Monocotyledons (Illustrated Handbook of Succulent Plants) 366256484X, 9783662564844

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Illustrated Handbook of Succulent Plants

Urs Eggli · Reto Nyffeler Editors

Monocotyledons 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, 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 colour photographs complete this inventory of succulent plants. More information about this series at http://www.springer.com/series/4452

Urs Eggli • Reto Nyffeler Editors

Monocotyledons Second Edition

With 339 Figures and 12 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

ISBN 978-3-662-56484-4 ISBN 978-3-662-56486-8 (eBook) ISBN 978-3-662-56485-1 (print and electronic bundle) https://doi.org/10.1007/978-3-662-56486-8 1st edition: © Springer-Verlag Berlin Heidelberg 2001 2nd edition: © Springer-Verlag GmbH Germany, part of Springer Nature 2020 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, 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-Verlag GmbH, DE, part of Springer Nature. The registered company address is: Heidelberger Platz 3, 14197 Berlin, Germany

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 Handbook 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 Bromelieaceae and Cactaceae were not included. The past more than 15 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. Fifteen years is a long stretch of time for any lexicon, and much new data have been generated by recent research in all fields of botany. During this time, especially molecular studies have led to an improved understanding of the living diversity of plants, and a firm phylogenetic backbone is now in place for most 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 in the future. 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 followed, with the necessary cautious comments (see, e.g., the genus Aloe).

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For this second edition, 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, and we are delighted about the willingness of the respective authors to accept this task. In contrast to the first edition of this volume, the succulent taxa of the pineapple family (Bromeliaceae) are now also included, thanks to the energetic help of a group of specialists of that family. With this addition, this volume now covers the monocotyledonous succulents in full with the notable exception of the orchids – the sheer number of possible succulent species makes their inclusion unfeasible. 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. Sukkulenten-Sammlung Zürich, Grün Stadt Zürich Zürich, Switzerland Institut für Systematische und Evolutionäre Botanik (ISEB), Universität Zürich Zürich, Switzerland December 2019

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 the first handbooks devoted to the family Cactaceae were already published in the nineteenth 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. For various reasons, the Lexicon project took much longer to be completed than anticipated. The Cactaceae part had not become available in time for vii

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From the Preface to the First Edition

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 (= 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 III 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 and 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 and 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 semiarid 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, for example, the Commelinaceae and Gesneriaceae (e.g., Columnea), and several other families. 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 semiarid conditions (see Ogburn and 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 and Eggli 2010a) showed some 2200 succulent species for this family, and a more detailed census (see the family treatment in this volume) arrives at a minimum count of 4500 succulent species – this

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General Introduction to the Series

makes the Orchidaceae the most important family containing succulents in term of species number. Fortunately, vast specialist literature covering orchids exists, numerous specialist societies are in existence, and the available lexicographic efforts (e.g., Pridgeon & al. 1999, 2001, 2003, 2005, 2009, 2014) do not need to be duplicated here. Also omitted from the series are the pteridophytes, as insufficient knowledge on the – possibly few – marginally succulent taxa is available.

Succulence Among 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 time 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 and 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 breakdown into volumes is as follows (see also Fig. 1): • Monocots: Eggli, U. & Nyffeler, R. (eds.): Illustrated Handbook of Succulent Plants. Monocotyledons. This volume. • Dicots A: Eggli, U. (ed.): Illustrated Handbook of Succulent Plants. Magnoliids and Dicotyledons (excluding Rosids). • Dicots B: Eggli, U. (ed.): Illustrated Handbook of Succulent Plants: Dicotyledons: Rosids. • 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. (2010), and Soltis & al. (2011). The estimated number of succulent species (modified from Nyffeler and Eggli 2010) is indicated to the right of the order names. Supraordinal names follow Soltis & al. (2011) and Chase and 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 60 Zygophyllales 40 Celastrales Malpighiales 1030 Oxalidales 60 Fabales 25 Rosales 30 Cucurbitales 100 Fagales Geraniales 155 Myrtales 3 Crossosomatales Picramnales Sapindales 37 Huerteales Malvales 30 Brassicales 70 Dilleniales Berberopsidales Santalales 3 Caryophyllales 3991 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. (Diagram by Urs Eggli) rats

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General Introduction to the Series

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 Caryophyllaceae Clusiaceae Commelinaceae Convolvulaceae Crassulaceae Cucurbitaceae Cyperaceae Didiereaceae Dioscoreaceae Doryanthaceae Ericaceae Eriospermaceae Euphorbiaceae Fabaceae Fouquieriaceae Francoaceae Geraniaceae Gesneriaceae Halophytaceae Hyacinthaceae Icacinaceae

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 Caryophyllales Malpighiales Commelinales Solanales Saxifragales Cucurbitales Poales Caryophyllales Dioscoreales Asparagales Ericales Asparagales Malpighiales Fabales Ericales Geraniales Geraniales Lamiales Caryophyllales Asparagales Icacinales

Monocot (this volume) 

Dicot A

Dicot B (Rosids)

Separate volume 

                                            (continued)

General Introduction to the Series

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Table 1 (continued) Family Lamiaceae 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 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 (this volume)

Dicot A     

Dicot B (Rosids)

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: 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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Chase, M. W. & Reveal, J. L. (2009) A phylogenetic classification of the land plants to accompany APG III. Bot. J. Linn. Soc. 161(2): 122–127. https:// doi.org/10.1111/j.1095-8339.2009.01002.x. Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions – succulence as an adaptive strategy. Bradleya 27: 13–36. https://doi.org/ 10.5167/uzh-28825. Moore, M. J. [& al. 2010], Soltis, P. S., Bell, C. D., Burleigh, J. G. & Soltis, D. E. (2010) Phylogenetic analysis of 83 plastid genes further resolves the early diversification of eudicots. Proc. Nation. Acad. Sci. USA 107(10): 4623–4628. https://doi.org/10.1073/pnas.0907801107. Nyffeler, R. & Eggli, U. (2010a) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/ 10.5167/uzh-35243. Nyffeler, R. & Eggli, U. (2010b) Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 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. Pridgeon, A. M. [& al. 1999], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (1999) Genera Orchidacearum. Volume 1. General introduction, Apostasioideae, Cypripedioideae. Oxford (GB): Oxford University Press. Pridgeon, A. M. [& al. 2001], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (2001) Genera Orchidacearum. Volume 2. Orchidoideae (part one). Oxford (GB): Oxford University Press. Pridgeon, A. M. [& al. 2003], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (2003) Genera Orchidacearum. Volume 3. Orchidoideae (part two), Vanilloideae. Oxford (GB): Oxford University Press. Pridgeon, A. M. [& al. 2005], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (2005) Genera Orchidacearum. Volume 4. Epidendroideae (part one). Oxford (GB): Oxford University Press. Pridgeon, A. M. [& al. 2009], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (2009) Genera Orchidacearum. Volume 5. Epidendroideae (part two). Oxford (GB): Oxford University Press. Pridgeon, A. M. [& al. 2014], Cribb, P. J., Chase, M.W.&Rasmussen, F. N. (eds.) (2014) Genera Orchidacearum. Volume 6. Epidendroideae (part three). Oxford (GB): Oxford University Press. Soltis, D. E. [& al. 2011], Smith, S. A., Cellinese, N., Wurdack, K. J., Tank, D. C., Brockington, S. F., Refulio-Rodriguez, N. F., Walker, J. B., Moore, M. J., Carlsward, B. S., Bell, C. D., Latvis, M., Crawley, S., Black, C., Diouf, D., Xi, Z., Rushworth, C. A., Gitzendanner, M. A., Sytsma, K. J., Qiu, Y.-L., Hilu, K. W., Davis, C. C., Sanderson, M. J., Beaman, R. S., Olmstead, R. G., Judd, W. S., Donoghue, M. J. & Soltis, P. S. (2011) Angiosperm phylogeny: 17 genes, 640 taxa. Amer. J. Bot. 98(4): 704–730. https://doi.org/10.3732/ajb.1000404.

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 cross-reference 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 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, the reader is referred to general botanical books that include keys to plant families. Rowley (1980) and Eggli (2008) provided keys for flowering and nonflowering 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 Compositae for Asteraceae) are not used. Important synonyms at family rank are indicated, especially when the synonymous names include succulents. The order to which the family belongs is also given. Within each family, the genera are treated in alphabetical sequence and the same applies to the sequence of species within genera. 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 among its succulent representatives. This is supplemented by notes on the distribution, phylogeny, classification, and economic importance of the family and the occurrence of xv

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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. The family concept adopted more or less follows APG III and APG IV, but with the modifications for Asparagales proposed by Nyffeler and Eggli (2010a).

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 and 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 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 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, for example, on population biology, pollination, and seed dispersal or ethnobotany, are 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 are enumerated immediately following the entry for the genus. Cultivars

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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 ICBN/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, that is, 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 numerous botanical glossaries, of which Stearn (1992) is cited by way of a most important and useful example.

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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 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 and 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.

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 ICBN or ICN articles violated. In many of these cases, the numbering still follows the “Vienna Code” (McNeill & al. 2006), but for more recent names, the numbering in the “Melbourne Code” (McNeill & al. 2012) is used; this is indicated by preceding the violated articles by “ICN.” Spelling variants are considered as invalidly published according to ICBN/ICN Art. 61. While invalid names are, strictly speaking, not considered as names in ICBN/ICN (see Art. 6.3), they are nonetheless listed (and flagged accordingly with the violated articles of ICBN/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(s) 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 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 (= heterotypic synonyms). Again, groups of combinations based on the same basionym are united with -signs. Basionyms are given first and in chronological order.

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

How to Use This Handbook

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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 1990s, especially affecting the former Cape Province, which has been split up into four units (North-West Province, Northern Cape, Western Cape, Eastern Cape). We 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, 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 and Cowan (1976–1988) and supplements (Stafleu and Mennega 1992–2000, Dorr and 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.

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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. Eggli, U. (1985) A bibliography of succulent plant periodicals. Bradleya 3: 103–119. https://doi.org/10.25223/brad.n3.1985.a6. 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. Eggli, U. & Nyffeler, R. (2009) Living under temporarily arid conditions – succulence as an adaptive strategy. Bradleya 27: 13–36. https://doi.org/ 10.5167/uzh-28825. 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. 2006], Barrie, F. R., Burdet, H. M., Demoulin, V., Hawksworth, D. L., Marhold, K., Nicolson, D. J., Prado, J., Silva, P. C., Skog, J. E., Wiersema, J. H. & Turland, N. J. (eds.) (2006) International Code of Botanical Nomenclature (Vienna Code). Ruggell (FL): A. R. G. Gantner Verlag KG. 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. (2010a) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/ 10.5167/uzh-35243. Nyffeler, R. & Eggli, U. (2010b) Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. https://doi.org/ 10.1002/tax.591021.

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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. 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. Stafleu, F. A. & Mennega, E. A. (1992–2000) Taxonomic literature. Königstein (DE): Koeltz Scientific Books. Supplements I–VI to Ed. 2; 6 vols. Stearn, W. T. (1992) Botanical Latin. Newton Abbot (GB): David & Charles Publishers. Ed. 4; 560 pp.

Contents

Volume 1 Introduction to the Classification of Monocotyledons . . . . . . . . . . . R. Nyffeler and U. Eggli

1

Part I

The Family Agavaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

7

Agavaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. Thiede and U. Eggli

9

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

21

Agave AGAVACEAE J. Thiede

Beschorneria AGAVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313 J. Thiede Furcraea AGAVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 J. Thiede Hesperaloe AGAVACEAE J. Thiede

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349

Hesperoyucca AGAVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357 J. Thiede Yucca AGAVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 363 J. Thiede Part II

The Family Amaryllidaceae . . . . . . . . . . . . . . . . . . . . . . . . . .

423

Amaryllidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 425 U. Eggli, E. Van Jaarsveld, and S. Arroyo-Leuenberger Boophone AMARYLLIDACEAE E. Van Jaarsveld

. . . . . . . . . . . . . . . . . . . . . . . . . . 429

Brunsvigia AMARYLLIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 433 E. Van Jaarsveld

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Contents

Cyrtanthus AMARYLLIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 435 E. Van Jaarsveld Haemanthus AMARYLLIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . 441 E. Van Jaarsveld Rauhia AMARYLLIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445 S. Arroyo-Leuenberger Part III The Family Anthericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . .

447

Anthericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 449 E. Van Jaarsveld and U. Eggli Chlorophytum ANTHERICACEAE . . . . . . . . . . . . . . . . . . . . . . . . 451 E. Van Jaarsveld Part IV

The Family Araceae

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

455

Araceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 457 J. Bogner Zamioculcas ARACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 461 J. Bogner Part V

The Family Asparagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . .

