for the coral-red fruits. Incl. Phialocarpus Deflers (1898). Type: Phialocarpus glomeruliflorus Schweinfurth ex Deflers. Incl. Gijefa (M. J. Roemer) Kuntze (1903). Type: Rhynchocarpa welwitschii Naudin. Incl. Calyptrosicyos Keraudren (1959) (nom. inval., ICN Art. 37.1). Type: not typified. Perennial scandent or trailing herbs or softly woody climbers from tuberous R, or shrublets with thickened stem and branches (C. glomeruliflorus), monoecious; stems often slightly succulent; tendrils simple, or rarely bifid or absent; L petiolate, simple, ovate to cordate-reniform in outline, usually palmately 3- to 5-lobed to finely dissected; Fl small, greenish-yellow, rotate; male Fl few to many in

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] G. N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya e-mail: [email protected]

subsessile or pedunculate short to elongated usually congested racemes; hypanthium campanulate; Sep small; Pet united at the base; St 5, sometimes 2 pairs
united giving the appearance of only 3 stamens; Fil free, short; Anth all 1-thecous but sometimes appearing as 2 2-thecous and 1 1-thecous, thecae straight; female Fl solitary or fasciculate, subsessile, similar to male flowers; staminodes 5 or absent; Sti 2- (to 3-) lobed; Fr solitary or clustered small berries to 2 cm, ovoid or ellipsoid, red, often beaked, circumscissile above a persistent green cup-like base; Se few to several, small, pear-shaped or rarely subglobose. – Cytology: n ¼ 13 (for C. epigaeus, cited from Schaefer & Renner (2011: 149)). A genus of 17 species (13 from Africa, 2 from Madagascar, 2 in Arabia, India and Pakistan according to Schaefer & Renner (2011). Only some of the species are moderately to distinctly succulent. C. bainesii (Hooker fil.) A. Meeuse (Bothalia 8: 41, 1962). Type: Botswana (Chapman & Baines s.n. [K]). – Distr: Namibia, Botswana, RSA (Limpopo, Mpumalanga, KwaZulu-Natal), Swaziland, Moçambique, Zimbabwe, Tanzania, Comoros, NW Madagascar; woodland and grassland, 350–1050 m. I: Keraudren (1966: 73); Jeffrey (1978: 489); Rowley (1987: 62).  Rhynchocarpa bainesii Hooker fil. (1871)  Kedrostis bainesii (Hooker fil.) Cogniaux (1881); incl. Corallocarpus poissonii Cogniaux (1881); incl. Corallocarpus sphaerocarpus

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_35

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202

Cogniaux (1888); incl. Corallocarpus sphaerocarpus var. hastatus Cogniaux (1895); incl. Corallocarpus sphaerocarpus var. scaberrimus Cogniaux (1895); incl. Corallocarpus bussei Gilg (1904); incl. Corallocarpus dinteri Cogniaux (1916). Climbers; stems to 1 m, usually branched at the base, esp. near the base thickened and slightly succulent, glabrous or scabrid; tendrils simple; L petiole 1.8–2.6 cm, lamina ovate or reniformcordate in outline, 3.6–5.4  5.2–7 cm, usually palmately 3- to 5-lobed, lobes elliptic, rhombic or obovate, softly hairy or hispid-hairy, slightly succulent; male Fl 3–12 in racemes to 0.5 cm on 1.5–5 cm peduncles; Ped 1.5–2 mm, minute; female Fl solitary or to
25 in sessile clusters or dense racemes; Fr ovoid or ovoid-conical, 0.9–1.6  0.5–1 cm, red. C. epigaeus (Rottler) C. B. Clarke (in Hooker fil., Fl. Brit. India 2: 628, 1879). Type [syn]: India (Rottler s.n. [K]). – Distr: Niger, Senegal, Kenya, Tanzania, Democratic Republic of the Congo [Zaïre], Sudan, Ethiopia, Yemen, Oman, W Pakistan, India; woodland and forest margins, 550–2100 m. I: Jeffrey (1967: 140), repeated by Jeffrey (1996: 26).  Bryonia epigaea Rottler (1803)  Rhynchocarpa epigaea (Rottler) Naudin (1862); incl. Rhynchocarpa corallina Naudin (1862)  Corallocarpus corallinus (Naudin) Cogniaux (1881); incl. Rhynchocarpa epigaea var. gracilipes Naudin (1862)  Corallocarpus gracilipes (Naudin) Cogniaux (1881); incl. Corallocarpus fenzlii Hooker fil. (1871) (nom. illeg., Art. 52.1); incl. Corallocarpus elegans Gilg (1904); incl. Corallocarpus hildebrandtii Gilg (1904); incl. Corallocarpus tavetensis Gilg (1904); incl. Corallocarpus bequaertii De Wildeman (1921). Climbers, with globose probably underground tuber; stems to 4 m,
glaucous, glabrous or finely pubescent, becoming thickened with age, with brownish papery bark; Br stout, succulent, jointed at the nodes; tendrils slender; L petiole robust, fleshy, 0.8–3.5 cm, lamina almost circular, 2.5–12  3.5–12 cm, palmately 3- to 5-lobed, lobes obovate or oblanceolate, often themselves

L. E. Newton and G. N. Njoroge


3-lobulate, finely hairy; male Fl 7–40 in racemes to 1 cm on 3–11 cm peduncles, often on leafless stems; Ped 1–5 mm; hypanthium 1.5–2.5 mm; Pet dull yellow or greenish-yellow, 1.5–2 mm; female Fl solitary or to 30 in racemes, often on contracted leafless stems; Ped 1–4 mm; hypanthium campanulate, 2 mm; Pet reflexed, 1.5–2.5 mm; Fr ovoid or ellipsoid, beaked, 1–2.1  0.4–0.9 cm, red except for the green base and beak, or sometimes green with white mottling. C. glomeruliflorus (Deflers) Cogniaux (in Engler, A. (ed.), Pflanzenr. IV.275(1): 174, ills., 1916). Type [syn]: Yemen (Schweinfurth 449 [P]). – Lit: Jeffrey & Thulin (1993: 221). Distr: Yemen, Somalia; rocky ground, usually gypsum hills, 200–1100 m.  Phialocarpus glomeruliflorus Deflers (1895)  Kedrostis glomeruliflora (Deflers) C. Jeffrey (1962); incl. Rhynchocarpa courbonii Deflers (1885) (nom. illeg., ICN Art. 53.1). Deciduous shrublets to 0.4 cm, stems succulent, ridged, rough, brownish, branched near the tips; L petiole 0.4–2 cm, lamina ovate to reniform, unlobed to shallowly 3- to 5-lobed, 0.7–2  0.8–2 cm, surface velvety-rough; male Fl in small sessile clusters; hypanthium globoseurceolate; Pet
3 mm; female Fl solitary; Fr ovoid, with short beak, furrowed, 1–1.5  0.5–1 cm, red. Beyenbach (2018: 231) illustrates a sterile leafless plant under this name from the Marsabit region in N Kenya, but the identification is highly questionable given the known range of the species. C. perrieri Keraudren (Bull. Soc. Bot. France 107(9): 349, 1960). Type: Madagascar, Toliara (Perrier 19118 [P]). – Distr: S Madagascar (Toliara); dry bush on sand, 0–70 m. I: Keraudren (1966: 69); Rauh (1998: 197) – Fig. 1.  Calyptrosicyos perrieri Keraudren (1959) (nom. inval., ICN Art. 35.1). Climbers from a roundish to pear-shaped subterranean tuber; stems annual, 5-angled, glabrous; tendrils simple; L petiole 2–4 cm, lamina dissected into 3 linear lobes, each further dissected

Corallocarpus CUCURBITACEAE

203

Fig. 1 Corallocarpus perrieri (Hoffmann s.n.: Madagascar; Toliara, 10 km W of Sakaraha; cult. Sukkuklenten-Sammlung Zürich). (Copyright: U. Eggli)

into secondary lobes, main lobes 3–6 cm, glabrescent; male Fl 4–10 in short racemes; Pet pale yellow, 2–3  1–1.5 mm; St 3; female Fl solitary or 2–3 together; hypanthium cylindrical, 2 mm; Fr 8  5 mm, fleshy, red. C. schimperi (Naudin) Hooker fil. (in Oliver, Fl. Trop. Afr. 2: 567, 1871). Type: Ethiopia (Schimper 413 [P]). – Distr: Ethiopia, Sudan, Somalia, Kenya, Yemen, Saudi Arabia, S Egypt, Iran, W Pakistan, India; deciduous bushland and woodland, 150–1100 m.  Rhynchocarpa schimperi Naudin (1862); incl. Rhynchocarpa pedunculosa Naudin (1862)  Corallocarpus pedunculosus (Naudin) Cogniaux (1881); incl. Rhynchocarpa courbonii Naudin (1863)  Corallocarpus courbonii (Naudin) Cogniaux (1881); incl. Rhynchocarpa ehrenbergii Schweinfurth (1868)  Corallocarpus ehrenbergii (Schweinfurth) Hooker fil. (1871); incl. Rhynchocarpa erostris Schweinfurth (1868)  Corallocarpus erostris (Schweinfurth) Hooker fil. (1871); incl. Corallocarpus etbaicus Hooker fil. (1871); incl. Corallocarpus velutinus C. B. Clarke (1879); incl. Corallocarpus brevipedunculatus Gilg (1904); incl. Corallocarpus longiracemosus Gilg (1904). Climbers; stems robust, little branched, succulent, angular-furrowed, slightly rough, developing reddish-brown or greyish papery bark, jointed at

the nodes; tendrils robust; L petiole 2.2–4 cm, lamina simple, ovate-cordate in outline, 2.5–10  3.5–15 cm, unlobed or usually with 5 obovate, rhombic or oblong lobes, slightly succulent, ash-grey, densely and finely scabrid-hairy; male Fl 8–50 in racemes to 6 cm on 0.8–12.5 cm long peduncles, often on contracted leafless stems; Ped 0.2–1.2 cm; hypanthium campanulate, 1.5–2 mm; Pet
2 mm, yellow with green veins; female Fl solitary or to 20 in sessile clusters, often on contracted leafless stems; Ped 0.1–0.3 cm; hypanthium
1 mm; Pet
1.5 mm, yellow with green veins; Fr ovoid or conical, with short beak, 1–3.5  0.3–1 cm, red, glabrous or minutely soft-hairy. C. welwitschii (Naudin) Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 831, 1867). Type: Angola (Welwitsch 799 [COI]). – Distr: Democratic Republic of the Congo [Zaïre], Angola, Namibia, RSA (“Transvaal”, Northern Cape)?; deciduous open woodland and bushland, 350– 1400 m.  Rhynchocarpa welwitschii Naudin (1862); incl. Corallocarpus welwitschii var. subintegrifolius Cogniaux (1881); incl. Corallocarpus congolensis Cogniaux (1914); incl. Corallocarpus gilgianus Cogniaux (1916); incl. Corallocarpus glaucicaulis Dinter & Gilg (1919) (nom. inval., ICN Art. 38.3); incl. Corallocarpus

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scaber Dinter & Gilg (1919) (nom. inval., ICN Art. 38.3). Tuberous perennials; caudex fusiform or napiform; stems annual; Br scrambling, to 2 m, slender to stout and succulent, almost angular, jointed at the nodes; tendrils usually filiform; L petiole 2–4 cm, lamina ovate to triangular, 4–7  almost 4–7 cm, palmately 5-lobed, lobes obovate-oblong or linear, acute, with short dense tufted hairs, ash-grey; male Fl 2–8 in capitate Inf, 2.5–4 cm ∅; hypanthium urceolate; Pet pale yellow; female Fl solitary, paired or few in fascicles; Pet pale yellow; Sti 2; Fr ellipsoid to oblong, 1.5–2  0.8–1.2 cm, scarlet. C. wildii C. Jeffrey (Kew Bull. 30(3): 488, 1975). Type: Zimbabwe (Wild 3967 [K, LISC, SRGH]). – Distr: Botswana, Tanzania, Zambia, Zimbabwe; Brachystegia and Colophospermum woodland, 750–1350 m. Incl. Melothria mildbraedii Gilg (1916)  Kedrostis mildbraedii (Cogniaux) C. Jeffrey (1962). Plants slender, from a tuberous Rstock; stems prostrate or scandent, branching, succulent, jointed at the nodes, glabrous except at the base, sparsely spiculate; tendrils slender; L petiole 1.2–4.2 cm, densely pubescent, lamina ovate or broadly ovate, cordate, 1.8–9  2–8 cm, plane or obscurely sinuate, minutely denticulate, both facs minutely scaberulous, palmately deeply 3- to 5-lobed, lobes triangular, broadly ovate, ovateelliptic, oblanceolate or linear-lanceolate, the central sometimes 3- (to 5-) lobulate; male Fl 5–10 or more in congested racemiform to subumbelliform pedunculate clusters; peduncle 1.3–12 cm; Ped 1.5–5.5 mm; hypanthium 1.5–2.4 mm; Sep lobes 1–1.8 mm, ovate-oblong to lanceolate; Pet 1.5–2 mm, yellow; female Fl 1–4, but usually solitary or paired; Ped 2–13 mm; Ov 5–7 mm, narrowly ovoid-rostrate; Fr 1.4–2.8 cm, ovoid,

L. E. Newton and G. N. Njoroge

rostrate, glabrous, red with green apex, 0.5–2.2 cm pedicellate; Se 5.1–6  2.8–3.4  1.9–2.6 mm, asymmetrically ovoid, fibrillose, with ridged 2-grooved margins. Jeffrey (1978) suggests that C. tenuissimus Buscalioni & Muschler (1913), a nomen dubium of uncertain application in the absence of a type, could be an earlier name for this species. – [L. E. Newton].

References Beyenbach, J. (2018) Auf Pflanzensafari durch Kenia. Teil 3: Der Norden, von Isiolo nach Marsabit/Teil 4 und Schluss: Der Norden, Marsabit und Moyale. Avonia 36(3): 170–177, (4): 224–233, ills., map. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. (1978) Cucurbitaceae. In: Launert, E. (ed.): Flora Zambesiaca, 4: 414–499, ills., keys. London (GB): Flora Zambesiaca Managing Committee. Jeffrey, C. (1996) Cucurbitaceae. In: Edwards, S. & al. (eds.): Flora of Ethiopia and Eritrea, 2(2): 17–59, ills., keys. Addis Ababa (ET): National Herbarium/ Uppsala (SE): Department of Systematic Botany. Jeffrey, C. & Thulin, M. (1993) Cucurbitaceae. In: Thulin, M. (ed.): Flora of Somalia, 1: 216–240, ills., keys. Kew (GB): Royal Botanic Gardens. Keraudren, M. (1966) Cucurbitacées. 185e famille. In: Humbert, H. (ed.): Flore de Madagascar et des Comores. Paris (FR): Muséum National d’Histoire Naturelle, Laboratoire de Phanérogamie. http://www. biodiversitylibrary.org/page/9186087 Rauh, W. (1998) Succulent and xerophytic plants of Madagascar. Vol. 2. Mill Valley (US): Strawberry Press. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10.