463

Asparagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 465 E. Van Jaarsveld and U. Eggli Asparagus ASPARAGACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 467 E. Van Jaarsveld and U. Eggli Part VI

The Family Asphodelaceae

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

473

Asphodelaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475 G. F. Smith and E. Figueiredo Algastoloba ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 483 U. Eggli Aloe ASPHODELACEAE L. E. Newton

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 485

Aloloba ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697 U. Eggli Alworthia ASPHODELACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . 699

Astroloba ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 701 N. L. Meyer and G. F. Smith

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Astroworthia ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . 709 N. L. Meyer and G. F. Smith Bayerara ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 711 U. Eggli Bulbine ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 713 E. Van Jaarsveld and P. I. Forster Bulbinella ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 741 E. Van Jaarsveld Cummingara ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . 743 U. Eggli Gasteraloe ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 745 L. E. Newton Gasterhaworthia ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . 749 U. Eggli Gasteria ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 751 E. Van Jaarsveld Gastroloba ASPHODELACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . 767

Haworthia ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 769 M. B. Bayer and E. Van Jaarsveld Kniphofia ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 819 G. F. Smith and E. Figueiredo Maysara ASPHODELACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . 823

Poellneria ASPHODELACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . 825

Poellnitzia ASPHODELACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 827 G. F. Smith Trachyandra ASPHODELACEAE E. Van Jaarsveld

. . . . . . . . . . . . . . . . . . . . . . . . . 829

Volume 2 Part VII

The Family Bromeliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . .

833

Bromeliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 835 U. Eggli and E. J. Gouda Acanthostachys BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . 849

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Contents

Aechmea BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 851 U. Eggli Billbergia BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 859

Biltanthus BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . 863

Cryptanthus BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 865 U. Eggli Deinacanthon BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 875 U. Eggli Deuterocairnia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . 877 N. Schütz and F. Krapp Deuterocohnia BROMELIACEAE N. Schütz

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

Dyckcohnia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 891 N. Schütz and F. Krapp Dyckia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 893 F. Krapp and U. Eggli Dycktia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 963 N. Schütz and F. Krapp Encholirium BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 965 F. Krapp Enchotia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 983 U. Eggli Fascicularia BROMELIACEAE U. Eggli Fosterella BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . 985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 989

Hechcohnia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 995 N. Schütz and F. Krapp Hechtia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 997 A. Espejo-Serna, A. R. López-Ferrari, and N. Martínez-Correa Neobergia BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . 1033

Neoglaziovia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1035 U. Eggli

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Neophytum BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 1039 U. Eggli Ochagavia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041 U. Eggli Orthocohnia BROMELIACEAE N. Schütz

. . . . . . . . . . . . . . . . . . . . . . . . . 1045

Orthoglaziovia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . 1047 U. Eggli Orthomea BROMELIACEAE U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . 1049

Orthophytum BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 1051 U. Eggli and E. J. Gouda Orthotanthus BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 1067 U. Eggli Puckia BROMELIACEAE N. Schütz and F. Krapp

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1069

Pucohnia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1071 N. Schütz and F. Krapp Puya BROMELIACEAE J. M. Manzanares

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1073

Sincoraea BROMELIACEAE U. Eggli and E. J. Gouda

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1101

Tillandsia BROMELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1107 E. J. Gouda Part VIII

The Family Commelinaceae . . . . . . . . . . . . . . . . . . . . . . . . 1165

Commelinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1167 D. R. Hunt Aneilema COMMELINACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1171 D. R. Hunt Callisia COMMELINACEAE D. R. Hunt

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

Cyanotis COMMELINACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1179 D. R. Hunt Tradescantia COMMELINACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 1183 D. R. Hunt

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Contents

Tripogandra COMMELINACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 1189 D. R. Hunt Part IX

The Family Cyperaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1193

Cyperaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1195 U. Eggli Part X

The Family Dioscoreaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . 1197

Dioscoreaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1199 U. Eggli and G. D. Rowley Dioscorea DIOSCOREACEAE G. D. Rowley and U. Eggli Part XI

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1201

The Family Doryanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 1209

Doryanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1211 P. I. Forster and U. Eggli Doryanthes DORYANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 1213 P. I. Forster Part XII

The Family Eriospermaceae . . . . . . . . . . . . . . . . . . . . . . . . 1215

Eriospermaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1217 E. Van Jaarsveld and U. Eggli Eriospermum ERIOSPERMACEAE . . . . . . . . . . . . . . . . . . . . . . . . 1219 E. Van Jaarsveld Part XIII

The Family Hyacinthaceae . . . . . . . . . . . . . . . . . . . . . . . . . 1223

Hyacinthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1225 E. Van Jaarsveld and U. Eggli Albuca HYACINTHACEAE E. Van Jaarsveld

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

Bowiea HYACINTHACEAE E. Van Jaarsveld

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1243

Dipcadi HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1247 U. Eggli Drimia HYACINTHACEAE E. Van Jaarsveld and U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1249

Lachenalia HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1265 E. Van Jaarsveld

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Ledebouria HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1269 E. Van Jaarsveld Massonia HYACINTHACEAE E. Van Jaarsveld and U. Eggli

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 1273

Merwilla HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1277 E. Van Jaarsveld Namophila HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1279 U. Eggli Ornithogalum HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 1281 E. Van Jaarsveld Spetaea HYACINTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1289 E. Van Jaarsveld Part XIV

The Family Laxmanniaceae . . . . . . . . . . . . . . . . . . . . . . . . 1291

Laxmanniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1293 C. C. Walker Cordyline LAXMANNIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1295 C. C. Walker Part XV

The Family Orchidaceae

. . . . . . . . . . . . . . . . . . . . . . . . . . . 1297

Orchidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1299 U. Eggli Part XVI The Family Poaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1305 Poaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1307 E. Van Jaarsveld and U. Eggli Dregeochloa POACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1311 E. Van Jaarsveld Part XVII

The Family Ruscaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1313

Ruscaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1315 U. Eggli Beaucarnea RUSCACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1319 C. C. Walker Dasylirion RUSCACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1329 C. C. Walker Dracaena RUSCACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1337 C. C. Walker

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Contents

Nolina RUSCACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1345 C. C. Walker Sansevieria RUSCACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1353 L. E. Newton Part XVIII

The Family Xanthorrhoeaceae . . . . . . . . . . . . . . . . . . . . 1387

Xanthorrhoeaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1389 U. Eggli Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1391

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. BG C clono

cult. cv. Distr Distr. E e.g. epi esp. Etym excl. fa. fig., figs. fl. Gr. Herb. holo Hort., hort.

Article of ICBN/ICN Botanical Garden central clonoytpe, 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 cultivated cultivar distribution District East, eastern for example epitype (nomenclatural type) especially etymology excluding forma (botanical rank) figure, figures “floruit,” i.e., lived Greek herbarium holotype (nomenclatural type) garden, of gardens xxxi

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List of Abbreviations and Symbols

Illustration International Code of Botanical Nomenclature (“Vienna Code,” 2006) ICN International Code of Nomenclature for Algae, Fungi, and Plants (“Melbourne Code,” 2012) ill., ills. illustration, illustrations; illustrated illeg. illegitimate (nomenclatural concept) in sched. “in schedis,” i.e., in the herbarium incl. including ING Index Nominorum Genericorum inval. invalid (nomenclatural concept) KG, KGW Karroo Garden, Worcester, RSA l.c. “loco citato,” i.e., at the place cited lecto lectotype (nomenclatural type) Lit literature ms. manuscript Mt., Mts. mountain, mountains mybp million years before present N North, northern NBG National Botanical Garden, RSA NE North-East, north-eastern neo neotype (nomenclatural type) nom. “nomen,” i.e., botanical name NW North-West, north-western p., pp. page, pages p.a. “per annum,” i.e., per year p.p. “pro parte,” i.e., partly pers. comm. personal communication pl. plate pp. pages Prov. Province publ. published q.v. “quod vide,” i.e., which see RSA Republic of South Africa S South, southern s.a. “sine anno,” i.e., without year s.l. “sensu lato,” i.e., in a wide sense s.n. “sine numero,” i.e., without (collection) number s.s. “sensu stricto,” i.e., in the strict/narrow sense SE South-East, south-eastern Sect. section (botanical rank) SEM Scanning electron microscope Ser. series (botanical rank) spp. several species ssp. subspecies (botanical rank) Subgen. subgenus (botanical rank) Subsect. subsection (botanical rank) I ICBN

List of Abbreviations and Symbols

Subser. SUG SW syn t., tt. unpubl. USA var. W WCSP

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subseries (botanical rank) Stellenbosch University Botanical Garden South-West, south-western syntype (nomenclatural type) “tabula,” i.e., illustration plate/plates unpublished United States 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/

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 Int ITep L Nec NGl NSc OTep Ov Pc Ped

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 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

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Per Pet Phy Poll R Rec Ri Ros Sc Se Sep Sp SpS St Sti Sty Tep Tu

List of Abbreviations and Symbols

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 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 a 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 for the vascular plant collections at the United Herbaria of the University and ETH Zurich, and he is a lecturer for plant systematics at the xxxv

xxxvi

About the Editors

Department of Systematic and Evolutionary Botany, University of Zurich. 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

S. Arroyo-Leuenberger Instituto de Botanica Darwinion, Buenos Aires, Argentina M. B. Bayer Hermanus, South Africa J. Bogner Gersthofen, Germany U. Eggli Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland A. Espejo-Serna Herbario Metropolitano, Depto. Biología, División de Ciencias Biológicas y de la Salud, Universidad Autónoma Metropolitana Iztapalapa, Ciudad de México, Mexico E. Figueiredo Department of Botany, Nelson Mandela University, Port Elizabeth, Eastern Cape, South Africa Centre for Functional Ecology, Departamento de Ciências da Vida, Calçada Martim de Freitas, Universidade de Coimbra, Coimbra, Portugal P. I. Forster Department of Science, Information Technology and Innovation, Brisbane Botanic Gardens, Queensland Herbarium, Toowong, QLD, Australia E. J. Gouda Curator University Botanic Gardens, Utrecht, The Netherlands D. R. Hunt Sherborne, UK F. Krapp Guxhagen, Germany A. R. López-Ferrari Herbario Metropolitano, Depto. Biología, División de Ciencias Biológicas y de la Salud, Universidad Autónoma Metropolitana Iztapalapa, Ciudad de México, Mexico J. M. Manzanares Herbario Nacional (QCNE), Sección Botánica del Museo Ecuatoriano de Ciencias Naturales del Instituto Nacional de Biodiversidad, Quito, Ecuador N. Martínez-Correa Herbario Metropolitano, Depto. Biología, División de Ciencias Biológicas y de la Salud, Universidad Autónoma Metropolitana Iztapalapa, Ciudad de México, Mexico

D. R. Hunt: deceased. xxxvii

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N. L. Meyer South African National Biodiversity Institute, Pretoria, South Africa L. E. Newton Barking, Essex, UK R. Nyffeler Institut für Systematische und Evolutionäre Botanik (ISEB), Universität Zürich, Zürich, Switzerland G. D. Rowley Reading, UK N. Schütz Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany G. F. Smith Department of Botany, Nelson Mandela University, Port Elizabeth, Eastern Cape, South Africa Centre for Functional Ecology, Departamento de Ciências da Vida, Calçada Martim de Freitas, Universidade de Coimbra, Coimbra, Portugal J. Thiede Hamburg, Germany E. Van Jaarsveld Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa C. C. Walker School of Environment, Earth and Ecosystem Sciences, The Open University, Milton Keynes, England

G. D. Rowley: deceased.

Contributors

Introduction to the Classification of Monocotyledons R. Nyffeler and U. Eggli

General Remarks Succulents of the Monocotyledons (“monocots”, subclass Liliidae or superorder Lilianae) contribute in an important manner to the overall plant diversity in many arid and semi-arid regions. Generally, they are easily recognized on the basis of their characteristic parallel-veined leaves often in rosettes or as geophytic bulbs. Prominent and landscape-dominating representatives are shrubby to arborescent monocots such as the “kokerboom” (Aloe dichotoma, Asphodelaceae) in the Namib desert of SW Africa, or tall-growing and branched Yucca species, e.g. the “Joshua tree” (Yucca brevifolia, Agavaceae), which form open forests in parts of the Mojave desert and surroundings in California, Unites States. The Monocotyledons comprise almost 25% of the total existing vascular plant biodiversity, i.e. c. 70,000 species. Overall, about 10%, i.e. 6460 monocot species, are recognized as succulents, present in five of the eleven orders commonly recognized today (APG 2009, 2016, Christenhusz et al. 2016). Molecular phylogenetic studies of the

R. Nyffeler Institut für Systematische und Evolutionäre Botanik (ISEB), Universität Zürich, Zürich, Switzerland e-mail: [email protected] U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

past three decades have provided a well-founded “backbone” of relationships among the distinct clades recognized as orders (e.g., Givnish & al. 2006, Moore & al. 2010, Soltis & al. 2011). Our current knowledge on the phylogenetic relationships among the monocot orders and their share of succulent representatives is illustrated in Fig. 1.

Family Classification in the Order Asparagales There is unanimous agreement about the circumscription of the monophyletic order Asparagales in all recent phylogenetic studies (Chase & al. 2000, Fay & al. 2000). This order, comprising in total some 36,000 species of which some 16% are succulent, includes several species-rich distinct succulent lineages. However, their classification into families is still dealt with controversially, as they include most of the former “Liliaceae” and relatives as recognized by Cronquist (1981) and other authors in the era before molecular phylogenetic studies. Relationships among lineages in the order Asparagales are reconstructed in rather stable form (Bogler & al. 2006, Pires & al. 2006, Seberg & al. 2012, Steele & al. 2012) and consensus is reached (e.g., Judd & al. 2016) concerning relative relationships among the major clades, similar to the relationships illustrated in Fig. 2. The order includes a grade of families referred to as “lower Asparagales”, including Orchidaceae and

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_113

1

2

R. Nyffeler and U. Eggli

Acorales (3 / −) (4560 / 1)

Alismatales

Petrosaviales (4 / −) Dioscoreales (1037 /

7)

Pandanales (1345 / −) Liliales (1558 / −)

Asparagales

Poales

(36205 / 5973)

(18875 / 500)

Commelinids

Arecales (2585 / −) Commelinales (812 / 25) Zingiberales (2185 / −)

Fig. 1 Summary phylogeny of the monocot orders following APG (2009, 2016). The numbers to the right of the ordinal names are the total species number and the number of succulent species. The widths of the triangles are proportional to the number of species per order. The proportion of succulents is illustrated by the black part of each triangle.

Except for the order Asparagales, all other orders only comprise a few dozen succulent species, with the only exception of the Poales where the bromeliads (family Bromeliaceae) contribute almost 500 species of succulents. (Copyright: U. Eggli)

Iridaceae, and a well-supported clade of “higher Asparagales” consisting of the broadly circumscribed families Asphodelaceae s.l. (changed for nomenclatural reasons from the previously used

Xanthorrhoaeaceae s.l.), Amaryllidaceae s.l. and Asparagaceae s.l. However, no final consensus has been reached for the classification into families, largely due to different concepts of

Introduction to the Classification of Monocotyledons

3

Orchidaceae

(26400 / 4500)

Boryaceae (12/ −) Blandfordiaceae (4/ −) Lanariaceae (1/ −) Asteliaceae (31/ −) Hypoxidaceae (120/ −) Ixioliriaceae (3/ −) Tecophilaeaceae (25/ −) Doryanthaceae (2 / 2) Iridaceae (2120/ −)

Asphodelaceae s.s. (1050 / 826) Agapanthaceae (9/ −) Alliaceae (795/ −) Amaryllidaceae s.s.

(870 / 21)

Asph. s.l. Amar. s.l.

Xeronemataceae (2/ −) Hemerocallidaceae (89/ −) Xanthorrhoeaceae s.s. (30 −) /

Aphyllanthaceae (1/ −) Themidaceae (62/ −) (920 / 86)

Anemarrhenaceae (1/ −) Agavaceae (445 / 392) Anthericaceae (285 / 4) Laxmanniaceae (180 / 1) Asparagaceae s.s. (250 / 10) Ruscaceae (375 / 126) Eriospermaceae (102 / 5)

Fig. 2 Summary phylogeny of the Asparagales (data from Judd & al. 2007, Mabberley 2008, Nyffeler & Eggli 2010, and Stevens (2001 onwards, accessed Dec. 2017). Family

Asparagaceae s.l.

Hyacinthaceae

concepts of APG (2016) are indicated to the right of the shaded boxes. The shading of the boxes relates to the proportion of succulent species in the families. (Copyright: U. Eggli)

4

R. Nyffeler and U. Eggli

Table 1 Comparison of family names used in this series (based on Nyffeler & Eggli 2010) with the family names proposed by APG (2009, 2016), and the subfamily names proposed by Chase & al. (2009) Nyffeler & Eggli 2010, used in this Handbook Asphodelaceae s.s. (1) Amaryllidaceae s.s. Agavaceae Anthericaceae Asparagaceae s.s. Eriospermaceae Hyacinthaceae Ruscaceae (2)

AGP IV (2016), Christenhusz & al. (2016) Asphodelaceae s.l. (3) Amaryllidaceae s.l. Asparagaceae s.l. Asparagaceae s.l. Asparagaceae s.l. Asparagaceae s.l. Asparagaceae s.l. Asparagaceae s.l.

Subfamilies following Chase & al. (2009) Asphodeloideae Amaryllidoideae Agavoideae Agavoideae Asparagoideae Nolinoideae Scilloideae Nolinoideae

Notes: (1) incl. Aloaceae; (2) incl. Dracaenaceae, Nolinaceae; (3) previously, e.g. APG (2009) Xanthorrhoaeaceae s.l.

inclusiveness at the rank of subfamilies versus families in this order. Hence, we identify the different concepts of inclusiveness relative to each other as sensu lato (s.l.) versus sensu stricto (s.s.). Nyffeler & Eggli (2010) have discussed the different options at length, and the situation has not changed during the past half decade (see Nordal & Sletten Bjorå 2016). The main point for discussion remains the number of families to accept, and their diagnostic circumscriptions. Traditional concepts such as the vast family Liliaceae s.l., or the utilization of superior versus inferior ovaries as basis for classificatory decisions, have been found to be not congruent with inferred phylogenetic relationships and these characters are no longer considered as providing valid concepts. While APG (1998) recognized almost 30 families for the Asparagales as a whole, subsequent discussions about the practicability of having so many families for the “higher Asparagales” led to a “lumping” proposal to recognize only the three widely circumscribed families Asphodelaceae (previously called Xanthorrhoaeaceae), Amaryllidaceae and Asparagaceae (APG 2003, 2009, repeated by APG 2016; see Christenhusz & al. 2015 for a history of the APG process and the rationale for the preferred lumping). Hence, and applying these circumscriptions, APG recognizes today 14 families for the order Asparagales. Adherents of this reduction in the number of families and the concomitant wide circumscriptions claim that this results in a gain in practicability, esp. for teaching purposes. On the other side, the proposed radical changes in family circumscriptions are disruptive

in comparison with the traditional “splitting” concepts. It should be noted that the discussion does not concern neither the topology of the phylogenetic backbone, nor accepting nonmonophyletic taxa above the rank of species, but merely affects at what level family names are attached to clades. Since the “lumping” approach of APG (2009, 2016) does not result in a gain of information, and because the alleged gains in practicability remain to be clearly identified, we here follow the traditional (splitting) concept, in line with the considerations of Schmid (2013), and in parallel with the recently updated edition of “Plant systematics: a phylogenetic approach” by Judd & al. (2016; Eriospermaceae not covered). Additional support for the recognition of at least some of the traditionally accepted families is discussed by Seberg & al. (2012) and Nordal & Sletten Bjorå (2016). Our decision keeps all links to previous literature and concepts intact, without concessions to phylogenetic relationships. For clarity, Table 1 lists the two classificatory approaches side-by-side, and also provides the subfamily names suggested by Chase & al. (2009).

References APG [Angiosperm Phylogeny Group] (1998) An ordinal classification for the families of Flowering Plants. Ann. Missouri Bot. Gard. 85: 531–553. https://doi.org/ 10.2307/2992015 APG [Angiosperm Phylogeny Group] (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot.

Introduction to the Classification of Monocotyledons J. Linn. Soc. 141: 399–436. https://doi.org/10.1046/ j.1095-8339.2003.t01-1-00158.x. 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): 105–121. 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 Bogler, D. J. [& al. 2006], Pires, C. & Francisco-Ortega, J. (2006) Phylogeny of Agavaceae based on ndhF, rbcL, and its sequences: Implications of molecular data for classification. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 313–328. https://doi.org/10.5642/ aliso.20062201.26. Chase, M. W. [& al. 2000], Bruijn, A. Y. de, Cox, A. V., Reeves, G., Rudall, P. J., Johnson, M. A. T. & Eguiarte, L. E. (2000) Phylogenetics of Asphodelaceae (Asparagales): An analysis of plastid rbcL and trnL-F DNA sequences. Ann. Bot. (Oxford), n.s. 86: 935–951. https://doi.org/10.1006/anbo.2000.1262. Chase, M. W. [& al. 2009], Reveal, J. L. & Fay, M. F. (2009) A subfamilial classification for the expanded Asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Bot. J. Linn. Soc. 161(2): 132–136. https://doi.org/10.1111/j.1095-8339.2009.00 999.x. Christenhusz, M. J. M. [& al. 2015], Vorontsova, M. S., Fay, M. F. & Chase, M. W. (2015) Results from an online survey of family delimitation in angiosperms and ferns: Recommendations to the Angiosperm Phylogeny Group for thorny problems in plant classification. Bot. J. Linn. Soc. 178(4): 501–528. https://doi. org/10.1111/boj.12285. Christenhusz, M.J.M. [& al. 2016], Fay, M. F. & Chase, M. W. (2016). Plants of the World. An Illustrated Encyclopedia of Vascular Plants. Chicago (US): University of Chicago Press. Cronquist, A. (1981). An integrated system of classification of flowering plants. New York (US): Columbia University Press. Fay, M. F. [& al. 2000], Rudall, P. J., Sullivan, S., Stobart, K. L., Bruijn, A. Y. de, Reeves, G., Qamaruz-Zaman, F., Hong, W.-P., Joseph, J., Hahn, W., Conran, J. V. & Chase, M. W. (2000) Phylogenetic studies of Asparagales based on four plastid DNA regions. In: Wilson, K. L. & Morrison, D. A. (eds.): Monocots: Systematics and evolution. Melbourne (AU): CSIRO. Givnish, T. J. [& al. 2006], Pires, J. C., Graham, S. W., McPherson, M. A., Prince, L. M., Patterson, T. B., Rai, H. S., Roalson, E. H., Evans, T. M., Hahn, W. J., Millam, K. C., Meerow, A. W., Molvray, M., Kores, P. J., O’Brien, H. E., Hall, J. C., Kress, W. J. & Sytsma, K. J. (2006) Phylogenetic relationship of Monocots based on the highly informative plastid gene ndhF: Evidence for widespread concerted convergence. In:

5 Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 28–51. https://doi.org/10.5642/aliso.20062201.04. Givnish, T. J. [& al. 2018], Zuluaga, A., Spalink, D., Soto Gomez, M., Lam, V. K. Y., Saarela, J. M., Sass, C., Iles, W. J. D., Lima de Sousa, D. J., Leebens-Mack, J., Pires, J. C., Zomlefer, W. B., Gandolfo, M. A., Davis, J. I., Stevenson, D. W., dePamphilis, C., Specht, C. D., Graham, S. W., Barrett, C. F. & Ané, C. (2018) Monocot plastid phylogenomics, timeline, net rates of species diversification, the power of multi-gene analyses, and a functional model for the origin of monocots. Amer. J. Bot. 105 (11): 1–23. https://doi.org/10.1002/ajb2.1178. Judd, W. S. [& al. 2007], Campbell, C. S., Kellogg, E. A., Stevens, P. F. & Donoghue, M. J. (2007) Plant systematics: A phylogenetic approach. Ed. 3. Sunderland (US): Sinauer Associates. Judd, W. S. [& al. 2016], Campbell, C. S., Kellogg, E. A., Stevens, P. F. & Donoghue, M. J. (2016) Plant systematics: A phylogenetic approach. Ed. 4. Sunderland (US): Sinauer Associates. Mabberley, D. J. (2008) Mabberley’s Plant-Book. A portable dictionary of plants, their classification and uses. Ed. 3. Cambridge (GB): Cambridge University Press Moore, M. J. [& al. 2010], Soltis, P. S., Bell, C. D., Burleigh, J. G. & Soltis, D. E. (2010) Phylogenetic analysis of 83 plastid genes further resolves the early diversification of eudicots. Proc. Nation. Acad. Sci. USA 107(10): 4623–4628. https://doi.org/10.1073/pnas.0907801107. Nordal, I. & Sletten Bjorå, C. (2016) Family delimitation in flowering plants, should we bother? - A case study of Asparagales. Symb. Bot. Upsal. 38: 41–48. 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 Pires, J. C. [& al. 2006], Maureira, I. J., Givnish, T. J., Sytsma, K. J., Seberg, O., Petersen, G., Davis, J. I., Stevenson, D. W., Rudall, P. J., Fay, M. F. & Chase, M. W. (2006) Phylogeny, genome size, and chromosome evolution of Asparagales. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 287–304. https://doi. org/10.5642/aliso.20062201.24. Schmid, R. (2013) Considerations for flora writers in adopting the new phylogenetic classifications. Taxon 62: 206–207. https://www.jstor.org/stable/24389345 Seberg, O. [& al. 2012], Petersen, G., Davis, J. I., Pires, J. C., Stevenson, D. W., Chase, M. W., Fay, M. F., Devey, D. S., Jorgensen, T., Sytsma, K. J. & Pillon, Y. (2012) Phylogeny of the Asparagales based on three plastid and two mitochondrial genes. Amer. J. Bot. 99(5): 875–889, ills. https://doi.org/10.3732/ajb.110468. Soltis, D. E. [& al. 2011], Smith, S. A., Cellinese, N., Wurdack, K. J., Tank, D. C., Brockington, S. F., Refulio-Rodriguez, N. F., Walker, J. B., Moore, M. J., Carlsward, B. S., Bell, C. D., Latvis, M., Crawley, S., Black, C., Diouf, D., Xi, Z., Rushworth, C. A., Gitzendanner, M. A., Sytsma, K. J., Qiu, Y.-L., Hilu, K. W., Davis, C. C., Sanderson, M. J., Beaman, R. S.,

6 Olmstead, R. G., Judd, W. S., Donoghue, M. J. & Soltis, P. S. (2011) Angiosperm phylogeny: 17 genes, 640 taxa. Amer. J. Bot. 98(4): 704–730. https://doi.org/ 10.3732/ajb.1000404. Steele, P. R. [& al. 2012], Hertweck, K. L., Mayfield, D., Mckain, M. R., Leebens-Mack, H. & Pires, J. C. (2012) Quality and quantity of data recovered from massively parallel sequencing: Examples in Asparagales and

R. Nyffeler and U. Eggli Poaceae. Amer. J. Bot. 99(2): 330–348. https://doi. org/10.3732/ajb.1100491. Stevens, P. F. (2001+) Angiosperm Phylogeny Website, Version 12, July 2012 [and more or less continuously updated since]. St. Louis (US): Missouri Botanical Garden. http://www.mobot.org/MOBOT/research/ APweb/