Cucumis CUCURBITACEAE L. E. Newton

Cucumis Linné (Spec. Pl. [ed. 1], 1011, 1753). Type: Cucumis sativus Linné [lectotype, designated by Britton & Wilson, Sci. Surv. Porto Rico, 6, Bot. Porto Rico & Virgin Isl., 1925].
Benincaseae
Lit: Kirkbride (1994: revision Cucumella); Renner & al. (2007: molecular phylogeny); Ghebretinsae & al. (2007a: molecular phylogeny & biogeography); Schaefer (2007: classification, key); Sebastian & al. (2010: biogeography). Distr: Africa, Asia, Australia. Etym: Lat. ‘cucumis’, cucumber, gherkin; i.e. the ancient name for C. sativus. Incl. Melo Miller (1754). Type: Cucumis melo Linné [lectotype, designated by Swart in Farr & al. (eds.): Index Nom. Gen. Pl., 2, 1979]. Incl. Mukia Arnott (1840). Type: Bryonia scabrella Linné fil. Incl. Oreosyce Hooker fil. (1871). Type: Oreosyce africana Hooker fil. Incl. Dicoelospermum C. B. Clarke (1879). Type: Dicoelospermum ritchiei C. B. Clarke. Incl. Hymenosicyos Chiovenda (1911). Type: Cucumis membranifolius Hooker fil. Incl. Cucumella Chiovenda (1929). Type: Cucumella robecchii Chiovenda.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

Incl. Myrmecosicyos C. Jeffrey (1962). Type: Myrmecosicyos messorius C. Jeffrey. Climbing or trailing annual or perennial herbs, monoecious or rarely dioecious, sometimes with tuberous Rstock; stems annual, usually  hispid or scabrid-hairy; tendrils simple, rarely absent, very rarely bifid; L petiolate, simple, palmately 3- to 7-lobed; Fl small to medium-sized, usually yellow; male Fl solitary or a few in clusters; hypanthium campanulate; Sep small, 5 or rarely 4; Pet 5, united at the base, St 3, free; Anth 2 2-thecous, 1 1-thecous, thecae lateral, straight, sometimes apically hooked, or triplicate; female Fl usually solitary; Pet as in male flowers; Fr subglobose, ellipsoid or elongated, sometimes beaked, fleshy, indehiscent, glabrous or pubescent, smooth or prickly or hairy; Se few to many, small to medium-sized, ovate to elliptic, globose or compressed, light-coloured, smooth or ornamented, glabrous or rarely puberulent, margin distinct.
Cytology: 2n ¼ 7, 12 (Darlington & Wylie 1955). In traditional classifications, Cucumis included some 33 species, with a centre of diversity in tropical and S Africa. Ghebretinsae & al. (2007a), Kocyan & al. (2007) and Renner & al. (2007) all investigated the phylogeny of Cucumis and related genera on the basis of molecular markers, and the genera Cucumella (11 species), Dicaelospermum (1 species), Mukia (6 species), and the monotypic Myrmecosicos and Oreosyce have been found to be embedded

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amongst Cucumis species. According to Ghebretinsae & al. (2007a), 6 major clades can be recognized. In this revised circumscription, Cucumis is a medium sized-genus of 52 (Schaefer 2007, Ghebretinsae & al. 2007b) to 65 species (Schaefer & Renner 2011, taking into account at least 9 undescribed species mentioned by Sebastian & al. 2010). Ghebretinsae & al. (2007a) postulate at least 4 migrations from Africa to Asia, and 1 migration in backwards direction, and Sebastian & al. (2010) date the divergence between the African and the Asian/Australian clade at 12 mybp. Schaefer (2007) presents an infrageneric classification recognizing 2 subgenera (Subgen. Humifructus from S Africa [only the non-succulent C. humifructus Stent and the succulent C. hirsutus] and subgen. Cucumis, comprising all other species, and divided into 5 sections) and a key to the 52 species he recognized. Some species are widespread in cultivation for their edible fruits, and have a long domestication history, including C. melo Linné with many commercially grown cultivars (incl. Cantaloupe Melon, Honeydew Melon etc.) and C. sativus Linné, the ordinary cultivated Cucumber. Only a handful of species can be considered to have succulent tubers. C. bryoniifolius (Merxmüller) Ghebretinsae & Thulin (Novon 17(2): 177 [20. June], 2007). Type: Zimbabwe (Dehn R25 [M]).
Lit: Jeffrey (1967: 114); Kirkbride (1994: 177); both as Cucumella. Distr: Tanzania, Zambia, Zimbabwe, Malawi, N RSA (Limpopo, Mpumalanga); woodland and grassland, usually on and among granite rocks, 750–1470 m.  Hymenosicyos bryoniifolius Merxmüller (1953)  Cucumella bryoniifolia (Merxmüller) C. Jeffrey (1962); incl. Cucumis umbrosus A. Meeuse & Strey (1962). Monoecious climbers or trailers; R fusiform; stems prostrate or scandent, herbaceous, slender, annual, sparsely hispid-spiculate; L petiole 0.9–3 cm, lamina ovate-cordate in outline, 1–2.9  1.5–3.8 cm, palmately 3- to 5-lobed, lobes triangular, shortly hispid-hairy; male Fl solitary or paired; Ped 1.2–2 cm; hypanthium campanulate,  3 mm; Pet  5 mm, yellow; female Fl

L. E. Newton

on Ped 0.4–1.7 cm; hypanthium 3 mm; Pet  5 mm; Fr ellipsoid, 2.1–2.9  1.3–1.7 (
2) cm, dark green becoming paler green or orange, obscurely striped with white to darker orange longitudinal bands; Se 3.2  2  0.5 mm, ovate, compressed, smooth. Sometimes erroneously described as annual, but merely with annually deciduous stems. The report for Moçambique is erroneous and based on a misidentification of another species (Lebrun & Stork 2003: 448). The occurrence in Namibia is based on a single specimen listed by Kirkbride (1994: 177) and needs confirmation. C. cinereus (Cogniaux) Ghebretinsae & Thulin (Novon 17(2): 177, 2007). Type: Namibia, Otjozondjupa (Dinter 1440 [Z, BR]).
Distr: Namibia, Angola, N RSA (North-West Prov., Limpopo, Mpumalanga), Kenya, Tanzania, S Madagascar (Toliara); seasonally deciduous forests on sand or limestone, 330–3240 m. I: Keraudren (1966: 55); Fernandes & Fernandes (1970: t. 29); both as Cucumella.  Kedrostis cinerea Cogniaux (1901)  Cucumella cinerea (Cogniaux) C. Jeffrey (1962)  Melothria cinerea (Cogniaux) A. Meeuse (1962); incl. Melothria breyeri Burtt Davy (1926); incl. Melothria hispidula Burtt Davy (1926); incl. Kedrostis gracilis R. Fernandes (1959). Monoecious tuberous vine, herb or subshrub, caudex with cortex splitting into scales; stem thickened and woody at the base, simple or often  branched; Br to 25 cm, slender, angularfurrowed, hirsute at first, hispidulous or retrorsestrigose, hairs 0.1–1 mm, sometimes longer on the ridges and shorter in the grooves; tendrils simple, filiform, 4–5 cm; L petiole 1.5–5.5 cm, hispidulous or retrorsely strigose, hairs to 6 mm, lamina ovate, 1.8–6.5  1.5–6 cm, 3-lobed or (rarely) palmately 5-lobed, cordate at the base, tip acute to broadly acute or obtuse, hispidulous on the upper face and on the veins below, margins crenate-curly, central lobe broadly triangular or shallowly ovate, lateral lobes shallowly triangular or shallowly ovate; male Fl 5–19 in short dense raceme; Ped 0.2–1 cm; hypanthium campanulate, 2.8–4 mm, hispidulous; Sep narrowly oblong, linear or triangular, 0.7–2.4 mm; Pet elliptic to

Cucumis CUCURBITACEAE

broadly elliptic or obovate, 2.6–5.6  2–5.6 mm, outside puberulent; female Fl solitary; Ped 0.28–1.4 cm; hypanthium hour-glass shaped, ridged, lower 2=3 – 3=4 fused to the ovary; Sep narrowly oblong or narrowly triangular, 1–1.3 mm; Pet obovate; Fr ovoid, 1.5–2.5  0.6–1 cm, hispidulous, orange, apex rostrate, beak 0.7–1 cm; Se ovate, 4.2–5.2  2.4–3 mm, glabrous.
[L. E. Newton & U. Eggli] C. hirsutus Sonder (in Harvey & Sonder, Fl. Cap. 2: 497, 1862). Type [lecto]: RSA, Mpumalanga (Zeyher 581 [MEL]).
Lit: Jeffrey (1967: 107–108); Keraudren (1967: 139–142, with ill.). Distr: S Sudan, Cameroon, Democratic Republic of the Congo [Zaïre], Zambia, Tanzania, Kenya, Angola, Moçambique, Botswana, Zimbabwe, RSA (North-West Prov., Limpopo, Gauteng, Mpumalanga, KwaZulu-Natal, Eastern Cape), Lesotho, Madagascar; deciduous woodland and bushland, grassland, 30–1500 (
2500) m. I: Wilkins (1994: 68).
Fig. 1. Incl. Cucumis sonderi Cogniaux (1881); incl. Cucumis welwitschii Cogniaux (1881)  Cucumis hirsutus var. welwitschii (Cogniaux) R. & A. Fernandes (1962); incl. Cucumis hirsutus var. dissectus Cogniaux (1895); incl. Cucumis seretii De Wildeman (1909); incl. Cucumis homblei Cogniaux (1916); incl. Cucumis wildemanianus Cogniaux (1916); incl. Cucumis hirsutus var. Fig. 1 Cucumis hirsutus (Zimbabwe; Bulawayo Road, Matesi Safari Area). (Copyright: B. T. Wursten / Online Flora of Zimbabwe)

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major Cogniaux (1924); incl. Cucumis hirsutus var. ovatus Cogniaux (1924); incl. Cucumis seretioides Suessenguth (1951). Dioecious trailing perennial herbs with large woody Rstock; stems to 2.5 m, angular, hairy; L petiole 0.3–5.5 cm, lamina ovate, ovate-triangular, elliptic or lanceolate in outline, 1.2–14.3  1–9.5 cm, unlobed or 3- to 5-lobed, softly to scabrid-hairy, esp. on the lower face; male Fl solitary or 2–14 together in 0.1–4.7 cm pedunculate groups; Ped 0.4–6 cm; hypanthium 3.5–7 mm; Pet united at the base, 6–21 mm, white, cream or pale yellow with green veins; female Fl 1–3; Ped 1.1–2.3 cm; hypanthium 4 mm; Pet united at the base, 7–10 mm; Fr subglobose, 4–4.7  3.7–4.2 cm, brownish-orange. C. kalahariensis A. Meeuse (Bothalia 8: 70–71, 1962). Type: Namibia (Story 5320 [PRE]).
Distr: S Namibia, Botswana, RSA (Northern Cape); deep sandy soil, to 1190 m. Dioecious trailers; R fibrous, with cylindrical or fusiform swellings to 20  3 cm; stems annual, to 3 m, angular-sulcate, minutely and sparsely hirsute when young, later scabrid to subglabrous, branched; tendrils short, thickened at the base; L petiole 1–5 cm, lamina ovate or ovate-oblong in outline, 5–15  3–9 cm, palmately 3- to 7-lobed, greyish-green, scabrid; male Fl solitary or sometimes several in fascicles; hypanthium

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campanulate,  7 mm; Pet 6–7 mm, yellow; female Fl solitary; Pet 9–15 mm, yellow; Fr ellipsoid or oblong, 3–5.5  2–3 cm, with thick soft prickles, greenish-white with brownishpurple longitudinal bands.

References Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Fernandes, R. & Fernandes, A. (1970) Cucurbitaceae. In: Exell, W. & al. (eds.): Conspectus Florae Angolensis. Vol. IV; pp. 232–289, ills., keys. Lisboa (PT): Junta de Investigações do Ultramar / Instituto de Investigação Científica de Angola. Ghebretinsae, A. G. [& al. 2007a], Thulin, M. & Barber, J. C. (2007) Relationships of cucumbers and melons unraveled: Molecular phylogenetics of Cucumis and related genera (Benincaseae, Cucurbitaceae). Amer. J. Bot. 94(7): 1256–1266. https://doi.org/10.3732/ajb.94.7.1256. Ghebretinsae, A. G. [& al. 2007b], Thulin, M. & Barber, J. C. (2007) Nomenclatural changes in Cucumis (Cucurbitaceae). Novon 17(2): 176–178. https://doi.org/ 10.3417/1055-3177(2007)17[176:NCICC]2.0.CO;2. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Keraudren, M. (1966) Cucurbitacées. 185e famille. In: Humbert, H. (ed.): Flore de Madagascar et des Comores. Paris (FR): Muséum National d’Histoire Naturelle, Laboratoire de Phanérogamie. http://www. biodiversitylibrary.org/page/9186087 Keraudren, M. (1967) Cucurbitacées. In: Flore du Cameroun, vol. 6. Paris (FR): Muséum National d’Histoire Naturelle.

L. E. Newton Kirkbride, J. H. jr. (1994) Revision of Cucumella (Cucurbitaceae, Cucurbitoideae, Melothrieae, Cucumerinae). Brittonia 46(3): 161–186, key, ills., map. https://doi.org/10.2307/2807230. Kocyan, A. [& al. 2007], Zhang, L.-B., Schaefer, H. & Renner, S. S. (2007) A multi-locus chloroplast phylogeny for the Cucurbitaceae and its implications for character evolution and classification. Molec. Phylogen. Evol. 44(2): 553–577. https://doi.org/10. 1016/j.ympev.2006.12.022. Lebrun, J.-P. & Stork, A. L. (2003) Tropical African flowering plants. Ecology and distribution. Volume 1: Annonaceae - Balanitaceae. Genève (CH): Éditions des Conservatoire et Jardin Botaniques. Renner, S. S. [& al. 2007], Schaefer, H. & Kocyan, A. (2007) Phylogenetics of Cucumis (Cucurbitaceae): Cucumber (C. sativus) belongs in an Australian/Asian clade far from African melon (C. melo). BMC Evol. Biol. 7: 58, 11 pp. https://doi.org/10.1186/1471-21487-58. Schaefer, H. (2007) Cucumis (Cucurbitaceae) must include Cucumella, Dicoelospermum, Mukia, Myrmecosicyos and Oreosyce: A recircumscription based on nuclear and plastid DNA data. Blumea 52(1): 165–177, key. https://doi.org/10.3767/ 000651907X612427. Schaefer, H. & Renner, S. S. (2011) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi.org/10. 1002/tax.601011. Sebastian, P. M. [& al. 2010], Schaefer, H., Telford, I. R. H. & Renner, S. S. (2010) Cucumber and melon have their wild progenitors in India, and the sister species of Cucumis melo is from Australia. Proc. Nation. Acad. Sci. USA 107: 14269–14273. Wilkins, M. H. (1994) Good gourds and succulent squash: A cucurbit review. Cact. Succ. J. (US) 66(2): 65–71, ills.