Part I The Family Agavaceae

Agavaceae J. Thiede and U. Eggli

Including Funkiaceae Horaninov (nom. illeg., Art. 18.3). Including Yuccaceae J. Agardh. Including Hesperocallidaceae Traub. Including Hostaceae B. Mathew. Including Chlorogalaceae Doweld & Reveal. Small to often large perennial Ros plants or (sometimes bulbous) herbs, or shrubs or trees, Ros monocarpic, either unbranched and plant monocarpic, or branched and whole plant polycarpic, terrestrial, very rarely epiphytic; stems none or short (then not rarely caespitose), or  arborescent (then mostly  branching), in part with secondary thickening growth (Furcraea, Yucca), partly with spreading or thick and upright rhizomes (Yucca p.p., Agave Subgen. Manfreda p.p., herbaceous non-succulent taxa); L spirally arranged in Ros, dorsiventral, lanceolate, linear or subulate, often broadest near the base or the middle and gradually tapering towards the tip, deciduous or more commonly evergreen and then usually thick and succulent, tough and fibrous or  thin and weak (Agave Subgen. Manfreda,

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

herbaceous non-succulent taxa), tip often a hard pungent spine (Yucca, Agave s.s.) or a  soft (more rarely hard) point (Beschorneria, Furcraea, Agave Subgen. Manfreda) or not differentiated, margins entire, hardened in most perennial taxa but soft in herbaceous taxa, or with horny marginal teeth (these often on prominences), or filiferous; Inf terminal, mostly large, 0.5–13 m, with few or numerous bracts, determinate and mostly complex much-branched panicles with cymose lateral part-Inf consisting of monochasial units; peduncle mostly present, more rarely short to nearly none; peduncular Bra mostly  similar to rosette leaves, diminishing in size upwards; floral Bra present; Ped normally present; Fl bisexual, 3-merous, actinomorphic or slightly zygomorphic, pendent or spreading, or  upright (Agave), anthesis diurnal and / or nocturnal; Tep 3 + 3, mostly whitish to yellow or greenish, rarely  reddish, pinkish or brownish, or lavender or blue, Tep  spreading or connivent to form a tube-like structure (Beschorneria, Hesperaloe p. p.), or fused for part of their length and forming a tubular to campanulate tube (Agave), sometimes abruptly widened or urceolate in the upper part; St 3 + 3, inserted at or somewhat above the base of the free tepals or within the perianth tube; Fil mostly long and slender-filiform, rarely short, sometimes widened basally (Furcraea) or apically (Yucca), mostly glabrous, sometimes puberulent (Yucca p.p.); Anth dorsifixed, 2-thecous, small to large, sagittate to hastate (Yucca, etc.) or oblong to linear (Agave, etc.), long or short,

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_112

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dehiscing introrsely with longitudinal slits; pollen primarily sulcate, rarely bisulcate (Agave Subgen. Manfreda p.p.), mainly released in monads, rarely also in tetrads (Beschorneria, Furcraea); Ov inferior (Agaveae) or superior (all other clades), 3-locular, each locule with 2 to several to mostly many ovules; Nec septal; Sty rather long and simple or apically with 3 free and short branches (Yucca); Sti mostly single and either capitate or 3-lobed or on 3 style branches (Yucca), surface dry or wet; Fr mostly loculicidal capsules, rarely septicidal capsules or berry-like (Yucca p.p.), with several to many seeds; Se various but mostly flat and plate-like, crescent-shaped or semicircular, but sometimes less compressed (Yucca), or globose, ovoid or pear-shaped (Chlorogaleae, Schoenolirion), black (due to phytomelans). — Cytology: x = 30 in the Agavoideae (bimodal, with 25 short and 5 long chromosomes) and Hosta (bimodal, with 4 large, 2 medium and 24 small chromosomes), or variable (n = 12, 15, 16, 17, 18, 26, 30, 48?) in the other clades (chromosomes all of equal length, or bimodal in Hesperocallis). Order: Asparagales Important Literature: Krause (1930: outdated synopsis); Schulze (1984: palynology, classification); Dahlgren & al. (1985: classification); Eguiarte & al. (1994: classification); Bogler & Simpson (1995: molecular phylogeny); Bogler & Simpson (1996: molecular phylogeny); Verhoek-Williams (1998: synopsis); Bogler & al. (2006: molecular phylogeny); Givnish & al. (2006: molecular phylogeny); Good-Avila & al. (2006: molecular phylogeny); Graham & al. (2006: molecular phylogeny); Rocha & al. (2006: molecular phylogeny & pollination); Smith & al. (2008: molecular phylogeny); Kim & al. (2010: molecular phylogeny); Seberg & al. (2012: molecular phylogeny); Steele & al. (2012: molecular phylogeny); Matiz & al. (2013: photosynthesis); Halpin & Fishbein (2013: molecular phylogeny Chlorogaloideae); Archibald & al. (2015: molecular phylogeny Chlorogaloideae); Heyduk & al. (2016: molecular phylogeny & photosynthesis); McKain & al. (2016: molecular phylogeny).

Distribution: S Canada, USA, Mexico (= distribution centre), C America to Panama, Caribbean Region, Colombia, Venezuela, Peru, Bolivia, E Asia (Hosta); widely cultivated throughout the world in suitable climates and often also naturalized. The Agavaceae s.l. are a family of 12 genera and some 445 species. Most are adapted to semiarid or arid conditions, and the majority are xeromorphic and somewhat to distinctly succulent rosette plants of semidesertic regions and drylands. Classification: The genera placed in Agavaceae s.s. have a complicated classification history (Nyffeler & Eggli 2010). Up to the 1950ies, Agave and the closely related genera with inferior ovaries were placed in Amaryllidaceae, while Yucca and superficially similar genera with superior ovaries were classified in Liliaceae. At times, superficially similar xeromorphic taxa such as Dracaena or Nolina (now placed in Ruscaceae s.l.) were also included in Agavaceae, e.g. by Hutchinson (1934), Matuda & Piña Luján (1980), Rowley (1987) or Verhoek-Williams & Hess (2002). In traditional classifications (e.g. VerhoekWilliams (1998)), Agavaceae s.s. are composed of 8 genera of mostly xerophytic to succulent species. In contrast to this narrow circumscription, already the early molecular phylogenies of Eguiarte & al. (1994), Bogler & Simpson (1995), Eguiarte (1995) and Bogler & Simpson (1996) showed a close relationship of Agavaceae s.s. with Hosta (traditionally classified as monogeneric Hostaceae; not succulent, Kubitzki (1998)), either as sister group, in a basal unresolved position, or nested within. The inclusion of Hosta is also supported by cytological data (Tamura 1995) as well as embryological (Cave 1948), serological (Chupov & Kutiavina 1981) and palynological data (Schulze 1984). Hesperocallis (monotypic Hesperocallidaceae; not succulent, from North America, treated as anomalous genus in Hostaceae by Kubitzki (1998: 260)) and Chlorogalum and related genera (traditionally classified as Hyacinthaceae subfam. Chlorogaloideae, Speta (1998)) are also associated with Agavaceae s.l. in various recent molecular phylogenies. According to the molecular phylogenies of

Agavaceae

Seberg & al. (2012) and Steele & al. (2012). Agavaceae should be treated in a wide sense and include not only Hosta, but also Anemarrhena (monotypic Anemarrhenaceae; not succulent, from China and Korea, see Conran & Rudall (1998)), Behnia (monotypic Behniaceae; not succulent, from SE Africa, see Conran (1998a)) and Herreria (bitypic Herreriaceae; not succulent, from S America and Madagascar, see Conran (1998b)), although support for the various topologies is low. The most recent well-sampled study is that of Archibald & al. (2015). These authors found the Chlorogaloideae as unambiguous (though not monophyletic in the traditional circumscription) part of Agavaceae, confirming the earlier expressed views as well as the results of Pfosser & Speta (1999). Behnia and Herreria, however, remain unplaced in that phylogeny, and for the time being should either be included in an expanded Anthericaceae, or probably best be accepted as separate families in sister-group position (Petersen & al. 2006: Fig. 2). Whether the Anthericaceae should also be treated as part of Agavaceae s.l. has not yet been fully clarified. While Graham & al. (2006), Givnish & al. (2006) and Kim & al. (2010) prefer to include them in a further enlarged Agavaceae s.l., Pfosser & Speta (1999), Good-Avila & al. (2006), Bogler & al. (2006) and Smith & al. (2008) stress the lack of support for this and prefer to treat the Anthericaceae as separate family in sister-group position to Agavaceae s.l. This is also the position followed in this handbook. There is no doubt that the Agavaceae s.l. belong to the higher core Asparagales, together with Asparagaceae s.s., Anthericaceae, Hyacinthaceae and Ruscaceae, to name just the families with succulent representatives. The families of the higher core Asparagales have recently been merged and treated as Asparagaceae s.l. (Janssen & Bremer 2004, Graham & al. 2006), and this classification is that advocated by APG (2009), and confirmed by APG (2016). Asparagaceae s.l. in that circumscription would be diagnostically circumscribed by racemose inflorescences (vs. umbellate inflorescences in the lower core Asparagales). Within the framework of such a classification, Agavaceae s.l. would be classified at the rank of subfamily as

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Agavoideae. However, support for some of the proposed relationships is either low or unconfirmed in recent molecular phylogenies (e.g. Kim & al. (2010)). Moreover, accepting Asparagaceae in the very broad sense of APG does not provide improved insights into the phylogeny of the order, but is unnecessarily disruptive to established traditional classifications (Nyffeler & Eggli 2010). Therefore, and in line with these authors and the recent textbook by Judd & al. (2016), Agavaceae in their new expanded circumscription is here treated as independent family. While Judd & al. (2016) also include the Anthericaceae, we maintain that support for this placement is insufficient, and accept Anthericaceae as separate family in this Handbook. Agavaceae s.l. is sister to Aphyllanthaceae + Themidaceae according to Seberg & al. (2012), while Steele & al. (2012) found just Aphyllanthaceae (monotypic, not succulent, W Mediterranean) as the more likely sister group. This is confirmed by the plastid phylogenomic study of Givnish & al. (2018) who also found Aphyllanthes as most likely sister of Agavaceae (as Agavoideae, including Anthericaceae), but support is low. The clade age of the expanded Agavaceae has been estimated at a mere  14.5 mybp but could also be 23.7  9.7 mybp (Smith & al. 2008a). The monocot-wide plastid phylogenomic study of Givnish & al. (2018) found an age of 24.51 mybp for the split of the expanded Agavaceae from Anthericaceae (including Herreria). This is much younger than the age of  35 mybp given by Wikström & al. (2001) for the split of Agave from (Behniaceae + Anthericaceae). The major radiations in Agave (starting 7.8–10.1 mybp) and Yucca (starting 13–18 mybp) of crown-group Agavaceae (Good-Avila & al. 2006, Rocha & al. 2006) fall into a time of increasing desertification of North America. The origin of the family in its presently wide circumscription is supposed to have been in W North America (Bogler & al. 2006). Classification of crown group Agavaceae: The crown group of Agavaceae is co-extensive with the “Agave+”-clade of Archibald & al. (2015) and is largely congruent with Agavaceae s.s. as covered in the first edition of this Handbook.

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J. Thiede and U. Eggli

Ch

Ch

Agavoideae Hosteae

SHH

Yucceae

Hosta

1 2 3

Yucca

Agaveae B

Fur

Agave

50 species Fig. 1 Summary phylogeny of Agavaceae s.l. mainly based on Archibald & al. (2015) (445 species total). Ch = Chlorogaloideae, 1 = Hesperoyucca, 2 = Hesperaloe;

3 = Schoenolirion; B = Beschorneria, Fur = Furcraea; SHH = SHH clade. (Copyright: U. Eggli)

Archibald & al. (2015) show this clade in a trichotomy with Hosta on the one hand, and their SHH-clade (Hesperaloe and Hesperoyucca plus the non-succulent Schoenolirion). – Fig. 1. Although the various molecular phylogenies cited above are not in full agreement about the topology of the genera that make up the “Agave +”-clade, there are good indications that Yucca is the sister of an Agave-clade that is composed of Beschorneria as sister to Agave + Furcraea. The remaining genera traditionally accepted in Agavaceae, i.e. Manfreda, Polianthes and Prochnyanthes, are all derived from within Agave s.s. (Bogler & al. 2006, Rocha & al. 2006). They cannot be separated from Agave s.s. on molecular grounds, and if they were accepted as separate genera, several clades within Agave s.s. would have to be recognized as separate genera as well to attain a monophyletic classification. Such a classification appears clearly unwarranted before the background of the frustratingly low sequence variation for Agave (Bogler & al. 2006) and the segregate Manfreda (Ritchie 2014, almost no variation at all amongst 19 specimens studied).

In addition, Manfreda can be successfully hybridized with Agave (= Mangave) (Klein 2010) and Polianthes (= Polifreda) (Lindstrom 2006a, Lindstrom 2006b, Ritchie & Lindstrom 2014). We interprete this as additional support for subsuming these genera under Agave. The mesomorphic characters exhibited by the species of Manfreda, Polianthes and Prochnyanthes have thus evolved secondarily from more arid-adapted evergreen taxa. The study of McKain & al. (2016) arrives at the same topology, but is overall of limited value since only single terminals were analyzed. Agavaceae s.l. can thus conveniently be classified into the following monophyletic clades – Fig. 1: • Chlorogaloideae Lotsy 1911 (= “core Chlorogaloideae” of Archibald & al. (2015); incl. Hesperocallidaceae Moore & Reveal 2012): Bulbous herbs; L keeled; Inf (double) racemes; Tep white, greenish or blue; Ov superior, with 2 or numerous (Hesperocallis) ovules per locule; Sti 3-lobed; Se pear-shaped to globose or ovoid; n = 15, 17, 18, 24 (Hesperocallis), 26, 30, bimodal in

Agavaceae

Hesperocallis (5–6 large, 18–19 small chromosomes). — 16 species (not succulent) in the 4 genera Camassia, Chlorogalum, Hastingsia and Hesperocallis from N America. • Hosta-clade (= Hosteae Traub 1972; = Hostaceae B. Mathew 1988): Rhizomatous, usually deciduous herbs; L linear-lanceolate to cordate-ovate, often long-petiolate, distinctly veined; Inf bracteate racemes, often  secund; Tep white, lavender or blue; Fil declinate; Ov superior; Se flattened, winged; x = 30, bimodal with 4 large, 2 medium and 24 small chromosomes. — Only genus Hosta (40 species, not succulent) from Japan, China, Korea and Russia. • SHH-clade (Archibald & al. 2015): Ov superior; Sti  capitate to slightly lobed. — This clade is morphologically heterogeneous: • Schoenolirion: Perennial shortly rhizomatous herbs with or without bulb; L grasslike, flat, without keel; Inf to 1.5 m; Tep white or yellow; Se  globose; x = 12, 15, 16, 24(?). — 3 species (not succulent) from N America. • Hesperaloe + Hesperoyucca: Perennial xerophytes with or without (Hesperoyucca) rhizomes and with monocarpic rosettes; L few to very numerous, tough, more or less to distinctly succulent, fibrous, rounded or keeled on both faces; Inf to 4.5 m, racemose-paniculate; Tep greenish-white, white, red to salmon or yellow; Se flat; x = 30, bimodal. — Composed of the genera Hesperaloe and Hesperoyucca, treated below in this Handbook. • Agavoideae = “Agave + clade” of Archibald & al. (2015): Mostly evergreen and L mostly fibrous and succulent, rarely hardly succulent and xerophytic, or herbaceous-succulent and  deciduous; Ov superior or inferior. — Divisible in 2 subclades: • Tribe Yucceae Bartlett 1930 (incl. subfam. Yuccoideae Kostelezky 1831; incl. Yuccaceae J. Agardh 1858): L xeromorphic, fibrous, not strongly succulent (except Y. endlichiana); L margins mostly filiferous, rarely serrulate; Fl pendent, actinomorphic; Tep free; Ov superior; Anth sagittate to

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hastate. — Only genus: Yucca, treated below in this Handbook. • Tribe Agaveae (incl. tribe Poliantheae Hutchinson 1934): L mostly xerophytic, evergreen and succulent, rarely deciduous and herbaceous and soft; L margins entire and often with teeth, or rarely filiferous; Fl pendent or  upright, actinomorphic or slightly zygomorphic; Tep free or  fused to form a tube; Ov inferior; Anth oblong to linear. — Consists of the genera Agave, Beschorneria and Furcraea, all treated below in this Handbook. Cytology: The striking bimodal karyotype (McKelvey & Sax 1933) with x = 30 represents a major distinguishing feature for Agavaceae s.s. Similar, but less pronounced bimodal karyotypes are present in Hesperocallis (x = 24) and Hosta (x = 30), but not in the other members of the Agavaceae s.l. According to McKain & al. (2012) the bimodal karyotype is due to two independent palaeopolyploidization events (identified as wholegenome duplication by McKain & al. (2016)), one predating the last common ancestor of Agave and Hosta, the other in the lineage that includes Anthericaceae. Subsequently, the bimodal karyotype was lost twice independently, once in the Hesperocallis - Chlorogalum - Camassia clade, once in Schoenolirion. Polyploidy is common esp. in species of Agave subgen. Agave. Ploidy ranges from diploid to octoploid, including (sterile) triploids (e.g. A. angustifolia var. letonae) and pentaploids (e.g. A. sisalana) (Robert & al. 2008). Polyploidization, at least in Agave, is likely of a comparatively young age (Robert & al. 2008) and probably linked to increased vegetative propagation and/or selected for by the conditions of an increasingly arid and harsh environment. Robert & al. (2008) did not find any dysploid changes, and early chromosome counts that deviate from the basal x = 30 are likely observation errors. Succulence: Leaf succulence in varying degrees prevails in the family, but stem succulence in the form of thickened rhizomes, pachycaulous stems, or subterranean corms, also occurs. The strongly succulent leaves of many Agave species show

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well-developed tissue succulence, visible in cross sections as a narrow peripheral band of chlorenchyma, and voluminous chlorophyll-free water storage parenchyma (hydrenchyma) with interspersed scattered fibrous vascular bundles (see Müller (1909) for illustrations). The demarcation between chlorenchyma and hydrenchyma is not very strict. Already Bray (1903) found the hydrenchyma to contain air spaces, which cause the whitish colour of this tissue. The massive trunks of stem-forming Yucca species such as Y. gigantea are reminiscent in architecture to those of Beaucarnea (Ruscaceae) and consist of numerous vascular bundles dispersed in a matrix of ground tissue. Within the outer cortex, a secondary thickening meristem develops (Fisher 1975, Stevenson 1980, Carlquist 2012: 113). Physiology: Crassulacean Acid Metabolism (CAM) is almost always present in Agave s.s., Furcraea and Hesperaloe. Yucca is heterogenous with respect to photosynthetic pathway, with constitutive CAM in the species of sect. Yucca, and C3 in the species of the other sections (Nobel 1988, Gibson 1996: 118, Sayed 2001, Heyduk & al. 2016). Agave amica (Polianthes tuberosa) and Beschorneria appear to use the C3 pathway (Heyduk & al. 2016). Data for Hesperoyucca is ambigous: Winter & Smith (1996) report CAM, but Heyduk & al. (2016) indicate C3. The non-succulent species of the former Chlorogaloideae, Hosta and the SHH-clade also use the C3 pathway (Heyduk & al. 2016). Heyduk & al. (2016) argue for 3 independent origins of CAM. Olivares & Medina (1990) report that CAM signature varies over the length of the leaves as well as between the leaves of a rosette in Furcraea acaulis (as F. humboldtiana); CAM is least pronounced in the vertically upright youngest leaves, and in the leaf base of older leaves. Chemistry: Steroidal saponins are probably universally present throughout the Agavaceae s.l., sometimes in considerable concentrations. Simmons-Boyce & Tinto (2007) and Sidana & al. (2016) present an overview of steroidal saponins and sapogenins found in species of Agave,

J. Thiede and U. Eggli

Furcraea and Yucca. The most common compound classes are furastanes, furospirostanes, and spirostanes. Some of these compounds show considerable biological activity, including anticancer effects, and some are used as starting material for the biosynthesis of steroidal hormones. Ecology: Many species of Agave and Yucca as well as Hesperoyucca are landscape-dominating elements throughout the arid and semi-arid regions of Mexico and the SW, S, and SE USA. Some species form vast populations locally and these are a spectacular sight when mass flowering occurs. Pollination systems vary from very specialized to generalist. The most striking specialized pollination system is found in Yucca and Hesperoyucca, with a closely knit mutualistic relationship between the pollinating yucca moths and Yucca, sometimes described as “nursery pollination” (Svensson & al. 2011). Althoff (2016) and the chapter on Yucca in this Handbook provide overviews of extant knowledge. In Agave, bats and hawkmoths during the night as well as hummingbirds and other birds and/or bees during the day are known as pollinators, depending on the species. The floral syndromes displayed are not always predictive of the main pollinators (e.g. Silva-Montellano & Eguiarte (2003a) for A. lechuguilla, Riffell & al. (2008a) and Riffell & al. (2008b) for the generalist pollination systems of A. palmeri and A. chrysantha). Flowering A. palmeri form a veritable “nectar corridor” and its geographical range closely overlaps the range of its migratory bat pollinators. In C Mexico, Agave and bats appear to mutualistically depend on each other, while in N Mexico and SW USA, the interdependence is less strict (Riffell & al. 2008b). Ethnobotany: Many species esp. of Agave and to a lesser extent of Yucca have been used for food, beverage and fibre since prehistoric times by numerous local ethnic groups. The use of these plants can be dated back for at least 9000 years (Gentry 1982: 5) and has continued for centuries among many different and unrelated cultures (Hodgson 2001: 14–51).

Agavaceae

Gentry (1982: 3–24) termed the intimate relationship between these cultures and the plants “man-Agave symbiosis”. There is evidence that some species of Agave (e.g. A. parryi) were extensively cultivated in prehistorical times for both food and fibre (Parker & al. 2014), leading to domestication of selected variants. Indeed, many relict populations continue to exist at ancient dwelling sites, and domestication of Agave probably started several 1000 years BCE. A. murphyi is one of several ancient cultigens, and clones persist at prehistorically inhabited sites as “living archaeological assemblages preserved in their cultural landscape for more than five centuries after their presumed cultivation” (Hodgson 2001). Several species of Agave are used to manufacture distilled alcoholic beverages, and are the basis of the Mexican pulque, mezcal and tequila industries. Enormous areas are devoted to this purpose: Chagoya-Méndez (2004) (cited from Martínez-Gutiérrez & al. (2013)) report an area of 15.500 hectares just for the Mexican state of

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Oaxaca, planted with A. angustifolia and yielding an annual 2.9 million litres of mezcal. A number of species of Agave and Furcraea continue to be important sources of fibre, and are cultivated in large-scale plantations up to today in suitable climates throughout the world. Another commodity obtained from species of Agave is agave syrup. Horticulture: Numerous taxa of Agavaceae s.s. have considerable horticultural potential (see Starr (2012) and Moore (2016) for Agave, Boeuf (2007) and Ondrovic & Ondrovic (2016) for Yucca, as well as the more general works on the horticultural use of perennial succulent monocots by Irish & Irish (2000) or Jacquemin (2000) and Jacquemin (2001)). Many species such as A. americana are frequently planted as ornamentals in broad-scale landscaping in suitable frostfree regions, esp. with Mediterranean climates. Numerous cultivars are available in the trade, including variously variegated forms (for Yucca, see, e.g. Boeuf & al. (2010)). Numerous species

Key to the genera with succulents 1a L margins often filiferous, rarely serrulate but never toothed; Fl pendent, actinomorphic; Tep free; 2 Anth sagittate to hastate; Ov superior 1b L margins entire and without or with (often strong) teeth, rarely filiferous; Fl pendent or  upright, 4 actinomorphic or slightly zygomorphic; Tep free or usually forming a  long tube; Anth oblong to linear; Ov inferior 2a Ros stemless or with short to massive stem and then arborescent; L margins entire or filiferous, Yucca L epidermis without papillate cells; Fl usually white or whitish and wax-like; Fil free, apically swollen and outcurved, sometimes puberulent; Sty thickened with 3 short branches; Sti 3 with 2 lobes each (S Canada to Guatemala) 2b Ros stemless; L margins filiferous or finely serrulate, L epidermis with papillate cells over the 3 veins; Fil adnate to the base or lower parts of the tepals, not swollen, straight, glabrous; Sty short and slender; Sti distinctly capitate and fringed with papillae 3a Habit  grass-like with succulent leaves; Ros few-leaved; L margins filiferous; Tep connivent and Hesperaloe Fl therefore narrowly campanulate, whitish to reddish (S USA (Texas), N and C Mexico) 3b Habit Yucca-like; Ros many-leaved; L margins finely serrulate; Tep openly spreading, whitish Hesperoyucca (SW USA, NW Mexico) 5 4a L margins entire or toothed; Fl pendent, actinomorphic; Tep (almost) free; Fil filiform or basally swollen; Sty swollen and with 3 basal ridges, distally abruptly narrowed 4b L margins entire or toothed, or filiferous; Fl  upright, actinomorphic or slightly zygomorphic; Tep Agave basally fused to form a  long tube; Fil filiform; Sty basally not swollen (S USA to Colombia and Venezuela, Caribbean region) 5a Plants polycarpic; L margins entire or minutely denticulate; Tep free but connivent and forming a Beschorneria tube-like structure, reddish to yellowish; Fil filiform (Mexico) 5b Plants monocarpic or polycarpic when branched; L margins mostly toothed; Tep openly spreading, Furcraea whitish to greenish; Fil basally swollen (Mexico to Bolivia, Caribbean region)

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have become naturalized outside their native occurrence, and today form an important feature of the landscape, often together with species of Opuntia (esp. O. ficus-indica, Cactaceae), which has a similar New-World origin.