Cucurbita CUCURBITACEAE L. E. Newton

Cucurbita Linné (Spec. Pl. [ed. 1], 1010, 1753). Type: Cucurbita lagenaria Linné [lectotype, designated by Britton & Brown, Ill. Fl. North US, ed. 2, 3: 291, 1913 (fide ING)]. – Cucurbiteae – Lit: Bemis & Whitaker (1969: ecology & morphology). Distr: USA, Mexico, Central America, West Indies, N South America; cultivated worldwide. Etym: Lat. ‘cucurbita’, gourd, in ancient times used for species of the modern genera Citrullus and Lagenaria, since Renaissance botany books used for this New World genus (Genaust 1996). Incl. Melopepo Miller (1754). Type: not designated. Incl. Pepo Miller (1754) (nom. illeg., ICN Art. 52.1). Type: Cucurbita lagenaria Linné. Incl. Ozodycus Rafinesque (1832). Type: Cucumis perennis E. James. Incl. Sphenantha Schrader (1838). Type: Sphenantha scabra Schrader. Incl. Mellonia Gasparrini (1847). Type: Cucurbita melopepo Linné. Incl. Pileocalyx Gasparrini (1847).Type: Pileocalyx elegans Gasparrini. Incl. Tristemon Scheele (1848).Type: Tristemon texanum Scheele.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

Climbing or trailing annual or perennial herbs, often with large thick tuberous R or taproots, usually monoecious; stem scabrous, angled or striate, with trichomes, sometimes rooting at the nodes; tendrils 2- to 7-fid, or rarely simple, or absent in some cultivars; L ovate to cordate to suborbicular in outline, usually pedately lobed with 3–5 (–7) lobes; Fl solitary in the leaf axils, showy and large, sometimes scented; male Fl hypanthium cylindrical to campanulate; Sep lanceolate, tip sometimes foliaceous; Pet 5, large, basally connate, yellow; St 3; Fil mostly free; Anth 2 2-thecous, 1 1-thecous, connate and forming a central head, thecae reflexed; female Fl similar to male flowers; Ov globose, oblong, cylindrical or pear-shaped with constricted apex and 3–5 placentae; Fr globose, large, fleshy, indehiscent, smooth or ribbed; Se many, ovate to elliptic, strongly compressed, smooth, cream-coloured or black, margined or not. – Cytology: x ¼ 10, 12 (Darlington & Wylie 1955) or n ¼ 20 (McKay 1931). A genus of 15 species, of which several are of economic importance and grown for their often enormous large fruits or seeds rich in oils and/or protein. The vernacular names Squash and Pumpkin refer to a variety of species including C. pepo Linné, C. moschata (Duchesne) Poiret and others. Cultivars of C. pepo include “Zucchinis” etc. grown as vegetables and eaten when still immature. A selection of taxa with particularly thick roots is here covered by way of example.

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C. cordata S. Watson (Proc. Amer. Acad. Arts 24: 50, 1889). Type: Mexico, Baja California (Palmer 584 [GH]). – Distr: NW Mexico (Baja California, Baja California Sur); sandy plains. I: Duthion (2003: 15). Incl. Cucurbita cylindrata L. H. Bailey (1943). Trailers with a long massive R, sometimes partly exposed; stems with dense rigid strigose reflexed pubescence; tendrils short, glandulartipped; L broadly cordate, 5–10.2 cm wide, with dense rigid strigose pubescence, not reflexed, 3- to 5-lobed nearly to the middle, with narrow sinuses, lobes coarsely and obtusely toothed; Fl solitary on peduncles 3.8 cm; Cl 5 cm long; Ov pubescent, subglobose; Fr globose, 7.6 cm ∅, smooth, green with 10 whitish longitudinal lines; Se white, 8.5–10.5 mm. C. digitata A. Gray (Pl. Wright. 2: 60, 1853). Type: USA, New Mexico (Wright 1088 [GH, PH]). – Lit: Hodgson (2001: 158–159, ethnobotany, with ill.); Austin (2010: 123–124, ethnobotany, with ill.). Distr: SW USA (SE California, S Arizona, S New Mexico, Texas), NW Mexico (Baja California, Sonora, Chihuahua); dry plains and mesas, sandy alluvium and along washes, to 1525 m. I: Duthion (2003: 17–18); Nesom (2015: 48). Trailers with tuberous R; stems glabrous, ribbed, whitish-pustulate; tendrils with 3–5 branches; L petiole stout, usually shorter than the lamina, lamina palmately 5-lobed, lobes 4–10 cm, with conical trichomes; Fl hypanthium cylindrical to campanulate, 2.5–3 cm; Cl narrowly campanulate, 4–7 cm; Fr (6-) 7–9.5 cm, pale yellow with longitudinal stripes; Se ovate, 8–11  7–8 mm, margins thick, obtuse. – Cytology: 2n ¼ 40 (McKay 1931). Similarly used as C. foetidissima. – Vernacular name: “Coyote Gourd”. C. foetidissima Kunth (in Humboldt & al., Nov. Gen. Sp. 1: 123, 1817). Type: Mexico, Guanajuato (Bonpland & Humboldt 4280 [P]). – Lit: Hodgson (2001: 158–159, ethnobotany, with ill.). Distr: SW to C & SE USA (California, Arizona, Colorado, Illinois, Missouri, Oklahoma, Texas), N to C Mexico (widespread, to Veracruz); sandy flats or plains, 300–2130 (–2400) m. I: Duthion (2003: 19). – Figs. 1 and 2.

Fig. 1 Cucurbita foetidissima (USA; Texas, Llano County, Oxford Ranch Campground). (Copyright: Mark Merriwether Vorderbruggen, Ph.D)

 Pepo foetidissima (Kunth) Britton (1913); incl. Cucumis perennis E. James (1820)  Ozodycus perennis (E. James) Rafinesque (1832)  Cucurbita perennis (E. James) A. Gray (1850). Trailers with large deep-reaching tuberous R; stems to 6 m, scabrous, crushed with foetid odour; tendrils thick; L petiole 5–15 cm, lamina triangular-ovate, 10–30 cm, lobes slightly angular, upper face white-pubescent, lower face scabrous; Fl hypanthium 1.27–1.7 cm; Cl 7.6–10 cm, hairy; Fr depressed-globose, 6–10 cm, green with whitish stripes; Se oblong-ovate, 12  6–7 mm. – Cytology: (McKay 1931). The roots were traditionally used as a laxative, while the seeds are a source of protein and oil (Hodgson 2001: 158–159). Vernacular names: “Buffalo Gourd”, “Chilicote”, etc. C. palmata S. Watson (Proc. Amer. Acad. Arts 10(1): 137, 1876). Type [syn]: USA, California (Cleveland s.n. [GH]). – Lit: Hodgson (2001: 158–159, ethnobotany, with ill.). Distr: S USA (California, Arizona), NW Mexico (Baja California,

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Fig. 2 Cucurbita foetidissima (USA; Arizona, Sedona). (Copyright: A. Reeves/ Flickr, CC BY-ND)

Sonora); dry sandy plains and mesas, to 900 m. I: Wiggins (1980: 388); Schlising (1993: 537). Incl. Cucurbita californica Torrey ex S. Watson (1876). Trailers with deep-reaching fusiform R; stems strongly ribbed, densely pilose when young; tendrils mostly 3-branched; L petiole as long as or exceeding the lamina, lamina to 10  7.6–11 cm, palmately 5-lobed, lobes triangular-lanceolate, upper face light green, hirsute to hispid, lower face hispid; Fl Ped 3–5 cm; hypanthium campanulate, 2–3.5 cm; Cl 4.5–7 cm; Fr depressed-globose, 7.6 cm ∅, dull light green with 10 faint stripes; Se ovate to oblong, 9–12  6–8 mm, white. – Cytology: 2n ¼ 40 (McKay 1931). Similarly used as C. foetidissima.

References Austin, D. F. (2010) Baboquivari Mountain plants. Identification, ecology and ethnobotany. Tucson (US): University of Arizona Press.

Bemis, W. P. & Whitaker, T. W. (1969) The xerophytic Cucurbita of northwestern Mexico and southwestern United States. Madroño 20(2): 33–41. http://www.jstor. org/stable/41423342 Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Duthion, N. (2003) Three Cucurbitaceae caudiciforms. Cact.-Avent. 57: 15–19, ills. Genaust, H. (1996) Etymologisches Wörterbuch der botanischen Pflanzennamen. Ed. 3. Basel (CH), etc.: Birkhäuser Verlag. Hodgson, W. C. (2001) Food plants of the Sonoran Desert. Tucson (US): University of Arizona Press. McKay, J. W. (1931) Chromosome studies in the Cucurbitaceae. Univ. Calif. Publ. Bot. 16: 339–350, ills. Nesom, G. L. (2015) Cucurbitaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 3–58, ills., keys. New York (US)/Oxford (GB): Oxford University Press. Schlising, R. L. (1993) Cucurbitaceae. In: Hickman, J. C. (ed.): The Jepson Manual; pp. 535–538, ills., key. Berkeley/Los Angeles (US), etc.: University of California Press. https://ucjeps.berkeley.edu/ eflora/ Wiggins, I. L. (1980) Flora of Baja California. Stanford (US): Stanford University Press.

Cyclantheropsis CUCURBITACEAE L. E. Newton

Cyclantheropsis Harms (Bot. Jahrb. Syst. 23(1–2): 169, 1896). Type: Gerrardanthus parviflorus Cogniaux. Triceratieae Distr: E & S tropical Africa, Madagascar. Etym: Gr. ‘-opsis’, similar to; and for the genus Cyclanthera (Cucurbitaceae). Climbers with tuberous Rstock, dioecious; stems herbaceous, becoming softly woody with age; tendrils apically forked; L simple, entire or 3- to 5-lobed; Fl very small, greenish-yellow; male Fl many, in axillary panicles; hypanthium shallow, saucer-shaped; Sep minute, 0.5–1 mm; Pet very small, 1 mm; St 1, in the flower centre; Anth with 2 horizontal semicircular thecae forming a split ring; female Fl 3–6, in thyrses or monochasia, otherwise as male flowers, with 3 staminodes; Sty 3; Sti bifid; Fr compressed, samaroid, elliptic in outline, to 5.5  2.2 cm, brown, indehiscent; Se 1 per fruit, elliptic, compressed, to 11 mm, pale brown, slightly rough. A small genus with 3 species (2 from Africa, 1 from Madagascar), of which only the following is occasionally grown as a caudiciform, and thus covered here:

C. parviflora (Cogniaux) Harms (Bot. Jahrb. Syst. 23(1–2): 169, 1896). Type: Tanzania, Zanzibar (Hildebrandt 1140 [W, L]). Distr: Ethiopia, Kenya, Tanzania, Angola, Namibia, Zambia, Zimbabwe, Malawi, Moçambique; lowland evergreen forests, deciduous woodland and AcaciaCommiphora bushland and grassland, 75–1150 m. I: Jeffrey (1967: 149), repeated by Jeffrey (1996: 21) and Schaefer & Renner (2011: 133). Fig. 1.  Gerrardanthus parviflorus Cogniaux (1881); incl. Gerrardanthus aethiopicus Chiovenda (1939). Climbers; stems perennial, to 5 m, base swollen and tuberous, to 6 cm ∅, semi-underground, stems puberulous when young, with grey bark when older; L petiole 1–4.1 cm, lamina ovate or reniformcordate in outline, 2.1–10.5  3.2–10 cm, usually with 3–5 triangular lobes, acute, apiculate, minutely puberulous, semi-succulent; male Fl in panicles to 12 cm; hypanthium saucer-shaped; Sep triangular, 0.7 mm; Pet 1 mm, pale greenish-yellow; female Fl 3–6 in 2.5–4.5 cm panicles or racemes on 0.5–1 cm peduncles; Ped 0.4–0.6 cm; Pet 1.2 mm; Fr compressed, samaroid, elliptic in outline, indehiscent, 3.1–5.5  1.5–2.2 cm, light brown.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_38

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References Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. (1996) Cucurbitaceae. In: Edwards, S. & al. (eds.): Flora of Ethiopia and Eritrea, 2(2): 17–59, ills., keys. Addis Ababa (ET): National Herbarium/ Uppsala (SE): Department of Systematic Botany. Schaefer, H. & Renner, S. S. (2011) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10.

Fig. 1 Cyclantheropsis parviflora (Kenya?, cult. Copenhagen Botanical Garden). (Copyright: E. Bihrmann)

Dendrosicyos CUCURBITACEAE L. E. Newton

Dendrosicyos Balfour fil. (Proc. Roy. Soc. Edinburgh 11: 513, 1882). Type: Dendrosicyos socotranus Balfour fil. – Coniandreae – Lit: Olson (2003: anatomy); Habrová & al. (2020: population biology). Distr: Yemen (Socotra). Etym: Gr. ‘dendron’, tree; and Gr. ‘sicyos’, cucumber; for the large size of the plants and the placement in this family. Arborescent monoecious stem succulents; stem erect, to 7  1.2 m ∅, branching sparsely above; Br few, somewhat pendulous, thin, surface rough, wrinkled, densely hispid at first; tendrils absent; L petiolate, simple, ovate-cordate, to 10  10 cm, deeply 4- to 6-lobed, surface rough and somewhat prickly from white bristles; Fl unisexual, 2.5 cm ∅, in axillary fascicles; Sep lanceolate, 5 mm; Pet 5, yellow, pubescent; male Fl Ped 0.7–1.1 mm; hypanthium funnelshaped, 19–25 mm; Pet linear-lanceolate, 12.5 mm; St 3, exserted, with straight thecae; female Fl Ped 1 mm; hypanthium cup-shaped, 15–25 mm; Pet oblong, 9.5 mm; staminodes 5; Ov ovoid; Sty long and slender; Sti 3; Fr cylindrical, to 4 cm, with apical beak, glandularhirsute, fleshy, orange; Se few, 6 mm, compressed.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

This monotypic genus is remarkable and unique in the family for the arborescent habit and the massively well-developed stem succulence. Schaefer & al. (2009: 849) found Dendrosicyos in a near-basal position in tribe Coniandreae in their dated molecular phylogeny. The lineage that includes Dendrosicyos was estimated to be 22 (confidence interval 30–14) my old, which would be older than the estimated age of some 10 my for the island of Socotra. This dating was subsequently revised and the calculated age of 8–19 Ma is more in line with the age of the island (Renner & Schaefer 2016). The anatomy of Dendrosicyos was studied by Olson (2003): The wood of the stem is highly parenchymatic, with mainly parenchyma tissue and only small strands of xylem in an interconnected network. Phloem is produced from each xylem strand, resulting in successive cambia. The leaves are thin and without succulence. Habrová & al. (2020) found, based on a grid study covering the whole island, that populations are generally healthy (which is at variance with the commonly seen opinion that it is endangered by grazing), with a good size-class distribution, and at densities of up to 6.03 trees per km2. Regeneration and establishment of juveniles was recorded for 77% of the studied populations. D. socotranus Balfour fil. (Proc. Roy. Soc. Edinburgh 11: 513, 1882). Type: Yemen, Socotra (Schweinfurth 243 [K]). – Distr: Yemen (Socotra); rocky places, to 500 m. I: Rowley (1987:

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53, 189); Mies & Zimmer (1993); Mies (1995); Mies (1998: 51–52); Mies (1999); Schaefer & Renner (2011: 147); Lenain (2012: 106–107). – Figs. 1 and 2.

L. E. Newton

Incl. Dendrosicyos jaubertianus (1895). Description as for the genus.

Baillon

References

Fig. 1 Dendrosicyos socotranus (Mies 672: Yemen; Socotra, Nuged Plain; cult. Sukkulenten-Sammlung Zürich). (Copyright: U. Eggli)

Fig. 2 Dendrosicyos socotranus (details as above). (Copyright: U. Eggli)

Habrová, H. [& al. 2020], Vahalík, P., Drápela, K. & Ali, A. S. S. (2020) Distribution and population characteristics of the Cucumber Tree (Dendrosicyos socotranus Balf. f.). Rendic. Lincei Sci. Fis. Nat. 31(1): 1–12. https://doi.org/10.1007/s12210-020-00927-5. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer. Mies, B. A. (1995) Die Sukkulenten des Sokotrischen Archipels (Jemen, Indischer Ozean). Teil 1. Kakt. and. Sukk. 46(4): 92–96, diag., maps, ills. Mies, B. A. (1998) Leben auf grossem Fuss. Untersuchungen in [sic!] Caudexpflanzen der Insel Sokotra (Jemen). Kakt. and. Sukk. 49(3): 49–52; (4): 85–90, ills. Mies, B. A. (1999) Bottle-trees and heat on Socotra (Yemen): A big-foot story. A temperature study of Adenium socotranum (Apocynaceae) and Dendrosicyos socotrana [sic!] (Cucurbitaceae). Cact. Succ. J. (US) 71(2): 60–73, ills. Mies, B. A. & Zimmer, H. (1993) Die Populationen von Adenium socotranum (Balfour) Vierhapper und Dendrosicyos socotrana [sic!] Balfour bei Ras Hebak (Insel Sokotra, Jemen) und ihre Gefährdung. Kakt. and. Sukk. 45(1): 1–5, ills. Olson, M. E. (2003) Stem and leaf anatomy of the arborescent Cucurbitaceae Dendrosicyos socotrana [sic!] with comments on the evolution of pachycauls from lianas. Pl. Syst. Evol. 239(3–4): 199–214, ills. https://doi.org/ 10.1007/s00606-003-0006-1. Renner, S. S. & Schaefer, H. (2016) Phylogeny and evolution of the Cucurbitaceae. In: Grumet, R. & al. (eds.): Genetics and genomics of Cucurbitaceae; pp. 13–23. Cham (CH): Springer International Publishing. https:// doi.org/10.1007/7397_2016_14. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Schaefer, H. [& al. 2009], Heibl, C. & Renner, S. S. (2009) Gourds afloat: A dated phylogeny reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous oversea dispersal events. Proc. Roy. Soc., Biol. Sci., Ser. B, 276(1658): 843–851, maps. https://doi. org/10.1098/rspb.2008.1447.

Doyerea CUCURBITACEAE L. E. Newton

Doyerea Grosourdy (Med. Bot. Criollo 1(2): 338, 1864). Type: Doyerea emetocathartica Grosourdy. – Coniandreae – Lit: Belgrano & Pozner (2017: 33–35). Distr: Mexico, Caribbean, Central & South America (Venezuela, Colombia, NE Brazil). Etym: For Louis Michel Français Doyère (1811–1863), French zoologist and agronomist. Incl. Anguriopsis J. R. Johnston (1905). Type: Anguriopsis margaritensis J. R. Johnston. Dioecious perennial climbers to 6 m with thick Rstock; stem forming a swollen trunk to 35 cm ∅ with fleshy Br; tendrils simple; L (1.5–) 3–4 (–8) cm petiolate, deciduous, lamina simple, cordateovate or 3- to 5-lobed or 3-foliolate, 4–9 (–14)  3.5–8 (–16) cm, pubescent to glabrous, margins denticulate; Fl minute, small, greenish-yellow, Sep valvate in bud; male Fl crowded in pedunculate up to 40-flowered racemes; Ped 2 mm; hypanthium 1.5–2 mm; Sep 5, 1–1.5 mm; Pet 5, 1–2  0.5–1 mm, ovate-oblong to ovate-lanceolate, greenish-yellow to cream-coloured or ochre; St 3, inserted in the mouth of the hypanthium, completely free; Fil short or almost none; Anth 3, 2 2-thecous, 1 1-thecous, thecae curved; female Fl subsessile in dense fascicles, hypanthium and Pet as in male flowers; Ov fusiform, 2–3 mm,

with 2 placentae; Sty thick, simple; Sti 2, fimbriate; Fr ellipsoid to oblong-ovoid, (0.8–) 1.3–2 (–3)  0.4–0.7 cm, shortly beaked, thin-walled, fleshy, green to maroon with white stripes or blotches turning red to dark purple when fully mature; Se 6–10 per fruit, asymmetrically ovoid to pear-shaped, 2.5–4.9 (–5.2)  2.3–3.4 mm, brown with paler margins. This monotypic genus has a wide distribution from Mexico to N South America, with an outlying area in NE Brazil, where its sole species was for long known as Apodanthera congestiflora. Belgrano & Pozner (2017) have shown this and another Apodanthera species to be conspecific with the only species of Doyerea. Mooney & al. (1992) found a root and stem water content of some 80%, with very little fluctuation between dry and wet season; flowering and fruiting occurs during the dry season. D. emetocathartica Grosourdy (Med. Bot. Criollo 1(2): 338, 1864). Type: Puerto Rico (Grosourdy s.n. [P]). – Lit: Dieterle (1976: 337–339, with ills.); Jeffrey & Trujillo (1992: 35–39, with ills.); Lira (2001: 49–52, with ills.); McVaugh (2001: 554–557, with ills.). Distr: Mexico (very widespread from Sinaloa and

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_40

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Fig. 1 Doyerea emetocathartica (Hammel & Pérez 24997 [CR, MO]: Costa Rica; Guanacaste, Nicoya Peninsula, 500 m). (Copyright: Barry Hammel)

Fig. 2 Doyerea emetocathartica (collection details as for Fig. 1). (Copyright: Barry Hammel)

L. E. Newton

Tamaulipas to Veracruz, Chiapas, and Quintana Roo), Central America (Guatemala, Belize, El Salvador, Honduras, Nicaragua, Costa Rica, Panama), Caribbean, N Colombia (Bolívar, Magdalena), Venezuela (widespread), Brazil (Minas Gerais, Bahia, Alagoas, Pernambuco, Piauí, Rio Grande do Norte, Sergipe); dry forests, 0–500 (–900) m. I: Nee (1993: 55); Machado (2009: as Apodanthera congestiflora); Paiva Lima (2010: 12, 23, 24, 86, as Apodanthera congestiflora); Belgrano & Pozner (2017: 35) – Figs. 1 and 2.  Corallocarpus emetocatharticus (Grosourdy) Cogniaux (1891); incl. Doyerea angosturensis Grosourdy (1864)  Corallocarpus angosturensis (Grosourdy) V. M. Badillo (1947); incl. Anguria glomerata Eggers (1879) (incorrect name, ICN Art. 11.4)  Corallocarpus glomeratus (Eggers) Cogniaux (1881); incl. Anguriopsis margaritensis J. R. Johnston (1905); incl. Apodanthera congestiflora Cogniaux (1916)  Melothria congestiflora (Cogniaux) Martinez Crovetto (1955); incl. Apodanthera trifoliata Cogniaux (1916)  Melothria trifoliata (Cogniaux) Martinez Crovetto (1954) (nom. illeg., ICN Art. 53.1). Description as for the genus.

Doyerea CUCURBITACEAE

References Belgrano, M. J. & Pozner, R. (2017) Sinopsis del género Apodanthera (Cucurbitaceae, Coniandreae). Darwiniana n.s. 5(1): 5–50, ills, key, maps. https://doi. org/10.14522/darwiniana.2017.51.716. Dieterle, J. V. A. (1976) Cucurbitaceae. In: Nash, D. L. (ed.): Flora of Guatemala. Fieldiana, Bot. 24(11: 4): 306–395, keys, ills. http://www.biodiversitylibrary.org/ page/2452031 Jeffrey, C. & Trujillo, B. (1992) Cucurbitaceae. In: Morillo, G. (ed.): Flora de Venezuela, Vol. V, Parte I; pp. 11–201, ills., keys. Caracas (VE): Fundación Instituto Botánico de Venezuela, Herbario Nacional de Venezuela/Fondo Editorial Acta Científica Venezolana. Lira, R. (2001) Familia Cucurbitaceae. In: Rzedowski, J. & Calderón de Rzedowski, G. (eds.): Flora del Bajío y de regiones adyacentes, Fasc. 92. Pátzcuaro (MX): Instituto de Ecología, Centro Regional del Bajío.

219 Machado, M. (2009) Apodanthera, caudiciform cucumbers of Bahía, Brazil. Cact. Succ. J. (US) 81(3): 147–149, ills. https://doi.org/10.2985/015.081.0309. McVaugh, R. (2001) Flora Novo-Galiciana. A descriptive account of the vascular plants of Western Mexico. Volume 3: Ochnaceae to Loasaceae. Ann Arbor (US): University of Michigan Herbarium. Mooney, H. A. [& al. 1992], Chu, C., Bullock, S. H. & Robichaux, R. (1992) Carbohydrate, water and nitrogen storage in vines of a tropical deciduous forest. Biotropica 24(2a): 134–139. https://doi.org/10.2307/2388666. Nee, M. (1993) Cucurbitaceae. In: Sousa, V. & GómezPompa, A. (eds.): Flora de Veracruz, fascículo 74. Xalapa (MX): Instituto de Ecología. https://doi.org/ 10.21829/fv.407.1993.74. Paiva Lima, L. F. (2010) Estudos taxonômicos e morfopolinicos em Cucurbitaceae brasileiras. Porto Alegre (BR): Universidade Federal do Rio Grande do Sul, Ph.D. thesis. http://www.lume.ufrgs.br/handle/ 10183/26294

Eureiandra CUCURBITACEAE L. E. Newton and G. N. Njoroge

Eureiandra Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 825, 1867). Type: Eureiandra formosa Hooker fil. – Coniandreae – Distr: Tropical & subtropical Africa, Yemen (Socotra). Etym: Application not explained but perhaps Gr. ‘eurys’, broad; and Gr. ‘aner, andros’, stamen; for the broad connective of the anthers mentioned in the protologue. Dioecious perennial climbers with tuberous Rstock to 20 cm ∅; stems herbaceous or woody; tendrils simple; L simple, entire or palmately 3- to 5-lobed; Fl cream to orange-yellow or rarely white, medium-sized to large, often on leafless shoots; male Fl solitary or in fascicles; hypanthium short, narrowly campanulate to funnel-shaped; Sep lanceolate, acute; Pet 5, free, to 3.5 cm, obovate, rounded-apiculate; St 5 or 3; Fil free or 2 pairs connate and 1 free; Anth all 1-thecous or 2 2-thecous and 1 1-thecous, thecae triplicate; female Fl solitary; similar to male flowers; staminodes 3 or 5; Sti 3-lobed; Fr large, to 13  8 cm, ovoid-ellipsoid to cylindrical, rostrate, indehiscent, fleshy, orange to red; Se many, ovate to 

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] G. N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya e-mail: [email protected]

globose, blackish, smooth or fibrillose, margin narrow or indistinct. Only 2 of the 8 species of the genus are here covered as being of interest to succulent plant collectors: E. fasciculata (Cogniaux) C. Jeffrey (Kew Bull. 15(3): 353, 1962). Type: Moçambique, Tete (Menyhart 931 [Z]). – Distr: SE Tanzania, Moçambique, Zimbabwe, Zambia, Malawi; woodland and wooded grassland, 10–1050 m. I: Jeffrey (1967: 42). – Fig. 1.  Momordica fasciculata Cogniaux (1897); incl. Coccinia petersii Gilg (1904); incl. Coccinia polyantha Gilg (1904). Climbers with tuberous Rstock; caudex 15 cm ∅,  10 cm below ground; stems herbaceous and pubescent when young, later gnarled and thickened, with pale papery bark; L petiole 0.7–2.8 cm, lamina ovate-cordate or triangularcordate in outline, 2.4–11.5  1.4–10 cm, palmate, with 3–5 lanceolate to elliptic lobes, rarely unlobed, bright olive-green, lower face paler and pubescent; Fl very variable in size, often appearing when the stems are almost leafless; male Fl 1–6 together; Ped 1–1.5 cm; hypanthium campanulate, expanding above, 2.5–7 mm; Pet obovate, 18–32  14–20 mm, pale creamyyellow or apricot-yellow, or sometimes white; St 3 or 5; female Fl on 0.4–1 cm peduncle; hypanthium 1.5–5 mm; Pet elliptic, 24–30  11–18 mm; staminodes 5; Fr ellipsoid, beaked, 5.7–6.2  2.4–2.5 cm; orange-red.

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PRE]). – Lit: Keraudren (1967: 67, 99–101, with ills.). Distr: Angola, Cameroon, Congo, Nigeria, Sudan, Democratic Republic of the Congo [Zaïre]; grassland (sometimes on ferrugineous crusts), forests and swampy areas. Incl. Eureiandra schweinfurthii Cogniaux (1881); incl. Adenopus diversifolius Cogniaux (1916); incl. Eureiandra congolensis Cogniaux (1916); incl. Eureiandra bequaertii De Wildeman (1922); incl. Eureiandra formosa var. hirtella Cogniaux (1924). Climbing herbs, loosely subtomentose; R becoming thickened, to 20 cm ∅; Br slender, slightly hirsute; tendrils simple; L palmately 3- to 5-lobed, 10–15 cm wide, lobes oblong-acuminate, pubescent on both faces; Fl golden-yellow; male Fl solitary or in fascicles 5 cm long; hypanthium subcylindrical, 12.5 mm; Cl 7.6–15.25 cm ∅; Pet lobes obovate; St 5; female Fl smaller than male flowers; Fr ovoid, (6–) 7–8 (–9)  2–4 cm, scarlet with white hairs in blotches.

References Fig. 1 Eureiandra fasciculata (Zimbabwe; near entrance gate to Gorongosa National Park). (Copyright: B. Wursten/ Online Flora of Zimbabwe)

E. formosa Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 825, 1867). Type: Angola, Cuanza Norte (Welwitsch 807 [LISU, COI, G-DC, K,

Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Keraudren, M. (1967) Cucurbitacées. In: Flore du Cameroun, vol. 6. Paris (FR): Muséum National d’Histoire Naturelle.