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J. Thiede and U. Eggli Soc. 158(2): 215–222, ill. https://doi.org/10.1111/ j.1095-8339.2008.00831.x. Rocha, M. [& al. 2006], Good-Ávila, S. V., MolinaFreaner, F., Arita, H. T., Castillo, A., García-Mendoza, A., Silva-Montellano, A., Gaut, B. S., Souza, V. & Eguiarte, L. E. (2006) Pollination biology and adaptive radiation of Agavaceae, with special emphasis on the genus Agave. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 329–344, maps. https://doi.org/ 10.5642/aliso.20062201.27. 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. Schulze, W. (1984) Beiträge zur Taxonomie der Liliifloren: XII. Der Umfang der Agavaceae. Wiss. Z. FriedrichSchiller-Univ. Jena, Math.-Naturwiss. Reihe 32(6): 965–979, ills., maps. Seberg, O. [& al. 2012], Petersen, G., Davis, J. I., Pires, J. C., Stevenson, D. W., Chase, M. W., Fay, M. F., Devey, D. S., Jørgensen, T., Sytsma, K. J. & Pillon, Y. (2012) Phylogeny of the Asparagales based on three plastid and two mitochondrial genes. Amer. J. Bot. 99(5): 875–889, ills. https://doi.org/10.3732/ ajb.1100468. Sidana, J. [& al. 2016], Singh, B. & Sharma, O. P. (2016) Saponins of Agave: Chemistry and bioactivity. Phytochemistry 130: 22–46. https://doi.org/10.1016/ j.phytochem.2016.06.010. Silva-Montellano, A. & Eguiarte, L. E. (2003a) Geographic patterns in the reproductive ecology of Agave lechuguilla (Agavaceae) in the Chihuahuan Desert. I. Floral characteristics, visitors, and fecundity. Amer. J. Bot. 90(3): 377–387, map, ill., diags. https://doi.org/ 10.3732/ajb.90.3.377. Simmons-Boyce, J. L. & Tinto, W. F. (2007) Steroidal saponins and sapogenins from the Agavaceae family. Nat. Prod. Commun. 2(1): 99–114. Smith, C. I. [& al. 2008], Pellmyr, O., Althoff, D. M., Balcázar-Lara, M., Leebens-Mack, J. & Segraves, K. A. (2008) Pattern and timing of diversification in Yucca (Agavaceae): Specialized pollination does not escalate rates of diversification. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 275: 249–258. https://doi. org/10.1098/rspb.2007.1405. Speta, F. (1998) Hyacinthaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 261–285, ills., key. Berlin (DE) etc.: Springer Verlag. https://doi.org/ 10.1007/978-3-662-03533-7_35. Starr, G. (2012) Agaves. Living sculptures for landscapes and containers. Portland (US)/London (GB): Timber Press. Steele, P. R. [& al. 2012], Hertweck, K. L., Mayfield, D., McKain, M. R., Leebens-Mack, H. & Pires, J. C. (2012) Quality and quantity of data recovered from massively

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Agave AGAVACEAE J. Thiede

Agave Linné (Spec. Pl. [ed. 1], 323, 1753). Type: Agave americana Linné [lectotype, designated by Britton & P. Wilson, Sci. Survey Puerto Rico, 5: 156, 1923 (fide ING)]. — Agavoideae — Lit: Trelease (1913: synopsis West Indies); Trelease (1915: synopsis Guatemala); Berger (1915: general synopsis); Hummelinck (1936: synopsis Caribbean); Hummelinck (1938: synopsis Caribbean); Gentry (1972: monograph Sonora); Verhoek-Williams (1975: revision Subgen. Manfreda); Gentry (1982: monograph USA, Mexico, C America); Piña Luján (1985: synopsis Subgen. Manfreda); Piña Luján (1986: synopsis Subgen. Manfreda); McVaugh (1989: Fl. Novo Galiciana); Hummelinck (1993: synopsis Caribbean); Lott & García-Mendoza (1994: Fl. Mesoamerica); Solano Camacho (2002: revision Polianthes); Heller (2006: ill. synopsis cultivated taxa); Castillejos-Cruz (2009: revision Manfreda); Solano & Ríos-Gómez (2011: key to Sect. Polianthes in Jalisco); Solano & al. (2014: anatomy Polianthes); Castro-Castro & al. (2015: key to Sect. Polianthes in Nueva Galicia); Guillot Ortiz (2015: key to naturalized taxa in Spain); Mottram (2015: nomenclature infrageneric names); Sidana & al. (2016: review steroidal saponins); CastroCastro & al. (2016: key to Polianthes, flower ills.); Boeuf & al. (2017: ill. synopsis); Thiede &

J. Thiede (*) Hamburg, Germany e-mail: [email protected]

al. (2019: sectional classification). Distr: S USA, Mexico, C America to Panama, whole Caribbean region, Colombia, Venezuela; cultivated worldwide in tropical and subtropical to frost-free temperate climates and often naturalized. Etym: Gr. ‘Agave’, daughter of Kadmos and sister of Semele in Gr. mythology, also the mother of Pentheus, who she murdered in an outburst of fury; also Gr. ‘agavos’, stately, noble, illustrious; for the stately nature of many species, but also for the ferocious leaf margin teeth present in many species. Incl. Polianthes Linné (1753). Type: Polianthes tuberosa Linné [typification by inference, only element included]. Incl. Pothos Adanson (1763) (nom. illeg., Art. 53.1). Type: not typified. Incl. Tuberosa Heister ex Fabricius (1769) (nom. illeg., Art. 52.1). Type: Polianthes tuberosa Linné. Incl. Bonapartea Willdenow (1814) (nom. illeg., Art. 53.1). Type: Bonapartea juncea Willdenow. Incl. Littaea Tagliabue (1816). Type: Littaea geminiflora Tagliabue. Incl. Bravoa Lexarza (1824). Type: Bravoa geminiflora Lexarza [typification by inference, only element included]. Incl. Coetocapnia Link & Otto (1828). Type: Coetocapnia geminiflora Link & Otto [typification by inference, only element included]. Incl. Robynsia Drapiez (1841) (nomen rejiciendum, Art. 56.1). Type: Robynsia geminiflora Drapiez

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_111

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[typification by inference, only element included]. Incl. Ghiesbreghtia Roezl (1861) (nom. illeg., Art. 53.1). Type: not typified. Incl. Manfreda Salisbury (1866). Type: Agave virginica Linné. Incl. Allibertia Marion (1882) (nom. illeg., Art. 53.1). Type: Allibertia intermedia Marion [typification by inference, only element included]. Incl. Prochnyanthes S. Watson (1887). Type: Prochnyanthes viridescens S. Watson. Incl. Leichtlinia H. Ross (1893). Type: Agave protuberans Engelmann ex Baker [typification by inference, only element included]. Incl. Delpinoa Ross (1897). Type: Delpinoa gracillima Ross [typification by inference, only element included]. Incl. Pseudobravoa Rose (1899). Type: Bravoa densiflora B. L. Robinson & Fernald. Incl. Runyonia Rose (1922). Type: Runyonia longiflora Rose [typification by inference, only element included]. Incl.
Mangave D. Klein (2010). Type: not typified. Incl.
Polifreda W. Ritchie (2014). Type: Polyfreda ‘Lindstrom’. Perennial xerophytic Ros plants, mostly L succulents, mono- or polycarpic but individual rosettes always monocarpic, terrestrial (very rarely epiphytic); R tough and fibrous, sometimes fusiformly thickened (Subgen. Manfreda), shallowly radiating; Ros acaulescent or stems short, rarely elongated (e.g. A. attenuata ssp. dentata), mostly thick, solitary or branched, sometimes rhizomatous (esp. in Subgen. Manfreda); Ros small to very large; L mostly long-lived, predominantly succulent and xeromorphic, more rarely  soft and annual (Subgen. Manfreda p.p.),  thick and fibrous, linear to lanceolate to ovate, usually rich in steroidal sapogenins, tip a terminal  strongly developed Sp or a soft point (Subgen. Manfreda); L margins entire, minutely to strongly toothed (then teeth partly on prominences of the L margin), or filiferous; Inf paniculate, small to up to 12 m tall (= complete Inf, i.e. peduncle and floriferous part); part-Inf cymose, consisting of monochasial units, either  short-stalked and

J. Thiede

mostly with few (often paired) flowers (Subgen. Littaea), or long-stalked, often several times compound, with many  densely arranged flowers (Subgen. Agave), or without a stalk and paired to single flowers (Subgen. Manfreda); Inf sometimes bulbilliferous, esp. in species of anthropogenic origin; peduncular Bra  similar to rosette leaves, diminishing in size upwards; Ped long to short or nearly none (esp. Subgen. Manfreda); Fl diurnal and/or nocturnal, actinomorphic or (slightly) zygomorphic (Subgen. Manfreda p.p., rarely in Subgen. Littaea), nearly always proterandrous (proterogynous in A. polianthiflora); Per tubular to campanulate, usually yellow or greenish to brownish, more rarely reddish, rarely white or pink (Subgen. Manfreda p.p.); Tep basally usually fused and forming a perianth tube, much varying in length, lobes of varying length; St usually exserted, but included in Sect. Polianthes; Fil usually filiform, normally  long, rarely  short, inserted in the tube or at the mouth of the tube or at the tepal base; Anth versatile, oblong, pollen released in monads, sulcate (rarely bisulcate); Ov inferior, thick-walled, 3-locular, with numerous axile ovules in 2 rows per locule, often constricted above to a  conspicuous neck; septal Nec present; Sty elongate, filiform, tubular, not yet fully expanded at anthesis; Sti 3-lobed, papillate-glandular; Fr dehiscent loculicidal capsules, apically often beaked (i.e. narrowly elongated); Se flattened, black. — Cytology: x = 30, with multiples present. The genus includes 265 species (plus several naturally occurring or artificial hybrids) of leaf succulents (only some species of Subgen. Manfreda are weakly succulent) of considerable ethnobotanical, economical, ecological, and horticultural importance. Many of its species are landscapedominating elements esp. in the Mexican semiarid and arid biomes. Classification History: Agave taxonomy is difficult due to the complex nomenclature with many old names of uncertain application and many poorly or even undocumented names of horticultural origin. Additional difficulties are caused by the pronounced variability and plasticity of

Agave AGAVACEAE

many taxa, including possible hybridization and introgression. Haworth (1812: 70–72) published a first rudimentary infrageneric classification of Agave based on leaf colour with 2 groups “Foliis glaucis” vs. “Foliis viridibus, vel virescentibus”. Salm-Dyck (1834: 7–8) classified Agave into 3 informal groups: Caulescentes, Acaules, and Herbaceae (which includes 2 species of Subgen. Manfreda) and later (Salm-Dyck 1859: 86–87) recognized 5 unranked taxa (“§”) based on leaf characters. In a subsequent publication, SalmDyck (1861) amended the classification and used the rank of section for these groupings, which is apparently the first formally ranked classification of the genus. Jacobi (1864: 498–501) proposed a classification based on leaf characters into 4 major unranked groupings (“Hauptgruppen”), the largest of which (Keratacanthae) is subdivided into 7 subgroups. Koch (1865) was apparently the first to use inflorescence types and proposed an informal classification into 2 divisions (“Abtheilungen”) (“Agavae paniculatae” and “Agavae spicatae”), subdivided into groups (“Gruppen”) based on leaf characters. Engelmann (1875: 296–297) provided apparently the first formal classification of Agave based on inflorescence types, proposing 3 sections: A. Sect. Singuliflorae with solitary flowers (conforming to the modern Subgen. Manfreda), A. Sect. Geminiflorae with long and narrow ‘subspicate’ panicles (conforming to Subgen. Littaea), and A. Sect. Paniculatae with broad panicles (conforming to Subgen Agave). Baker (1877a: 171), Baker (1877b: 808), and Baker (1888a: 164) subdivided Agave into 3 subgenera based on inflorescence types corresponding to Engelmann’s 3 sections: A. Subgen. Euagave (= Agave), A. Subgen. Littaea, and A. Subgen. Manfreda. In a simultaneous alternative classification based on leaf characters, he (Baker 1877b: 808, Baker 1888a: 164–166) divided Agave into 4 series, 3 of which are subdivided into 13 informal groups. Terracciano (1885) recognized 2 subgenera, Cladagave (conforming to Subgen. Agave), and Aplagave (conforming to Subgen. Manfreda + Subgen. Littaea), the latter further subdivided into Engelmann’s Sect. Singuliflorae

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and Sect. Geminiflorae, both further divided into subsections. Berger (1898) first largely followed Baker’s classification. In his later monograph, Berger (1915) recognised 3 subgenera (Manfreda, Littaea, Euagave (= Agave)) and subdivided Subgen. Littaea into 7 formal sections, while Subgen. Euagave was divided into 18 informal “Reihen”. Berger’s classification became rather influential because it was adopted in Jacobsen’s handbooks (e.g., Jacobsen 1954: 78–89). Trelease (1913), in his monograph of the Caribbean species, established 7 informal groups widely used up to today. The highly influential monograph of the continental North American species of the genus by Gentry (1982) recognized 2 subgenera only (Littaea and Agave), thus excluding Manfreda from Agave s.s. Both subgenera are subdivided into informal groups “tantamount to sections” (Gentry 1982: xi); these were mainly taken from Baker and Trelease or also established as new, and were and are widely used. Several of these informal groups were formalized at sectional rank e.g. by Ullrich (1993b: 53, 56: Sect. Crenatae) and Starr & Webb (2015). For the treatment of Agave in the first edition of this handbook, the present author converted molecular phylogenetic results that showed Manfreda, Polianthes and Prochnyanthes to be nested within Agave s.s. into a revised, monophyletic circumscription of Agave s.l. that included Manfreda, Polianthes and Prochnyanthes in Agave, together classified as A. Subgen. Manfreda. Hochstätter (2015), in his Agave (s.s.) compilation, published many new names at sectional, series and subseries rank (many of the latter seem superfluous when there is only one series within a section or one subseries within a series). Mottram (2015) provided an annotated checklist of the infrageneric taxa of Agave. Classification: The circumscription of the infrageneric groupings applied in the first edition of this handbook is largely retained, but names at formal sectional rank for all infrageneric taxa are used. This classification applies, whenever available, names at sectional rank published earlier by