Gerrardanthus CUCURBITACEAE L. E. Newton

Gerrardanthus Harvey ex Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 840, 1867). Type: Gerrardanthus macrorhizus Hooker fil. – Zanonieae – Lit: Meeuse (1962: revision S Africa); Jeffrey (1967: Flora Tropical East Africa). Distr: Tropical & S Africa. Etym: Gr. ‘anthos’, flower; and for William Tyrer Gerrard (1831–1866), British naturalist and traveller, collecting in RSA and Madagascar in the 1860s. Incl. Atheranthera M. T. Masters (1871). Type: Atheranthera paniculata M. T. Masters. Dioecious climbers from large often partly exposed tubers or tuberous Rstocks; stems basally woody; tendrils apically forked; L petiolate, simple, ovate-cordate, unlobed or palmately 3- to 5-lobed; Fl small, yellowish to brownish-orange; male Fl few to many in panicles; hypanthium small, rotate; Sep 5, small and tooth-like; Cl rotate or widely campanulate; Pet 5, distinct, usually 2 slightly larger; St 5, usually 1 reduced to a staminode; Fil free; Anth in 2 pairs and one single, all 1-thecous, thecae straight, horizontal; female Fl solitary or few in small groups, similar to male flowers, with 5 staminodes; Ov trigonous, 3-locular near the tip and 1-locular at the base; Sty

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

3; Sti reniform, 2-lobed; Fr flask-shaped dry capsules dehiscing at the top, somewhat trigonous, pale yellow to brownish; Se several, fusiform, to 2 cm, (pale) brown, with membranous distal wing to 2 cm. Only 3 of the 5 species form succulent caudices (morphological origin not described but probably derived from the hypocotyl) as far as known. G. lobatus (Cogniaux) C. Jeffrey (Kew Bull. 15: 353, 1962). Type: Kenya (Scheffler 473 [B, K]). – Distr: Nigeria, Democratic Republic of the Congo [Zaïre], Uganda, Ethiopia, Kenya, Tanzania, Malawi, Moçambique; deciduous bushland and wooded grassland, 0–2000 m. I: Jeffrey (1967: 146), repeated by Jeffrey (1996: 19) and Schaefer & Renner (2011: 134); Lenain (2012: 144); Dortort (2018: 240). – Fig. 1.
Gerrardanthus grandiflorus var. lobatus Cogniaux (1916). Tuberous climbers; caudex exposed, domeshaped,  50 cm, greenish; stems to 6 m, semisucculent; L petiole 1.5–8 cm, lamina ovate to reniform-cordate, 3–10  4.4–12.4 cm, usually deeply 5-lobed, lobes triangular, shortly acuminate, slightly fleshy, somewhat glaucous, glabrous; male Fl few to 45 in 2–10 cm long panicles on 0.2–1.5 cm long peduncles; Ped 2–20 mm; hypanthium shallow, lobes 2–4 mm; Pet brownish-green or dull brownish-orange, unequal, 2 erect, oblanceolate, 10–18  2.5–5.5 mm, 3 reflexed, linear, paler, 9–16  1–2 mm; St 4, plus 1 subulate staminode; female

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Fig. 1 Gerrardanthus lobatus (without known wild origin; cult. Sukkulenten-Sammlung Zürich). (Copyright: U. Eggli)

Fl solitary or usually 2–3 in panicles on 1.5–3 cm long peduncles; Ped 1–8 mm; Pet unequal, 3 erect, 8–10  2–2.5 mm; Fr 4.5–6.3  1.8–2.8 cm, faces obscurely 2-veined. G. macrorhizus in the sense of Cogniaux (1916) belongs here. G. macrorhizus Harvey ex Hooker fil. (in Bentham & Hooker, Gen. Pl. 1: 840, 1867). Type: RSA (Gerrard s.n. [K]). – Distr: S Moçambique, E RSA (Eastern Cape, KwaZulu-Natal), Swaziland; lowland forests, 5–610 m. I: Rowley (1987: 58). Incl. Gerrardanthus portentosus Naudin ex Durand (1866); incl. Gerrardanthus megarhizus Decaisne & Harvey (1868). Tuberous climbers; caudex discoid, 30–60 cm, to 150 cm ∅; stems woody, to 1.6 m; L petiole 1.5–4 cm, lamina ovate to triangular, 3–8  3–8 cm,  angular or with 5–7 lobes, terminal lobe longest, tips subacute, glabrous; male Fl in short fascicles; Ped 5–30 mm; Sep oblong, 3 mm; Cl rotate,  12 mm ∅, brownish; female Fl solitary on 1–2 cm long peduncle; Fr obconical-subcylindrical, faintly angular, 5–6.5  1.5–2.2 cm, straw-coloured, smooth. G. tomentosus Harvey ex Hooker fil. (Curtis’s Bot. Mag. 109: t. 6694 þ text, 1883). Type: RSA,

KwaZulu-Natal (Wood 451 [K, NH]). – Lit: Crouch & al. (1999: with ills.); Crouch & al. (2003: with ills.). Distr: RSA (KwaZuluNatal); amongst rocks in semi-shade on steep bushy slopes. Tuberous climbers, caudex at soil-level, to 60 m ∅; stem to 15 m, greyish, furrowed, shortly and densely tomentose when young; tendrils long, branched, densely and shortly puberulous; L petiole 3–5 cm, lamina reniform-suborbicular, 6–12  6–12 cm, with 5–7 triangular or ovate lobes, upper face deep green, shortly and sparsely hairy, lower face greyish-green, densely tomentose, esp. on the prominent veins; male Inf to 10 cm, with flowers in bracteate fascicles at the nodes; female Inf a raceme with 3–4 flowers or flower solitary; all Fl 12–18 mm ∅, rotate; Pet brown, ovate-oblong; Fr narrowly bell-shaped, 10-ribbed, 6–7 cm. Very rare, with only 5 collections from 3 areas near Durban (Crouch & al. 1999).

References Cogniaux, A. (1916) Cucurbitaceae – Fevilleae et Melothrieae. In: Engler, A. (ed.): Das Pflanzenreich, IV. 275. I (Heft 66). Leipzig (DE): Verlag von Wilhelm Engelmann. http://bibdigital.rjb.csic.es/ idurl/1/10009

Gerrardanthus CUCURBITACEAE Crouch, N. R. [& al. 1999], Prentice, C., Smith, G. F. & Symmonds, R. (1999) South Africa’s rarest caudiciform cucurbit, Gerrardanthus tomentosus. Bradleya 17: 95–100, ills. https://doi.org/10.25223/brad.n17.1999.a6. Crouch, N. R. [& al. 2003], Smith, G. F., Symmonds, R. & Condy, G. (2003) Gerrardanthus tomentosus. Flow. Pl. Afr. 58: 134–142, ill., t. 2200, map. Dortort, F. (2018) Commiphoras, Cucurbits, Adenias and more – Kenya 2017. Cact. Succ. J. (US) 90(4): 238–257, ills. https://doi.org/10.2985/015.090.0403. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Jeffrey, C. (1996) Cucurbitaceae. In: Edwards, S. & al. (eds.): Flora of Ethiopia and Eritrea, 2(2): 17–59,

225 ills., keys. Addis Ababa (ET): National Herbarium/ Uppsala (SE): Department of Systematic Botany. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer. Meeuse, A. D. J. (1962) The Cucurbitaceae of southern Africa. Bothalia 8(1): 1–111, keys. https://doi.org/10. 4102/abc.v8i1.1611. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10.

Halosicyos CUCURBITACEAE L. E. Newton

Halosicyos Martinez Crovetto (Bol. Soc. Argent. Bot. 2: 84, 1947). Type: Halosicyos ragonesei Martinez Crovetto. – Coniandreae – Distr: C Argentina. Etym: Gr. ‘hals, halos’, salt; and Gr. ‘sicyos’, cucumber; for the preferred occurrence on saline soils and the placement in this family. Dioecious perennials with thick R to 15–20 mm ∅ terminating in globose tubers 2–6.5  2.5–8 cm, tubers juicy-fleshy; stems slender, slightly grooved, with white hairs, esp. at the nodes, or scabrous; tendrils simple, slender; L 0.3–0.5 cm petiolate, lamina suborbicular in outline, dissected into lobes 0.6–1  0.2 cm, hairy or scabrous; Fl small, greenish; male Fl 4–8 in 10–25 cm long racemes; Ped 1–2 mm; hypanthium subcampanulate, 3  2.5 mm, pilose in the throat; Pet 5, oblong-spatulate with rounded papillose tips, 3.5–4  1.8–2 mm; St 3; Fil distinct; Anth 3, 2 2-thecous, 1 1-thecous, thecae curved, coherent; female Fl solitary, slightly smaller than the male flowers; hypanthium cylindrical, 3  0.8–1 mm; Pet as in the male flowers; staminodes 5; Fr 7–8 mm long, laterally

compressed, 10–12 mm broad and 6–8 mm thick, tip with short beak, ripening red, glabrous; Se 4–8 per fruit, pear-shaped, 5–5.5  2  1.5 mm, sculptured to rugose, slightly winged. Ruiz Leal (1958) found the root tubers to be very juicy and fleshy, with a water content of 88% and a starch content of merely 5%. – A single species only: H. ragonesei Martinez Crovetto (Bol. Soc. Argent. Bot. 2: 87, ills., 1947). Type: Argentina, Córdoba (Ragonese 68399 [CORD]). – Lit: Ruiz Leal (1958: with ills.). Distr: C Argentina (Córdoba, Corrientes, La Rioja, San Luis, Mendoza); sandy soils in low bushland, often in saline conditions, at low altitudes. Description as for the genus.

References Ruiz Leal, A. (1958) Una Cucurbitácea nueva para la flora mendocina. Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 7(1): 49–61, ills. http://bdigital.uncu.edu.ar/7382.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_44

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Ibervillea CUCURBITACEAE L. E. Newton

Ibervillea Greene (Erythea 3: 75, 1895). Type: Sicydium lindheimeri A. Gray.
Coniandreae
Lit: Kearns (1994a: monograph); Lira & al. (2015: phylogeny). Distr: S USA, Mexico, C America. Etym: Unexplained, but perhaps for Pierre Le Moyne d’Iberville (1661–1706), French-Canadian soldier, captain, explorer, adventurer, trader and founder of the French colony of Louisiana. Incl. Sicydium A. Gray (1850) (nom. illeg., ICN Art. 53.1). Type: Sicydium lindheimeri A. Gray. Incl. Maximowiczia Cogniaux (1881) (nom. illeg., ICN Art. 53.1). Type: Sicydium lindheimeri A. Gray. Incl. Tumamoca Rose (1912). Type: Tumamoca macdougalii Rose. Incl. Dieterlea E. J. Lott (1986). Type: Dieterlea fusiformis E. J. Lott. Perennial dioecious climbers; R thick, forming nearly globose tubers, top usually exposed above ground; stems annual or perennial, slender; tendrils simple, small; L simple, with 3–5 lobes and several secondary lobes, glabrous or scabridpubescent; Fl small; Pet 5, white or yellowish,

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

densely pubescent inside, tips bifid (I. fusiformis) or entire; male Fl in short racemes or corymbs; hypanthium tubular to narrowly campanulate; St 3, from the mouth of the hypanthium tube; Fil connate into a central column; Anth 2 2-thecous, 1 1-thecous, thecae straight; female Fl solitary; hypanthium more slender than in male flowers; staminodes 3–5 or absent; Sti 3-lobed; Fr globose, ovoid or fusiform, 1.5–15  1.5–7 cm ∅, fleshy, indehiscent, red to yellowish; Se several to many, irregularly ovoid, scarcely compressed, transversely ridged or  smooth, margins raised.
Cytology: 2n ¼ 11, 12 (Darlington & Wylie 1955). Lira & al. (2015) presented a phylogeny of the genus based on molecular and morphological data. Dieterlea (monotypic, accepted as separate genus by Schaefer & Renner (2011b), but included in Ibervillea by Schaefer & Renner 2011a), is firmly embedded in Ibervillea (first shown in the molecular phylogeny of Kocyan & al. 2007), which is shown as sister to Tumamoca. Ibervillea s.s. falls in 2 unnamed clades in this analysis. On the basis of their molecular data, Lira & al. (2015) propose to include the monotypic Tumamoca into a broadened Ibervillea as the two genera only differ in a single character (Tumamoca monoecious, Ibervillea s.s. dioecious) which is known to be labile in tribe Coniandreae, and Nesom (2015: 30) specifically mentions that I. macdougalii (as Tumamoca) can apparently be monoecious or dioecious depending on rainfall and plant size. In this wide

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_45

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circumscription, Ibervillea has 9 species, all covered below. I. fusiformis (E. J. Lott) Kearns (Madroño 41(1): 15, 1994). Type: Mexico, Jalisco (Magallanes 4252 [MEXU, CAS, ENCB, MICH, MO, NY]).
Lit: Lott (1986: protologue Dieterlea, with ills.); McVaugh (2001: 548–551, with ills.). Distr: W-C Mexico (Sonora, Sinaloa, Colima, Jalisco, Michoacán); tropical deciduous forests. I: Lott (1986: as Dieterlea); Lira & al. (2015: 201).
Figs. 1 and 2.  Dieterlea fusiformis E. J. Lott (1986). Tuberous climbers, dioecious; caudex to 40 cm ∅, pale brown; stems perennial, coarse; tendrils small; L petiole 1.5–5 cm, lamina orbicular-ovate, 6–15  7–17 cm, deeply 3- to 5-lobate, lobes obtuse to acute, with trichomes usually having a conspicuous cystolithic base, surface rough; Fl nocturnal, fragrant; hypanthium narrowly cylindrical, glabrous outside, densely villous inside in the throat; Pet white or yellow-white, tips bifid; male Fl 10–12 in 8–15 cm long racemes; Ped Fig. 1 Ibervillea fusiformis (thickened rootstock; Mexico; Sonora, Álamos, NW of Yocogigua). (Copyright: T. R. Van Devender)

L. E. Newton

4–5 mm; hypanthium 3–4.5 cm; Pet 1.5–2.5  1–1.5 cm, free, glandular-puberulent; St 3, female Fl solitary, axillary, 9–10 cm long; Ped 4–20 mm; hypanthium 5 cm; Pet 3 cm; staminodes 5; Fr indehiscent, fusiform, rostrate, with 10–11 longitudinal ridges, 9–12  3–4.5 cm, glabrous, orange-red at maturity. I. guatemalensis (Standley & Steyermark) Kearns (Madroño 41(1): 17, 1994). Type: Guatemala, Zacapa (Standley 74623 [F, K [photo]]).
Distr: S Mexico (Chiapas), Guatemala (Chiquimula, El Progreso, Zacapa), Honduras; low-growing deciduous forests, 200–400 (
1300) m.  Corallocarpus guatemalensis Standley & Steyermark (1944). Vigorous creepers to climbers, likely monoecious, with long horizontally oriented Rstock; stems thickened, strigillose when young, terete, becoming woody; tendrils simple; L petiole 3–7.5 cm, hairy, lamina 6.5–8  10.5–12.5 cm, broadly ovate-cordate to subreniform, 3- to

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Mexico (Nayarit, Jalisco, Colima, Michoacán, Guerrero, Oaxaca, Sinaloa?) to El Salvador;  Corallocarpus hypoleucus Standley (1937). Climbers with thin somewhat angled stems, glabrous; L petiole 2–3.5 cm, sparsely minutely tomentellous or glabrous, lamina thickishcoriaceous, 7–13  8–15 cm, circular to deltoidcircular in outline, 3-lobed to the middle, base deeply cordate, lobes broadly to narrowly circular, acuminate, terminal lobe entire, lateral lobes roundedly auriculate, upper face glabrous, lower face densely grey- to white-tomentose with appressed hairs; Inf and Fl not described; Fr globose-ovoid, 6  5 cm, apex obtuse to rounded and umbonate with persistent style, glabrous, colour not described; Se not known.

Fig. 2 Ibervillea fusiformis (leaf and unripe fruit; Mexico; Sonora, Álamos, SSE of Álamos). (Copyright: A. L. Reina-Guerrero)

5-lobed, somewhat coriaceous, both faces densely appressed-tomentose, margins scarcely minutely denticulate; male Inf congested racemes, 1–1.8 cm pedunculate; male Fl Ped  1 mm, pilose; hypanthium 2–6 mm, campanulate; Sep lobes 1–2 mm, ovate-triangular; Pet 3–5  1.5–4 mm, ovate-triangular, both faces pilose; femal Fl, Fr and Se unknown. Considered as a synonym of Doyerea emetocathartica by Dieterle (1976) but this has been found to be incorrect by Jeffrey (1978) and Kearns (1994a: 18). Poorly known but very similar to and closely related if not conspecific with I. hypoleuca (Lira & al. 2015). I. hypoleuca (Standley) C. Jeffrey (Kew Bull. 33(2): 349, 1978). Type: Mexico, Guerrero (MacDaniels 266 [F, K [photo]]).
Distr:

I. lindheimeri (A. Gray) Greene (Erythea 3: 75, 1895). Type: USA, Texas (Lindheimer 612 [GH, K, MO]).
Distr: S USA (Texas), Mexico (widespread from Durango and Nuevo León to Yucatán); dry scrub to low-growing (sub-) deciduous forests, 0–200 m.  Sicydium lindheimeri A. Gray (1850) (incorrect name, ICN Art. 11.4)  Maximowiczia lindheimeri (A. Gray) Cogniaux (1881) (incorrect name, ICN Art. 11.4); incl. Ibervillea convolvulacea hort. (s.a.) (nom. inval., ICN Art. 29.1); incl. Bryonia abyssinica Gouault (1853) (nom. illeg., ICN Art. 53.1); incl. Sicydium tenellum Naudin (1862) (incorrect name, ICN Art. 11.4)  Maximowiczia tripartita var. tenella (Naudin) Cogniaux (1881) (incorrect name, ICN Art. 11.4)  Ibervillea tenella (Naudin) Small (1903); incl. Sicydium tripartitum Naudin (1862) (incorrect name, ICN Art. 11.4)  Maximowiczia tripartita (Naudin) Cogniaux (1881) (incorrect name, ICN Art. 11.4)  Ibervillea tripartita (Naudin) Greene (1895). Dioecious; R large and fleshy; stems thin, glabrous, minutely striate, procumbent or climbing; tendrils very small, divided; L petiole 1–3.5 cm, lamina succulent, palmate, 2.5–7.5 (
12) cm ∅, 3- to 5-partite, again divided, secondary lobes linear, coarsely dentate, upper face glabrous, lower face pockmarked, hirsute; Fl yellow or geenish, diurnal, 8.3–12.5 mm long; male Fl funnel-shaped; Pet oblong, narrowed below,

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Fig. 3 Ibervillea macdougalii (USA; Arizona, Tucson Mts. W of Tucson, road to Gates Pass). (Copyright: F. W. Reichenbacher)

obscurely ciliate; female Fl with Cal tube constricted above the ovary and then prolonged into a beak and then funnel-shaped; Pet as in male flowers; Fr globose, 1.5–3  2.5 cm, reddish; Se roundish-oval, 6.25 mm, turgidly lenticular. Material usually referred to as I. tripartita from the Yucatán peninsula and Oaxaca probably represents a new species (Lira & Monro 2009: 25). I. macdougalii (Rose) Lira & al. (Phytotaxa 201(3): 204, 2015). Type: USA, Arizona (MacDougal s.n. [US 591589]).
Lit: Turner & al. (1995: 392–393, map, as Tumamoca). Distr: SW USA (S-C Arizona), NW Mexico (Sonora, Baja California?, N Sinaloa?, Zacatecas?); dry soil among rocks in desert scrub or grassland, to 700 (
1000) m. I: Kearns (1994b: 25); Nesom (2015: 29); both as Tumamoca.
Fig. 3.  Tumamoca macdougalii Rose (1912); incl. Tumamoca mucronata Kearns (1994). Dioecious or monoecious perennials with clusters of R tubers 5–15 cm long, with a persistent woody stem to 15 cm; annual stems slender, to 1.5 m, glabrous; L petiole 1–2 cm, lamina simple, to 4.5 cm, with 3 linear to linear-lanceolate lobes each 2–4  0.2–1 cm, upper face glabrous, lower facee with hairs to 0.1 mm with multicellular bases; Fl diurnal; male Fl 2–19 in 3.5–10 cm long racemes with 1–2.5 cm long peduncle; hypanthium cylindrical, 5–9 mm; Pet 4–4.5 mm,

pale yellow or greenish-yellow; female Fl solitary, with peduncle 0.5–1.5 cm; hypanthium cylindrical, 5.5–7.5 mm; Pet 5–6 mm, pale yellow or greenish-yellow; Fr globose, fleshy, 0.8–1 cm ∅, red, rarely yellow; Se 2 to several per fruit, ovoid, tuberculate-rugose, 7–8 mm. Formerly considered as a species of the separate genus Tumamoca, but the alleged differences between the two genera are minimal (Lira & al. 2015). These authors also reduced the second species of former Tumamoca, T. mucronata, to a synonym here as the alleged differences between the two taxa are continuous. The inclusion of T. mucronata (from Zacatecas, Mexico) here results in a very discontinuous range, which calls for further investigations.
Vernacular name: “Tumamoc Globeberry”. Plants can be monoecious or appear to be dioecious, but according to Nesom (2015: 30) plants often produce only male flowers early in the season, and female flowers follow later in response to rain, and small plants may not produce female flowers at all. I. maxima Lira & Kearns (Sida 14(2): 223–226, ills., 1990). Type: Mexico, Nayarit (Kearns & Kearns 390 [MEXU, BM, BRIT, CHAPA, CAS, F, GUAD, IBUG, K, MICH, MO, NY, TEX, UC, US, XAL]).
Lit: Lira (2001: 65–67, with ills.); McVaugh (2001:

Ibervillea CUCURBITACEAE

551–554, with ills., as Dieterlea). Distr: Mexico (Sinaloa, Nayarit, Jalisco, Michoacán, Guerrero); tropical deciduous and semi-deciduous forests, on dry slopes and lava fields, 140–1200 (
1850) m.  Dieterlea maxima (Lira & Kearns) McVaugh (2001). Climbing deciduous perennials with branched fleshy tuberous Rstock; stems perennial, to 12 m, terete, becoming glabrous; tendrils simple; L petiole 3–7.5 cm, lamina ovate-cordate to subreniform, 9.5–15  12.5–20 cm, slightly 3-lobed, surface hispid-scabrous, denser on the lower face, margin obscurely and sparsely denticulate; male Fl 4–10 in shortened racemes; Ped 10–25 mm; hypanthium cylindrical, 11–18 mm, densely pubescent; Pet yellow with greenish centre; St 3; female Fl solitary, similar to male flowers; Fr ellipsoid berries with short beak, 13–15   6 cm, glabrous, bright orange. I. millspaughii (Cogniaux) C. Jeffrey (Kew Bull. 33(2): 348–349, 1978). Type: Mexico, Yucatán (Gaumer 842 [F, NY]).
Distr: E & S Mexico (Tamaulipas, Veracruz, Oaxaca, Chiapas, Campeche, Yucatán, Quintana Roo), Belize, Guatemala?; (sub-) deciduous to semi-evergreen forests, 0–10 m.  Corallocarpus millspaughii Cogniaux (1896). Vigorous climbers or creepers; stems to 15 m and more, somewhat succulent, glabrous, terete (but becoming angled when dry); tendrils enlarged; L petiole 2.6–4.5 (
5.3) cm, glabrous, lamina 4.5–9  5–14 cm, narrowly ovate-cordate, coriaceous, entire or 3- (rarely 5-) lobed, lobes acute, obtuse or rounded, both faces glabrous and pustulate, margins entire to inconspicuously dentate; male Fl in short 5- to 10- (to 12-) flowered racemes or axillary fascicles, 8–11 mm pedunculate; Ped 5–10 mm, sparsely glandularpubescent; hypanthium 4–7 mm, cylindricalcampanulate with globose base, outer face puberulous; Sep lobes 10–15 mm, triangular, puberulous; Pet 5–7 mm, obovate, inner face papillose-puberulous, outer face glandularpuberulous; female Fl solitary; Ped 14 mm, otherwise similar to male flowers; Ov oblong-ovoid, glabrescent; Fr 2–4  2.5–3 cm, globose to

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ellipsoid or ovoid, green with whitish blotches turning yellow and finally orange to red, with stiff pericarp when dry; Se 7–9  5–6 mm. Similar to and perhaps conspecific with I. lindheimeri according to Jeffrey (1978). Considered as a synonym of Doyerea emetocathartica by Dieterle (1976) but this has been found to be incorrect by Jeffrey (1978) and Kearns (1994a: 18). I. sonorae (S. Watson) Greene (Erythea 3: 75, 1895). Type: Mexico, Sonora (Palmer 283 [GH, K, NY, US]).
Lit: Wiggins (1980: 390–391, with ills.). Distr: NW Mexico (Baja California, Baja California Sur, Sonora, Sinaloa, and offshore islands); foothill valleys and desert plains, in sandy to gravelly or rocky ground, to 300 m. I: Rowley (1987: 214); Lenain (2012: 148–149).
Fig. 4.  Maximowiczia sonorae S. Watson (1889) (incorrect name, ICN Art. 11.4); incl. Ibervillea

Fig. 4 Ibervillea sonorae (Supthut s.n.: Mexico; Baja California Sur, 22 km S of San Pedro; cult. SukkulentenSammlung Zürich). (Copyright: U. Eggli)

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magdalensis Hort. Christa’s (s.a.) (nom. inval., ICN Art. 29.1); incl. Maximowiczia insularis Brandegee (1916) (incorrect name, ICN Art. 11.4)  Ibervillea insularis (Brandegee) Wiggins (1980); incl. Maximowiczia sonorae var. brevicaulis I. M. Johnston (1924) (incorrect name, ICN Art. 11.4); incl. Maximowiczia sonorae var. peninsularis I. M. Johnston (1924) (incorrect name, ICN Art. 11.4)  Ibervillea sonorae var. peninsularis (I. M. Johnston) Wiggins (1980); incl. Ibervillea guarequi MacDougal (1931) (nom. inval., Art. 36.1). Perennial climbers; R tuberous, bottle-shaped, partly exposed above ground; stems to 3 m, slender, glabrous, with short internodes; L petiole  as long as the lamina, lamina circular to ovate in outline, 4–10 cm, with 3 primary lobes and 7–9 secondary lobes, glaucous and glabrous, sometimes with coarse white hairs on the lower face; Fl diurnal; male Fl in 15 cm long racemes; hypanthium tubular, glabrous; Pet 4–5 mm, villous, yellow; female Fl solitary; Fr globose or ovoid, 3–5 cm, reddish; Se irregularly ovoid, flattened, angles raised as low narrow ridges. I. tenuisecta (A. Gray) Small (Fl. Southeast. US, 1136, 1138, 1903). Type [lecto]: USA, New Mexico (Wright 221 [GH, K]).
Distr: S USA (Arizona, New Mexico, Texas), Mexico (Chihuahua, Coahuila, Durango, Nuevo León?, Zacatecas?, San Luis Potosí?); desert flats and grassy plains. I: Beyenbach (2004).  Sicydium lindheimeri var. tenuisectum A. Gray (1852) (incorrect name, ICN Art. 11.4)  Maximowiczia lindheimeri var. tenuisecta (A. Gray) Cogniaux (1881) (incorrect name, ICN Art. 11.4)  Maximowiczia tenuisecta (A. Gray) Cogniaux (1881) (incorrect name, ICN Art. 11.4)  Ibervillea lindheimeri var. tenuisecta (A. Gray) M. C. Johnston (1963). Climbers with thick nearly globose R tuber; stems slender, glabrous, finely striate; tendrils very small, jointed 1–3 mm above the enlarged base; L petiole 1–2 cm, lamina to 6 cm wide, dissected to almost the base with 3–5 linear to filiform lobes, irregularly lobulate, upper face glabrous, lower face pustulate-hispid; Fl diurnal; male Fl in racemes; hypanthium campanulate;

L. E. Newton

Pet spreading, 3–4 mm, yellow; female Fl solitary, small; Pet puberulent; Fr globose to ovoidglobose, 1.5–3 cm, reddish to bright red. Lira & al. (2015) chose to place this species as variety of I. lindheimeri, reverting to the original classification of the taxon under Sicydium lindheimeri, but the evidence from their moleculuar study is not compelling, as I. lindheimeri, I. tenuisecta and I. millspaughii (upheld at species rank) form an unresolved clade.

References Beyenbach, J. (2004) Ibervillea tenuisecta (A. Gray) Small. Avonia 22(2): 31–33, ills. Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Dieterle, J. V. A. (1976) Cucurbitaceae. In: Nash, D. L. (ed.): Flora of Guatemala. Fieldiana, Bot. 24(11: 4): 306–395, keys, ills. http://www.biodiversitylibrary.org/ page/2452031 Jeffrey, C. (1978) Further notes on Cucurbitaceae: IV. Some New World taxa. Kew Bull. 33(2): 345–380. https://doi.org/10.2307/4109590. Kearns, D. M. (1994a) The genus Ibervillea (Cucurbitaceae): An enumeration of the species and two new combinations. Madroño 41(1): 13–22, key. http://www.jstor.org/stable/41426234 Kearns, D. M. (1994b) A revision of Tumamoca (Cucurbitaceae). Madroño 41(1): 23–29, ills., key. http://www.biodiversitylibrary.org/page/47944598 Kocyan, A. [& al. 2007], Zhang, L.-B., Schaefer, H. & Renner, S. S. (2007) A multi-locus chloroplast phylogeny for the Cucurbitaceae and its implications for character evolution and classification. Molec. Phylogen. Evol. 44(2): 553–577. https://doi.org/10. 1016/j.ympev.2006.12.022. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer. Lira, R. (2001) Familia Cucurbitaceae. In: Rzedowski, J. & Calderón de Rzedowski, G. (eds.): Flora del Bajío y de regiones adyacentes, Fasc. 92. Pátzcuaro (MX): Instituto de Ecología, Centro Regional del Bajío. Lira, R. & Monro, A. K. (2009) Cucurbitaceae. In: Davidse, G. & al. (eds.): Flora Mesoamericana. Volumen 4, parte 1: Cucurbitaceae a Polemoniaceae; pp. 1–49, ills. México, D.F. (MX) / St. Louis (US) / London (GB): Universidad Nacional Autónoma de México / Missouri Botanical Garden / Natural History Museum (London). http://www.tropicos.org/Project/FM Lira, R. [& al. 2015], Sosa, V., Legaspi, T. & Dávila, P. (2015) Phylogenetic relationships of Ibervillea and Tumamoca (Coniandreae, Cucurbitaceae), two genera of the dry lands of North America. Phytotaxa 201(3): 197–206, ills. https://doi.org/10.11646/phytotaxa.201.3.3.

Ibervillea CUCURBITACEAE Lott, E. J. (1986) Dieterlea, a new genus of Cucurbitaceae from Mexico. Brittonia 38(4): 407-410, ills. https://doi. org/10.2307/2807088. McVaugh, R. (2001) Flora Novo-Galiciana. A descriptive account of the vascular plants of Western Mexico. Volume 3: Ochnaceae to Loasaceae. Ann Arbor (US): University of Michigan Herbarium. Nesom, G. L. (2015) Cucurbitaceae. In: FNA Editorial Committee: Flora of North America north of Mexico, vol. 6: 3–58, ills., keys. New York (US) / Oxford (GB): Oxford University Press. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press.