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Salm-Dyck (1861), Engelmann (1875), Bentham in Bentham & Hooker (1883), Berger (1915), Ullrich (1992l), Ullrich (1993b), Webb & Starr (2015), and Hochstätter (2015), including 3 sectional names recently published as new (Sect. Conicae, Sect. Intermediae, Sect. Hibernicae). Thiede & al. (2019) provide the necessary new combinations for those groupings for which no formal name at sectional rank has previously been published, and also provides typifications and a synopsis of the infrageneric classification of Agave. [1] Subgen. Littaea (Tagliabue) Baker 1888 ( Littaea Tagliabue 1816): Inf with  short-stalked to nearly sessile part-Inf mostly with few, mostly geminate (= paired) Fl, or Fl 3–4 (–8), rarely more or rarely single (‘spicate’ Inf): [A] ‘Weakly armed group’; L margins entire, serrulate, filiferous or with weak teeth: [a] L margins firm, not filiferous, L surface without white marks left by the central bud, striate, not soft, L margins finely serrulate; Tep tube well-developed; Ov without neck: [1a] Sect. Juncineae Salm-Dyck 1861 ( Group Striatae Baker 1877  Subsect. Striatae (Baker) A. Terracciano 1885  Sect. Schoenagave A. Berger 1915; incl. Abtheilung Loriformes Jacobi 1864  Sect. Chonanthagave A. Berger 1915): Plants perennial, often forming large clumps; L numerous, narrow, linear, hard, not softly succulent, usually striate, margins finely serrulate; Fl geminate; Tep forming a well-developed tube often longer than the lobes; Ov half-inferior, without neck, merging into a welldeveloped tepal tube; Fil usually inserted in the middle of the tube, rarely higher up, frequently at 2 levels. — 9 species in NE, C and S Mexico. Some species appear to flower repeatedly from “lateral” inflorescences which are actually likely terminal on rudimentary lateral rosettes. Molecular data (Bogler & Simpson 1996, Bogler & al. 2006, Gil-Vega & al. 2007) show

J. Thiede

that Sect. Juncineae is the earliest branch-off in Agave s.l. and sister to the remainder of the genus. Such a basal position coincides with its plesiomorphic morphological features, esp. as seen in A. dasylirioides (see the section on the phylogeny below, and under A. dasylirioides). [b] L not striate, margins firm, not filiferous, softly succulent, surface without white marks left by the central bud, margins smooth or irregularly serrulate; Tep tube short or none; Ov with a neck: [1b] Sect. Inermes Salm-Dyck 1861 ( Sect. Anoplagave A. Berger 1915  Group Amolae Gentry 1982 (nom. inval., ICN Art. 37.1); incl. Abtheilung Margine integerrimae Jacobi 1864 (nom. inval., ICN Art. 21.2 Ex. 2 & 32.1c); incl. Ser. Yuccifoliae A. Terracciano 1885  Sect. Yuccifoliae (A. Terracciano) B. Ullrich 1996; incl. Group Serrulatae Baker 1877; incl. Ser. Nizandensae B. Ullrich 1991  Sect. Nizandensae (B. Ullrich) B. Ullrich 1991): L soft, margins entire, unarmed, terminal Sp usually present, rarely soft or absent; Inf with densely arranged Fl; Tep tube shallow, much shorter than the lobes; Fil usually inserted at the mouth of the tepal tube. — 10 species in N, C and S Mexico (mainly Sierra Madre Occidental). [1c] Sect. Choritepalae Hochstätter 2015 (incl. Group Choritepalae Gentry 1982 (nom. inval., ICN Art. 37.1)): L soft to fleshy-leathery, without (A. bracteosa, A. ellemeetiana) or with terminal Sp (A. guiengolensis, A. gypsicola); Inf dense; Tep tube (nearly) lacking and Tep arising from a discoid receptacle, or very short; Fil inserted on the receptacle or at the base of the tube. — 4 species from N and S Mexico (Oaxaca). Regarded as artificial by Ullrich (1990b), and this is corroborated by the AFLP data of Gil-Vega & al. (2007) where the 3 studied taxa are not forming a monophyletic group.

Agave AGAVACEAE

[c] L margins filiferous (i.e. decomposing into white threads), L surface with white marks left by the central bud: [1d] Sect. Littaea (Tagliabue) Bentham 1883 ( Littaea Tagliabue 1816;  Sect. Xysmagave A. Berger 1915; incl. Group Filiferae Baker 1877  Subsect. Filiferae (Baker) A. Terracciano 1885): Ros small to medium-sized, 20–50 (–200) cm ∅; L numerous, narrow, unarmed, filiferous; Fl usually geminate, campanulate, 30–55 mm, Tep tube usually much (rarely slightly) shorter than the lobes; Fil usually inserted at the rim of the tube (rarely slightly below). — 8 species in N and C Mexico (mainly Sierra Madre Occidental). [1e] Sect. Parviflorae Hochstätter 2015 ( Group Parviflorae Gentry 1982 (nom. inval., ICN Art. 37.1)): Ros mostly small, compact, 12–60 (–80) cm ∅, solitary or freely suckering; L numerous, short (6–15 to very rarely exceeding 30 cm), narrow, unarmed, filiferous; Fl small, 13–42 (–50) mm; Tep forming a distinct tube; Fil shortly exserted, inserted usually deep in the tube. — 4 species from the N Sierra Madre Occidental (Mexico) and adjacent Arizona (USA). Starr & Devender (2011) supply a key to the species of the group. [B] ‘Strongly armed group’: L margins generally with large teeth (except some species of Sect. Micracanthae and Sect. Heteracanthae): [a] Plants mainly polycarpic; stems branched; Tep tube deep, lobes 1–2
as long as the tube: [1f] Sect. Micracanthae Salm-Dyck 1861 (incl. Abtheilung Subcarinatae Jacobi 1864;  Abtheilung Aloideae Jacobi 1868;  Sect. Anacamptagave A. Berger 1915; incl. Group Polycephalae Gentry 1982 (nom. inval., ICN Art. 37.1)): Ros usually surculose; L few, 13–36, broad, softly succulent, fleshy, margins usually with small, weak and closely set teeth or denticles; terminal Sp weak and flexible to strong; Tep tube often 3-angled

25

and  grooved, lobes 1–3
as long as the tube; Ov often 3-angled and  grooved. — 9 species from NE to E-C and S Mexico. A mostly mesophytic and predominantly tropical group. Many (or even all) species appear to flower repeatedly from “lateral” inflorescences which are actually likely terminal on rudimentary lateral rosettes. See Cházaro Basáñez & Vázquez-García (2013: 59) for a key. [b] Plants mainly monocarpic; stems simple; Ros often surculose; Tep tube shallow, lobes 2–6
as long as the tube (except A. pelona): [1g] Sect. Heteracanthae Salm-Dyck 1861 ( Sect. Geminiflorae Engelmann 1875; incl. Abtheilung Marginatae Jacobi 1864  Group Marginatae Baker 1877  Subsect. Marginatae (Baker) Terracciano 1885  Sect. Pericamptagave A. Berger 1915; incl. Sect. Brachysolenagave A. Berger 1915; incl. Sect. Bechtoldiae Hochstätter 2015): L stiff and straight, usually with conspicuous and continuous horny margins and with conspicuous marginal teeth; Fl usually small with short,  openly funnel-shaped Tep tube, lobes 3–8 (–10)
as long as the shallow tube, frequently involute around the filaments during or only after anthesis. — 25 species from the S USA (S New Mexico, S Texas), Mexico and Guatemala. [1h] Sect. Urceolatae Hochstätter 2015 ( Group Urceolatae Gentry 1982 (nom. inval., ICN Art. 37.1)): L with small or weakly attached marginal teeth; Ros usually small and in small tight clusters; Tep lobes broadly overlapping,  2.5–3
as long as the short, broadly funnelshaped Tep tube. — 1 species (A. utahensis) in the USA (NW Arizona, S Utah, Nevada, S California). [2] Subgen. Agave: Inf with long-stalked, often several times compound part-Inf with many  densely arranged flowers (‘umbellate’ part-Inf).

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[A] ‘Large-sized group’: Plants medium-sized to (very) large; Ros 1–2.5 m tall; L 1–2.5 m. — Continental Species: [a] L generally not ensiform but lanceolate to ovate, much less than 10
longer than wide; Fr  oblong: [2a] Sect. Agave ( Sect. Macracanthae Salm-Dyck 1861 (nom. inval., ICN Art. 22.1 & Ex. 2)  Abtheilung Carnosae Jacobi 1864 (nom. inval., ICN Art. 22.1 & Ex. 2)  Sect. Paniculatae Engelmann 1875 (nom. inval., ICN Art. 22.1 & Ex. 2)  Group Americanae Baker 1877 (nom. inval., ICN Art. 22.1 & Ex. 2)  Subsect. Americanae (Baker) A. Terracciano 1885 (nom. inval., ICN Art. 22.1 & Ex. 2)  Reihe Americanae A. Berger 1915 (nom. inval., ICN Art. 22.1 & Ex. 2); incl. Abtheilung Subcoriaceae Jacobi 1864): Ros medium-sized to large, (freely) surculose; stems short; L light glaucous-grey to pale green, marginal teeth well-developed; Inf axis with smaller chartaceous Bra; fertile part of the Inf usually linear to long-oval in outline, part-Inf several times compound, not crowded; Fl yellow, rather slender; Tep tube furrowed, lobes (much) longer than the tube, not wilting until after anthesis; Fil inserted mostly at or near mid-tube, rarely at the mouth. — 7 mainly cultivated species. [2b] Sect. Salmianae (A. Berger) Verloove & Thiede in Thiede & al. 2019 ( Reihe Salmianae A. Berger 1915;  Group Salmianae Gentry 1982 (nom. inval., ICN Art. 37.1)): Ros large, massive, usually closely and freely surculose; stems short, thick; L light green to green, mostly very large, very thick towards the base; Inf axis with large appressed imbricate fleshy Bra; Inf pyramidal to ellipsoid in outline, part-Inf widely branching, several times compound; Fl large, succulent; Tep tube broad, thick-walled, lobes longer than or  as long as the tube,

J. Thiede

dimorphic, becoming incurved when wilting after anthesis; Fil inserted at, above or below mid-tube, usually at 2 levels. — 3 mainly cultivated species. [2c] Sect. Sisalanae Thiede & Gideon F. Smith 2019 ( Unterreihe Sisalanae A. Berger 1915 (nom. illeg., ICN Art. 53.1)): Ros medium to large, usually freely surculose; stems short to elongate; L without marginal teeth, or teeth reduced to prickles or irregularly arranged; terminal Sp small and weak; Inf lax, frequently bulbilliferous; Tep tube as long as wide, as long as to equalling the lobes; Fil inserted at or near mid-tube. — 3 cultivated species. Ullrich (1990d) abandoned this group in the sense of Gentry (1982) as an artificial assemblage and rearranged its species in other groups. This is not followed here. [2d] Sect. Crenatae (A. Berger) B. Ullrich 1993 ( Unterreihe Crenatae A. Berger 1915; incl. Group Crenatae Gentry 1982 (nom. illeg., ICN Art. 53.3, and nom. inval., ICN Art. 37.1)): Ros medium-sized to large, usually solitary, rarely surculose; stems short; L green to yellowish-green, young pruinose, usually with clearly visible marks left by the central bud, L margins deeply crenate and undulate, marginal teeth large, irregular in size and spacing, frequently with small interstitial teeth, on distinct prominences; Inf tall, narrow; Tep red or purple in bud, yellow at anthesis, lobes 2.5–5
as long as the tube; Fil inserted at, above or below mid-tube. — 7 species mainly from the Mexican Sierra Madre Occidental and the Trans-Mexican Volcanic Belt. [2e] Sect. Campaniflorae (Trelease) R. H. Webb & G. D. Starr 2015 ( Group Campaniflorae Trelease 1912b  Reihe Campaniflorae A. Berger 1915): Ros small to large, rather open, mostly solitary (except var. capensis); stems short; L rather softly succulent, 35–150 cm, green,

Agave AGAVACEAE

margins not horny, marginal teeth usually 4–7 mm, uniform in size and spacing; Fl campanulate; Tep tube thin-walled, deep, broad (15–22 mm), Tep red to purplish in bud, opening (orange-) yellow, lobes 1.5–2
as long as the tube; Fil deeply attached in the tube. — 1 species (A. aurea) from Baja California (Mexico). [2f] Sect. Umbelliflorae (Trelease) R. H. Webb & G. D. Starr 2015 ( Group Umbelliflorae Trelease 1912  Reihe Umbelliflorae A. Berger 1915): Stems commonly branching from L axils, frequently developing long branching stems resulting in fragmented clones; L broad and short (600 flowers, axis reddish; Fl thickly fleshy, (70–) 80–90 (–100) mm; Ped 10–20 mm; Tep red to purple in bud, at anthesis yellowish to yellowish-green within, tip reddish, tube deeply furrowed, thick-walled, 10–15