235 Schaefer, H. & Renner, S. S. (2011a) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi. org/10.1002/tax.601011. Schaefer, H. & Renner, S. S. (2011b) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Turner, R. M. [& al. 1995], Bowers, J. E. & Burgess, T. L. (1995) Sonoran Desert plants. An ecological atlas. Tucson (US): University of Arizona Press. Wiggins, I. L. (1980) Flora of Baja California. Stanford (US): Stanford University Press.

Kedrostis CUCURBITACEAE L. E. Newton and G. N. Njoroge

Kedrostis Medikus (Philos. Bot. 2: 69, 1791). Type: Bryonia africana Linné. – Coniandreae – Distr: Tropical & subtropical Africa, Arabia, Madagascar, Asia. Etym: Gr. ‘kedrostis’, White Bryony, i.e. the ancient name of a scrambling cucurbit, used as generic name for a different group of plants by Medikus. Incl. Coniandra Schrader ex Ecklon & Zeyher (1836). Type: not designated. Incl. Cyrtonema Schrader ex Ecklon & Zeyher (1836). Type: not designated. Incl. Rhynchocarpa Schrader ex Endlicher (1839). Type: Trichosanthes foetidissima Jacquin. Incl. Achmandra Arnott (1840). Type: Bryonia rostrata Rottler [lectotype, designated by Pfeiffer, Nomencl. Bot. 1: 60, 1871, sub Aechmandra]. Incl. Bryonopsis Arnott (1840) (nom. illeg., ICN Art. 53.1). Type: Bryonopsis courtallenis Arnott. Incl. Aechmandra Arnott (1841) (nom. inval., ICN Art. 61.1). Type: Bryonia rostrata Rottler

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] G. N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya e-mail: [email protected]

[lectotype, designated by Pfeiffer, Nomencl. Bot. 1: 60, 1871]. Incl. Pisosperma Sonder (1862). Type: Pisosperma capense Sonder. Incl. Cerasiocarpum Hooker fil. (1867). Type: Aechmandra zeylanica Thwaites. Incl. Toxanthera Hooker fil. (1883). Type: Toxanthera natalensis Hooker fil. Climbing or trailing perennial herbs, rarely woody or subshrubs, often with tuberous Rstock, monoecious, or only rarely dioecious; stems climbing or creeping, glabrous or hirsute, sometimes rough, sometimes basally thickened and succulent, or tuberous herb with creeping underground branches with subsidiary tubers (K. psammophila); tendrils simple, sometimes forked, rarely absent (K. psammophila); L usually simple, ovate-lanceolate or hastate to suborbicular in outline,  palmately lobed, rarely with 3 leaflets; Fl small to large, rotate, greenish-white, greenishyellow or yellow; male Fl few to many, in pedunculate racemes; hypanthium shortly campanulate; Sep small, lanceolate; Pet 1.5–8 mm, ovate, united at the base; St 5, sometimes 2 pairs united, giving the appearance of a total of 3 stamens; Anth all 1-thecous or 2 2-thecous and 1 1-thecous, thecae  straight; female Fl solitary, paired or few in fascicles, similar to male flowers (but hypanthium narrowly cylindrical, 35–70  2–3 mm and Pet to 12 mm in K. psammophila); staminodes (3–) 5; Ov ovoid, smooth or finely papillate; Sty slender; Sti 2- (to 3-)fid; Fr subglobose to ovoid-

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_46

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rostrate, or conical or fusiform, to 9  3 cm, fleshy, sometimes hairy, orange to red at maturity (white and subterranean in K. psammophila), indehiscent or dehiscent by a longitudinal slit or valves; Se 1–10 per fruit, asymmetrically pearshaped to globose, small, smooth. A genus of 20 (Schaefer & Renner 2011b) to 28 species (Schaefer & Renner 2011a). In addition to those described below, some others have underground tubers and could possibly be grown as caudiciform plants, with the tuber at soil level. K. africana (Linné) Cogniaux (in A. & C. de Candolle, Monogr. Phan. 3: 643, 1881). Type: [icono]: Herman, Parad. Bat., t. 23, 1698. – Distr: Namibia, RSA (widespread throughout the country); grassland, vleis, stream banks and disturbed places, 190–2115 m. I: Rowley (1987: 57); Eggli (2008: 283); Lenain (2012: 162–163). – Figs. 1 and 2.  Bryonia africana Linné (1753)  Coniandra africana (Linné) Sonder (1862)  Rhynchocarpa africana (Linné) Ascherson (1867); incl. Bryonia ciliata Moench (1794); incl. Bryonia dissecta Thunberg (1794)  Corallocarpus dissectus (Thunberg) Hort. Kirstenbosch (s.a.) (nom. inval., ICN Art. 29.1)  Coniandra dissecta (Thunberg) Schrader Fig. 1 Kedrostis africana. (Copyright: E. Bihrmann)

L. E. Newton and G. N. Njoroge

(1836)  Rhynchocarpa dissecta (Thunberg) Naudin (1861)  Coniandra glauca var. dissecta (Thunberg) Sonder (1862)  Kedrostis glauca var. dissecta (Thunberg) Cogniaux (1881); incl. Bryonia pinnatifida Burchell (1822); incl. Coniandra glauca Schrader (1834)  Kedrostis glauca (Schrader) Cogniaux (1881); incl. Coniandra pinnatisecta Schrader (1836); incl. Cyrtonema digitatum Schrader ex Ecklon & Zeyher (1836); incl. Coniandra grossulariifolia E. Meyer ex Arnott (1841); incl. Bryonia multifida E. Meyer ex Drège (1843) (nom. inval., ICN Art. 38.1a); incl. Bryonia grossulariifolia E. Meyer ex Drege (1844) (nom. inval., ICN Art. 38.1a); incl. Kedrostis grossulariifolia C. Presl (1845); incl. Coniandra punctulata Sonder (1862)  Kedrostis punctulata (Sonder) Cogniaux (1881). Climbing or trailing, tuberous, monoecious; caudex to 50 cm, simple or branched, white or sometimes turning green when exposed; stems annual, to 6 m, much branched, glabrous or nearly so; tendrils filiform; L petiole 0.4–1.2 cm, sparsely hirsute, lamina triangular in outline, 6–10 cm, with 3–5 laciniate lobes, surface slightly rough; male Fl few to 12 in short racemes on 2–8 cm long peduncles; Ped 0.2–0.5 cm; hypanthium 0.5–1.5 mm; Pet 1–2 mm, finely papillose, yellowish-green, whitish or cream-coloured;

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or linear pointed lobes and scattered teeth or further small lobes, grey-green, surface with short tufted hairs; male Fl few or up to 20 in racemes to 6 cm; Ped to 3 cm; hypanthium 3–6 mm, shortly hairy; Pet to 8 mm, pale yellow or greenish-yellow, hairy; female Fl solitary, few or many, similar to male flowers but Ped to 1 cm; staminodes 3–5; Fr  globose, shortly beaked, 1.5–3 cm ∅; Se up to 10.

Fig. 2 Kedrostis africana. (Copyright: E. Bihrmann)

female Fl solitary or few in fascicles; Pet yellowish-green, whitish or cream-coloured; Fr subglobose, 0.8–1.5 cm ∅, orange-red; Se few. A hybrid with Zehneria scabra (as Melothria punctata) has been reported by England (1990). The tuber is used as emetic, purgative and diuretic, to treat dropsy, and against obesity, and aqueous extracts have been found to be safe to use (Unuofin & al. 2018). K. capensis (Sonder) A. Meeuse (Bothalia 8(1): 28, 1962). Type [syn]: RSA, Free State (Zeyher 593 [K]). – Distr: S Namibia (Karas), Lesotho, RSA (Free State, Northern Cape, Western Cape); grassland and thickets, disturbed places, 10–1830 m.  Pisosperma capense Sonder (1862). Tuberous, monoecious; caudex subglobose or napiform, to 10 cm ∅; stems annual, erect becoming prostrate, to 50 cm, branching; Br slender, to 30 cm, angular, striate, roughly hairy, rarely glabrous; tendrils short or absent; L petiole 0.4–1.5 cm, lamina oblong to ovate-oblong in outline, to 7  5 cm, with 5–7 oblong lanceolate

K. crassirostrata Bremekamp (Ann. Transvaal Mus. 15: 260, 1933). Type: RSA, Limpopo (Bremekamp & Schweickerdt 40 [PRE]). – Distr: Namibia, Botswana, RSA (Limpopo, North-West Prov., Northern Cape, Gauteng); dry sandy areas, 1020–1120 m. I: Ettelt (2018: 256). Climbing, tuberous, monoecious; stems annual, to 2 m, slender, furrowed; tendrils simple; L suborbicular in outline, 2.5–7  2.5–7 cm, digitately 3- to 5-lobed, segments pinnatifid, sessile, with white spots on the upper face, slightly fleshy; male Fl in short racemes on 0.5 cm long peduncles; hypanthium campanulate,  1 mm; Pet  1.5 mm, outer surface papillose-hairy; female Fl solitary, subsessile, similar to male flowers; Fr ovoid or pear-shaped, beaked, 1.4–1.7 cm, red; Se 1–5. K. foetidissima (Jacquin) Cogniaux (in A. & C. de Candolle, Monogr. Phan. 3: 634, 1881). Type: [icono:] Jacquin, Collectanea, t. 624, 1790. – Distr: Tropical Africa ro RSA (widespread), tropical Asia through India to Burma; rain forest margins, deciduous and semi-evergreen woodland and bushland, wooded grassland, 0–1830 m.  Trichosanthes foetidissima Jacquin (1789)  Bryonia foetidissima (Jacquin) Schumacher & Thonning (1827)  Rhynchocarpa foetidissima (Jacquin) Walpers (1843)  Melothria foetidissima (Jacquin) Roberty (1954); incl. Melothria foetida Desrousseaux (1797); incl. Bryonia rostrata Rottler (1803)  Aechmandra rostrata (Rottler) Arnott (1841)  Rhynchocarpa rostrata (Rottler) Naudin (1862)  Kedrostis rostrata (Rottler) Cogniaux (1881); incl. Bryonia perottetiana Seringe (1828)  Kedrostis foetidissima var. perottetiana (Seringe) Cogniaux

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(1898); incl. Bryonia pilosa Roxburgh (1832); incl. Rhynchocarpa foetida Schrader (1838) (nom. illeg., ICN Art. 52.1); incl. Cyrtonema convolvulaceum Hochstetter (1842); incl. Bryonia obtusiloba E. Meyer (1843); incl. Cyrtonema divergens A. Richard (1847)  Kedrostis foetidissima var. divergens (A. Richard) Cogniaux (1898); incl. Bryonia prostrata D. Dietrich (1852); incl. Zehneria obtusiloba Sonder (1862)  Kedrostis obtusiloba (Sonder) Cogniaux (1916)  Kedrostis foetidissima ssp. obtusiloba (Sonder) A. Meeuse (1962); incl. Cyrtonema foetens Hochstetter ex Hooker fil. (1871); incl. Kedrostis foetidissima var. microcarpa Cogniaux (1881); incl. Melothria obtusiloba Spruce ex Cogniaux (1881); incl. Kedrostis foetidissima var. glandulifera A. Zimmermann (1922). Perennial herbs from a tuberous Rstock; stems glabrous to densely hairy, often glandular, climbing or trailing to 2 m, becoming thickened at the base when older; tendrils simple; L petiole 0.5–5 cm, lamina sagittate, hastate, deltoid, orbicular-cordate, ovate-cordate or pentagonal,  scabrid, hairy (sometimes densely, esp. beneath), 1.9–9.3  1.3–9.2 cm, entire or palmately 3- (to 5-) lobed, lobes subtriangular, not or rarely narrowed below; male Fl 4–11 (rarely fewer) in lax fascicles; peduncle filiform, 2–25 mm; Ped filiform, 2–10 mm; hypanthium tube campanulate, simply hairy or glandularhairy,  2 mm, lobes filiform, 1–2 mm; Pet yellow, 3.5–4  1.5–2 mm; St 3; female Fl solitary or paired on stalks 2–5 mm long, sometimes coaxillary with male flowers; hypanthium tube campanulate, simply hairy or glandular-hairy,  2 mm; lobes subulate, 1–2 mm; Pet  4–5  2 mm; Ov beaked, 7–9  2.5–3 mm, densely glandular-hairy; Fr on 1–14 mm long stalks, pear-shaped, usually  beaked, hairy, scarlet, 0.9–2.7  0.6–1.3 cm; Se 4–6  2–4  1.5–2 mm, pear-shaped, subcompressed and narrower at the apex, inflated below, smooth, when fresh with narrow gelatinous marginal wing. – [L. E. Newton]. K. leloja (J. F. Gmelin) C. Jeffrey (Kew Bull. 15(3): 354, 1962). Type: Yemen (Forsskål

L. E. Newton and G. N. Njoroge

s.n. [C]). – Distr: Yemen, Somalia, Kenya, Tanzania; deciduous bushland and wooded grassland, 10–550 m. I: Jeffrey (1967: 132); Wilkins (1994: 69); Eggli (2008: 283); all as K. hirtella; Lenain (2012: 164–165).  Turia leloja J. F. Gmelin (1791); incl. Cyrtonema hirtellum Hochstetter ex Neudin (1862); incl. Rhynchocarpa hirtella Naudin (1862)  Kedrostis hirtella (Naudin) Cogniaux (1881); incl. Toxanthera natalensis Hooker fil. (1883)  Kedrostis natalensis (Hooker fil.) A. Meeuse (1962); incl. Kedrostis longipedunculata Cogniaux (1895); incl. Toxanthera kwebensis N. E. Brown (1909); incl. Toxanthera lugardiae N. E. Brown (1909); incl. Kedrostis rautanenii Cogniaux (1910); incl. Kedrostis eminens Cogniaux (1916); incl. Kedrostis gilgiana Cogniaux (1916); incl. Kedrostis ledermannii Cogniaux (1916); incl. Kedrostis rigidiuscula Cogniaux (1916); incl. Kedrostis abdallai A. Zimmermann (1922); incl. Kedrostis cufodontii Chiovenda (1934). Climbing or trailing, monoecious, with a tuberous Rstock and additional R tubers from adventitious roots of stem branches; stems annual, to 2 m, sparsely hairy, becoming glabrous; tendrils simple or bifid; L petiole 1.5–6.5 cm, lamina ovate or reniform-cordate in outline, 0.6–10  3.5–13 cm, usually palmately 3- to 5-lobed, lobes obovate or ovate-triangular, softly hairy to scabrid-hairy; male Fl 6–29 in lax 3–9 cm long racemes on peduncles 1.6–12.6 cm long; Ped 0.2–0.8 cm; hypanthium campanulate, 1–3 mm; Pet 1.5–6 mm, greenish, pale greenish-white or greenish-yellow; St 5, included; female Fl solitary; Ped 0.15–3 cm; hypanthium campanulate, 1–2 mm; Pet 2–6 mm; Fr ovoid, cylindrical, conical or fusiform, tip often with long beak, 3.8–9  1.5–2.6 cm, dehiscing by a longitudinal slit, glabrous, orange. K. nana (Lamarck) Cogniaux (in A. & C. de Candolle, Monogr. Phan. 3: 637, 1881). Type: cult. Paris (Anonymus s.n. [P]). – Distr: RSA (Western Cape, Eastern Cape, KwaZulu-Natal).  Bryonia nana Lamarck (1783). Meeuse (1962) commented: “The three varieties as defined here are not sharply separable, but

Kedrostis CUCURBITACEAE

the extremes are so different in habit that I hesitate to unite them all”, and this approach is here followed. K. nana var. nana – Distr: RSA (Eastern Cape, Western Cape, KwaZulu-Natal); climbing on shrubs in sandy soil, 5–245 m. Incl. Sicyos angulatus P. J. Bergius (1767) (nom. illeg., ICN Art. 53.1); incl. Bryonia africana Thunberg (1794) (nom. illeg., ICN Art. 53.1); incl. Bryonia triloba Thunberg (1794) (nom. illeg., ICN Art. 53.1)  Cyrtonema trilobum (Thunberg) Schrader (1834); incl. Cyrtonema molle G. Kunze (1847)  Coniandra molle (G. Kunze) Sonder (1862)  Kedrostis mollis (G. Kunze) Cogniaux (1881); incl. Coniandra thunbergii Sonder (1862); incl. Zehneria hederacea Sonder (1862)  Melothria hederacea (Sonder) Cogniaux (1881); incl. Kedrostis velutina Cogniaux (1908); incl. Kedrostis angulata Fourcade (1932). Perennial monoecious climbing herbs with thick tuberous R; stems slender, climbing or rarely prostrate, branching, furrowed, sparsely hirsute becoming glabrous; tendrils filiform, short; L petiole 1–4 cm, slender, with short hairs, lamina fleshy, cordate-reniform or suborbicular-cordate, 2–7  2–7 cm, simple or with 3–5 short lobes, pale to deep green, glabrous or with sparse short hairs, margins entire or slightly undulating, apex sometimes mucronate; male Fl to 20 in racemes on 2–10 cm long peduncles; Ped 0.3–1.6 cm; hypanthium campanulate to subcylindrical, 2–4 mm, usually  hairy; Sep narrowly triangular, 1–2 mm; Pet ovate-oblong or ovate-triangular, 4–9 mm, light yellow, densely papillose outside; female Fl solitary on 0.8 cm long peduncle; Pet greenish-yellow; Ov oblongfusiform,  beaked; Fr ovoid, conical-acute to rostrate, 1.5–2  0.7–0.8 cm, orange to red, smooth and glabrous, few-seeded; Se 6–7  3–4 mm, smooth. K. nana var. schlechteri (Cogniaux) A. Meeuse (Bothalia 8(1): 31, 1962). Type: RSA, Eastern Cape (Schlechter 6053 [Z, BOL, GRA]). – Distr: RSA (Eastern Cape, Western Cape); sandy loam in scrub, on slopes, 60–750 m.

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 Kedrostis schlechteri Cogniaux (1906); incl. Bryonia digitata Thunberg (1794)  Coniandra digitata (Thunberg) Sonder (1862)  Kedrostis digitata (Thunberg) Cogniaux (1881). Differs from the other two varieties: L lamina usually deeply dissected with narrow (tongueshaped) lobes or ultimate lobes, upper or both faces with distinct white pustules, rarely without. K. nana var. zeyheri (Schrader) A. Meeuse (Bothalia 8(1): 31, 1962). Type: RSA, Eastern Cape (Ecklon & Zeyher 1775 [L, NBG]). – Distr: RSA (Eastern Cape, Western Cape, KwaZulu-Natal); stony soils in scrub and forest, 15–915 m.  Coniandra zeyheri Schrader (1834)  Kedrostis zeyheri (Schrader) Cogniaux (1881); incl. Cyrtonema sphenolobum Schrader (1836); incl. Coniandra zeyheri var. angustiloba Sonder (1862)  Kedrostis zeyheri var. angustiloba (Sonder) Cogniaux (1881); incl. Kedrostis digitata var. major Cogniaux (1881). Differs from var. nana: L lamina dissected beyond the middle with rhomboid to oblongcuneiform, often 3-lobed, acute or obtuse lobes, usually glabrous. K. psammophila Bruyns (Bothalia 23(2): 233–235, ills., 1993). Type: RSA, Northern Cape (Bruyns 4569 [BOL, K, M, PRE]). – Distr: RSA (Northern Cape, Western Cape); fine reddish sands, 80 m. Tuberous, monoecious; R tuber to 20  10 cm; stems creeping horizontally below the soil surface to 1 m, much branched, aerial shoots to 15 cm, spreading, grey-green, hairy; tendrils absent; L petiole 0.8–1.5 cm, lamina simple, 0.6–2  1–2.5 cm, almost entire to palmately 3- to 5-lobed, grey-green, hairy; Inf arising from underground stems; male Fl  25 on 8 cm long peduncles; Ped 1.2–2.2 cm; hypanthium 4 mm long; Pet 5–7 mm, green, with darker longitudinal stripes on the inner side, hairy; St 3; female Fl solitary on 0–2 cm long peduncles; hypanthium cylindrical, 35–70 mm long, mostly underground; Pet 10–12 mm, pale greenish outside, bright yellow inside; staminodes 5; Fr  globose, 2.2 cm ∅, partly underground, green above ground,

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white below, finely pubescent; Se 8, white, flattened-ovoid, 10  7 mm, not further described. This species is unique in the genus, with stems that are partly below ground, and fruits also partly underground.

References Eggli, U. (2008) Sukkulenten. Ed. 2. Stuttgart (DE): Eugen Ulmer. England, A. (1990) Some do and some don’t. Anacampseros 6(3): 43–46. Ettelt, J. (2018) Empirische Erfahrungen mit der Freilandhaltung und Überwinterung sukkulenter Pflanzen im Keller. Avonia 36(4): 250–263, ills. Jeffrey, C. (1967) Cucurbitaceae. In: Milne-Redhead, E. & Polhill, R. M. (eds.): Flora of Tropical East Africa. London (GB): Crown Agents for Oversea Governments and Administrations. Lenain, H. (2012) Plantes à caudex de A à Z. Paris (FR): Ulmer.

L. E. Newton and G. N. Njoroge Meeuse, A. D. J. (1962) The Cucurbitaceae of southern Africa. Bothalia 8(1): 1–111, keys. https://doi.org/10. 4102/abc.v8i1.1611. Rowley, G. D. (1987) Caudiciform and pachycaul succulents. Pachycauls, bottle-, barrel- and elephant-trees and their kin: A collector’s miscellany. Mill Valley (US): Strawberry Press. Schaefer, H. & Renner, S. S. (2011a) Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60(1): 122–138, ills.; (3): 924 [erratum]. https://doi. org/10.1002/tax.601011. Schaefer, H. & Renner, S. S. (2011b) Cucurbitaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants, volume X; pp. 112–174, ills., key. Heidelberg (DE) etc.: Springer. https://doi.org/10.1007/978-3-64214397-7_10. Unuofin, J. O. [& al. 2018], Otunola, G. A. & Afolayan, A. J. (2018) Acute and subacute toxicity of aqueous extract of the tuber of Kedrostis africana (L.) Cogn in Wistar rats. J. Complem. Integrat. Med. 15(4): 11 pp. https://doi.org/10.1515/jcim-2017-0139. Wilkins, M. H. (1994) Good gourds and succulent squash: A cucurbit review. Cact. Succ. J. (US) 66(2): 65–71, ills.

Linnaeosicyos CUCURBITACEAE L. E. Newton

Linnaeosicyos H. Schaefer & Kocyan (Syst. Bot. 33(2): 350, 2008). Type: Trichosanthes amara Linné. – Sicyoeae – Distr: Hispaniola: Dominican Republic. Etym: For Carl Linné (Carolus Linnaeus; 1707–1778), famous Swedish botanist and originator of the binomial system of naming plants; and Gr. ‘sicyos’, cucumber; for the placement in this family. Perennial dioecious climbers or trailers with long fleshy Rstock; stems to 6 ( 10) m, to 8 ( 30) mm ∅, ribbed, scabrid to glabrescent; tendrils simple, to 12 cm; L petiole 2–3.2 cm, lamina reniform to suborbicular, entire to deeply 3-lobed, 1.2–4.5  1.6–4.2 cm, upper face pustulate with numerous trichomes on multicellular discoidal cystolith-bearing bases; Fl solitary, unisexual, white with green veins, nocturnal; Ped 3–5 cm; male Fl hypanthium broadly campanulate, hypanthium tube 29 mm, 5 mm ∅; Sep 5, narrowly triangular, entire,  10 mm; Pet 5, ovate, 30  12 mm, fimbriate with fringes to 30 mm; St 3; Fil free,  1–3 mm; Anth 3, connate into a head, 2 2-thecous, 1 1-thecous, thecae triplicate; female Fl similar to male flowers, with minute staminodes; Ov ellipsoid,  25 mm, placentae 3; Sti 3-lobed, lobes capitate; Fr turbinate to cylindrical-ellipsoid, green, pendent, 8–12 

3–4 cm, indehiscent, with soft whitish pulp; Se very many, 4–5  1.5–2 mm, linear-oblong, compressed, yellowish-brown, margin distinct. The single species of this genus was found to be misplaced in Trichosanthes Linné ( 100 species in Asia and Australia, no succulents), where it was traditionally placed on account of its deeply fimbriate petals. In the molecular analysis presented in the protologue, Linnaeosicyos is the basal sister to the New World genera of tribe Sicyoeae (Schaefer & al. 2008). Mitchell & al. (2015) studied the pollination biology of this strictly nocturnal taxon, and also illustrate the striking flowers. Several hawkmoths of different sizes but esp. Eumorpha labruscae were observed as visitors, and a small number of carpenter bee visits were also recorded. The flowers are unscented to the human nose but emit compounds known to be associated with hawkmoth pollination. L. amarus (Linné) H. Schaefer & Kocyan (Syst. Bot. 33(2): 350, ills. (pp. 353–354), 2008). Type: [lecto – icono]: Plumier, Descr. Pl. Amér., 86, t. 100, 1693. – Lit: Mitchell & al. (2015: with ills.). Distr: Hispaniola: Dominican Republic; thickets on limestone hills and along roadsides in dry forest, 0–400 m.  Trichosanthes amara Linné (1753). Description as for the genus.

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected] © Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_47

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References Mitchell, T. C. [& al. 2015], Dötterl, S. & Schaefer, H. (2015) Hawk-moth pollination and elaborate petals in Cucurbitaceae: The case of the Caribbean endemic Linnaeosicyos amara [sic!]. Flora (Jena) 216: 50–56, ills. https://doi.org/10.1016/j.flora.2015.08.004.

L. E. Newton Schaefer, H. [& al. 2008], Kocyan, A. & Renner, S. S. (2008) Linnaeosicyos (Cucurbitaceae): A new genus for Trichosanthes amara, the Caribbean sister species of all Sicyeae. Syst. Bot. 33(2): 349–355, ills. https:// doi.org/10.1600/036364408784571707.

Marah CUCURBITACEAE L. E. Newton

Marah Kellogg (Proc. Calif. Acad. Nat. Sci. 1: 38–39, 1854). Type: Marah muricata Kellogg.
Sicyoeae
Lit: Dunn (1913: outdated monograph); Stocking (1955: taxonomy & ecology); Martin (2009: ethnobotany); Nesom (2015: 21–25, Flora North America). Distr: W USA, NW Mexico. Etym: From Hebrew ‘marah’, bitter; for the bitter taste of the roots. Incl. Megarrhiza Torrey & Gray (1860).Type: Megarrhiza oregona Torrey & Gray. Monoecious or sometimes temporarily dioecious climbers or trailers with (sometimes very) large sometimes partly exposed hypocotylderived tubers; stems coarse; Br slender; tendrils simple or bifid; L with 3–9 lobes; Fl rotate to  cup-shaped, 3–15 mm ∅; Sep 5 and filiform, or absent; male Fl in simple or compound racemes or rarely solitary; hypanthium campanulate; Pet 5, triangular to ovate, fused for ½, white, whitish or cream to yellowish-green; St 3, rarely 4; Fil united; Anth twisted together and forming a head; female Fl solitary, usually larger than male flowers; staminodes 0–3; Fr ovoid to globose or oblong, large, 3–20  3–6 cm, fleshy becoming dry, irregularly dehiscent, covered with stout

L. E. Newton (*) Herbarium, Royal Botanic Gardens, Kew, UK e-mail: [email protected]

smooth spines; Se few to many, large, >1 cm, globose to ovoid, turgid and not or moderately compressed, smooth, margin indistinct.
Cytology: 2n ¼ 15 (Parfitt & al. 1990). A small genus of 7 species. All occur in semiarid climates and have fleshy tuberous and sometimes “gigantic” roots, and are covered below. The genus was suggested to be closely related to Echinocystis and Echinopepon by Stocking (1955: 120), and this was corroborated by the molecular phylogeny of Schaefer & Renner (2011: fig. S5). Tuber morphology is described in some detail by Stocking (1955) for several species: Tubers develop from the hypocotyl and are first fusiform for several years and only with time assume a globose shape. The heaviest tuber recorded was for M. fabacea, and weighted 58 kg. The interior tissue is a “starchy parenchyma”. Schlising (1969) studied the germination and early seedling growth of M. oregana: Seeds are buried by nocturnal rodents and germinate underground. First, the bases of the cotyledons elongate downwards and form a hollow tube, pushing the seedling proper deeper into the soil for 5–25 cm. Only now the radicle starts growing downwards, and the epicotyl grows upwards through the hollow tube. Concurrently, the hypocotyl starts to enlarge and forms a tuber that persists through the ensuing dry season. The tuber of this and other species of Marah are thus hypocotyl tubers and help with seedling establishment. All Californian species of Marah show the same mode of development (Schlising 1969: 557).

© Springer Nature Switzerland AG 2023 U. Eggli, R. Nyffeler (eds.), Dicotyledons: Rosids, Illustrated Handbook of Succulent Plants, https://doi.org/10.1007/978-3-030-93492-7_48

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All species appear to have been been used in folk medicine as emetics, purgatives, vermifuges and against malaria, and for ceremonial purposes. The seeds are rich in oil, and seed remains esp. of M. macrocarpa are present at many archaeological sites (Martin 2009). The grammatical gender of the name Marah was treated as masculine in the protologue but changed to feminine by the publishing author himself in a later publication; today, both versions are used. Here, we follow the usage of IPNI (accessed Jan. 2021) and treat it as feminine. M. fabacea (Naudin) Greene (Leafl. Bot. Observ. 2: 36, 1910). Type: USA, California (Anonymous s.n. [P †; [lecto
icono:] Ann. Sci. Nat., Bot., sér. 4, 12: t. 9, 1859]).
Distr: SW USA (California, W Nevada); coastal strand, streamsides and washes, rock outcrops and grassland or wooland, 20–1400 m.  Echinocystis fabacea Naudin (1859)  Micrampelis fabacea (Naudin) Greene (1890); incl. Micrampelis fabacea var. agrestis Greene (1891)  Marah fabacea var. agrestis (Greene) Stocking (1955); incl. Echinocystis inermis Congdon (1901)  Marah inermis (Congdon) Dunn (1913)  Echinocystis fabacea var. inermis (Congdon) Jepson (1936); incl. Echinocystis scabrida Eastwood (1903). Stems to 7 m, to 3 mm ∅, slightly pubescent to subglabrous, Int 7–10 cm; tendrils usually bifid; L petiole 3–6 cm, lamina suborbicular, 5–10  5–10 cm, glabrous to somewhat scabrous,  deeply 5- to 7-lobed, lobes