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10–20 mm, bulging at the insertion of the stamens, lobes triangular or oblong, erect, conduplicate-revolute, thickly fleshy, unequal, 30–40
4–6 (–9) mm, incurved at the tips; OTep slightly larger; ITep with very thick keel and thin margins, red with paler margins; Fil large, flattened, 70–80 (–90) mm, purplish or red-spotted, inserted in the middle or at the mouth of the tube, yellowish or with purple dots; Anth excentric, straight to sinuous, (25–) 30–43 mm, yellow(ish) to bronze; Ov cylindrical, tapering at the base, 3- to 6-angled, 30–50
(4–) 6–8 mm, neck thick, furrowed, 4–7 mm; Sty 80–110 mm, much longer than the stamens after anthesis; Sti clavate, 4–5 mm ∅; Fr ovoid, ellipsoid or clavate, 40–50 (–65)
20–30 mm, 5–12 mm stipitate, tapering from the apex; Se crescent-shaped, 7–9
4–6 mm, shiny, with inconspicuous wing. — Cytology: 2n = 180 (Simpson & al. 2011). A very large species with affinities to species of Sect. Guatemalenses (as Group Hiemiflorae) from Chiapas and Guatemala, which flower in winter, produce straight and strong peduncles, and with tightly arranged flowers in bracteolate globose purplish clusters in narrow panicles (Gentry 1982: 468–476). Karwinski’s type locality “Monte Tanga” (see Piña Luján (1994)) was re-named as Monte Fiscal-Santos and is located W of Ciudad Oaxaca (David 2009), and this is supported by Karwinski’s letters (Phillips 2013). Earlier authors (Baker, Berger, Trelease) confused the species with A. salmiana (Gentry 1982). Gentry’s records from NE Puebla and C Veracruz apparently refer to cultivated plants of var. mirabilis (see there). Ullrich (1990a) reports on a new locality from Guerrero. The records from México state (Ávila-Akerberg & al. 2008: 617) (specimens Castan˜eda Rojas 27, MEXU 1047763 & 1047765) need verification. A. macroculmis is a synonym to be placed here (Ullrich 1990i), and was misinterpreted by Gentry (1982: 598) for plants later described as A. gentryi and A. montana (see there). Leaves frequently show necrotic spots caused by a herbivorous fly whose larvae develop inside the leaves of A. atrovirens and A. salmiana (Brunel & Rull 2010).

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A. atrovirens var. mirabilis (Trelease) Gentry (Agaves Cont. North Amer., 473, ill. (p. 476), 1982). Type: Mexico, Veracruz (Trelease 7 p.p. [MO, DES]). — Lit: García-Mendoza (2011a: 20–21, with ills.). Distr: Mexico (NE & S Puebla, C Veracruz, N Oaxaca); cool montane habitats, in pine-oak forests and cultivated, 2150–2480 m. I: García-Mendoza (2002: 179); Richter (2011: 83); Spracklin (2014); Hochstätter (2015: VI: 6).  Agave mirabilis Trelease (1920)  Agave atrovirens ssp. mirabilis (Trelease) Hochstätter (2015). [2o] Differs from var. atrovirens: L broadly lanceolate, consistently light grey-glaucous; Fl smaller, 60–80 mm, without large bracteoles; Ov 25–30 mm. A light glaucous form cultivated for pulque and as fences about Las Vigas, Veracruz, without confirmed occurrence elsewhere according to Gentry (1982: 473). Photographs from the type locality were published on the web by Spracklin (2014). Gentry’s records for var. mirabilis from NE Puebla and C Veracruz apparently belong here according to GarcíaMendoza (2011a), who also reports the taxon from Oaxaca. A. attenuata Salm-Dyck (Hort. Dyck., 303, 1834). Type: [neo — icono]: Curtis’s Bot. Mag. 88: t. 5333, 1862. — Distr: Mexico (Sinaloa, SW Durango, Nayarit, Jalisco, Colima, Michoacán, México, Guerrero, Oaxaca). The neotype cited above was designated by Gentry (1982: 68). A. attenuata ssp. attenuata — Lit: Gentry (1982: 66–68); Ullrich (1990h); Cházaro Basáñez & al. (1998); Vázquez-García & al. (2007b: 47, t. F); Ullrich (2007); all with ills. Distr: Mexico (Jalisco, Colima, Michoacán, México); rocky outcrops in pine forests, 1900–2500 m; flowers November to December. I: Curtis’s Bot. Mag. 88: t. 5333, 1862, as A. glaucescens; Irish & Irish (2000: t. 5); Heller (2006: 21, 37, 41, 66, 67); Etter & Kristen (2012: 87); Pilbeam (2013: 38); Meer (2014: as A. cernua); Hochstätter (2015: VIII: 38); Moore (2016: 246). – Fig. 7.

J. Thiede

Fig. 7 Agave attenuata ssp. attenuata (Etter & Kristen 1569: Mexico; Michoacán, between Morelia and Ciudad Hidalgo, 2260 m). (Copyright: J. Etter & M. Kristen)

Incl. Ghiesbreghtia mollis Roezl (1861) (incorrect name, Art. 11.4); incl. Agave glaucescens Hooker (1862) (nom. illeg., ICN Art. 53.1); incl. Agave attenuata var. compacta Jacobi (1865); incl. Agave attenuata var. subundulata Jacobi (1866); incl. Agave attenuata var. coarctata Jacobi (1866) (nom. inval., ICN Art. 38.1a); incl. Agave ghiesbreghtii [?] mollis Hort. Belg. ex Jacobi (1866) (nom. inval., ICN Art. 36.1c); incl. Agave attenuata var. elliptica Jacobi (1868) (nom. inval., ICN Art. 38.1a); incl. Agave spectabilis Roezl ex Baker (1877) (nom. inval., ICN Art. 36.1c); incl. Agave attenuata var. glauca Carrière (1879) (nom. inval., ICN Art. 36.1c); incl. Agave attenuata var. latifolia Salm-Dyck ex A. Terracciano (1885); incl. Agave attenuata var. brevifolia Jacobi ex A. Terracciano (1885) (nom. inval., ICN Art. 38.1a); incl. Agave attenuata var. paucibracteata Trelease (1914); incl. Agave cernua A. Berger (1915); incl. Agave compacta Hort. Besaucèle ex

Agave AGAVACEAE

A. Berger (1915) (nom. inval., ICN Art. 36.1c/ 38.1a); incl. Agave elliptica Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/ 38.1a); incl. Agave spectabilis Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave virens Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/ 38.1a); incl. Agave attenuata ‘Nova’ Kimnach & Trager (1990) (nom. inval., Art. ICNCP 17.9); incl. Agave attenuata ‘Boutin Blue’ C. C. Walker (2000). [1b] Stems 1 to several, usually ascendingcurved, 0.5–1.5 m, becoming naked in age; L relatively short-lived, ovate-acuminate, softly succulent, broadest in the middle, flat to concave, 50–70
12–16 cm, light glaucous-grey to pale yellowish-green, margins smooth, straight, without teeth, leaf tip finely tapered, soon fraying, soft; Inf 2–3.5 m, ‘spicate’, densely flowered, part-Inf shortly pedunculate ‘fascicles’ with 3–8 flowers in the axils of chartaceous Bra; Fl 35–50 mm; Tep greenish-yellow, tube shallowly funnel-shaped, 3–5 mm, lobes linear-oblong, thin, recurving, equal, 16–24 mm, inner frequently broader, with broad low keel, mucronate with white-floccose tips; Fil slender, flattened, tapered, 35–45 mm, whitish, inserted on the rim of the tube; Anth centric, 15–20 mm, greenish-yellow; Ov fusiform, 15–25 mm, green, neck constricted; Fr oblong, 20–30
10 mm or smaller, shortly beaked, stipitate; Se crescent-shaped to deltoid, 3–3.5
2–2.5 mm, marginal wing low. — Cytology: 2n = 45–62 (Lv & al. 2009). The presence of a flower tube groups this species in Sect. Inermes (Group Amolae) and not in Sect. Choritepalae, where the otherwise similar A. bracteosa and A. ellemeetiana are placed. It is closely related to A. gilbertii (Gentry 1982: 70). For relationships, see also under ssp. dentata. Cházaro Basáñez & al. (1998) provide a new record for Colima. A. attenuata ‘Boutin Blue’ (Walker 2000: 97–98), first invalidly named A. attenuata ‘Nova’, differs in its leaves with an extremely blue colouration and its erect inflorescence; it was collected in 1970 as Kimnach & Boutin 3019 (ARIZ, DES, HNT) in the Sierra de Manantlán (Jalisco). A. attenuata ‘Emery Stripy’ (Walker 2000) and

63

‘Myron’s Magic’ (Smith 2004) are variegated cultivars (Alsemgeest 2006), and ‘Blue Flame’ is a hybrid with A. shawii (Trager 2005, Mays 2005: 44, Starr 2014: 219). The latter and further cultivars are illustrated by Guillot Ortiz & Meer (2009b). ‘Blue Glow’ is a hybrid with A. ocahui (Starr 2012: 56–58, with ills.), and ‘Snow Glow’ (Starr 2014: 224, with ill.) is a variegated sport selected from it. The taxon is reported as casually naturalized in Gran Canaria (Canary Islands) (Verloove 2013: 80) and in Portugal (Silva & al. 2015: 73), and introduced in Madeira and Libya (Euro+Med 2006+: accessed Jan. 2019). A. attenuata ssp. dentata (Roezl) B. Ullrich (Haseltonia 12: 27, ills. (pp. 23–27), 2007). Type [neo]: Mexico, Guerrero (Rzedowski & McVaugh 289 [ENCB, MICH]). — Lit: Gentry (1982: 79–82, with ills.); McVaugh (1989); Cházaro Basáñez & al. (1998); Vázquez-García & al. (2007b: 47, t. V-W); González-Elizondo & al. (2009: 58–60, with ill.); all as A. pedunculifera. Distr: Mexico (Sinaloa, SW Durango, Nayarit, Jalisco, Michoacán, Guerrero, Oaxaca); mountain slopes in tropical deciduous or pine-oak forests, 300–2200 m; flowers November to February. I: Richter (2011: 10, 61, 111, as A. pedunculifera); Etter & Kristen (2012: 88); Pilbeam (2013: 166–167, as A. pedunculifera); Hochstätter (2015: VIII: 39).  Ghiesbreghtia dentata Roezl (1861) (incorrect name, Art. 11.4); incl. Agave pruinosa Lemaire ex Jacobi (1865); incl. Agave debaryana Jacobi (1868); incl. Agave kellockii Jacobi (1868); incl. Agave ghiesbreghtii [?] dentata Hort. Belg. ex Jacobi (1868) (nom. inval., ICN Art. 36.1c); incl. Agave attenuata [?] dentata J. Verschaffelt (1876); incl. Agave dentata hort. ex Baker (1877) (nom. illeg., ICN Art. 53.1); incl. Agave attenuata var. subdentata Cels ex Carrière (1879); incl. Agave attenuata var. serrulata A. Terracciano (1885)  Agave cernua var. serrulata (A. Terracciano) A. Berger (1915); incl. Agave pedunculifera Trelease (1920). [1b] Differs from ssp. attenuata: Stems short; L mostly ovate-acuminate and 50–70
15–18 cm, or lanceolate and 80–90
11–15 cm,

64

pale green to glaucous-white, margins narrowly lined with brown or white, with closely set denticles 0.5–2 mm, otherwise smooth; terminal Sp acicular, weak, 10 mm; Inf erect or recurving (on cliffs), part-Inf with 2 or 4 flowers; Ped 20–30 mm; Tep lobes 12–22
5–6 mm; Fil 30–35 mm; Fr 15–20 mm; Se 2.5
2 mm. Known since Trelease (1920) as A. pedunculifera. Gentry (1982) considered it closely related to A. attenuata based on leaf and flower characters, but distinguished by being nearly stemless throughout all observed populations. The subspecies exhibits considerable variability in leaf form and size and depth of the flower tube, but the different forms are linked by intermediates (Gentry 1982: 80). GonzálezElizondo & al. (2009) provide a new record from Durango. The recently described blueleaved A. manantlanicola from Jalisco is closely related (see there). There has been some argument which name is the correct basionym for this taxon. Since Ghiesbreghtia dentata is merely incorrect, rather than invalid, its use as basionym is not precluded. A. aurea Brandegee (Proc. Calif. Acad. Sci., ser. 2, 2: 207, 1889). Type [lecto]: Mexico, Baja California Sur (Brandegee s.n. [UC, CAS, MO]). — Lit: Trelease (1912b: 44, 49–51, tt. 32–37); Berger (1915: 168–170); Gentry (1978: 70–83, tt. 3–4); Gentry (1982: 309–322, with ills.); Webb & Starr (2015: 68–71, 97–98, with ills.). Distr: Mexico (C & S Baja California Sur). Gentry (1978) and Gentry (1982) recognized 3 species (A. aurea, A. capensis and A. promontorii) within Sect. Campaniflorae (as Group), which, at the latitude of Todos Santos, become similar to each other, suggesting interfertility (Gentry 1982: 313). Based on detailed field studies, Webb & Starr (2015) recognized a single species only and reduced A. promontorii to subspecific and A. capensis to varietal status. Brandegee’s protologue does not indicate a type, but according to Trelease (1912a: 50), the type is “Brandegee, Feb. 13, 1889”. The designation of “Brandegee s.n.” at UC by Gentry (1972: 78) represents a lectotypification.

J. Thiede

A. aurea ssp. aurea — Lit: Trelease (1912b: 49–50, tt. 32–34); Berger (1915: 168–169); Gentry (1978: 71, 74, 77–81, t. 4); Gentry (1982: 311–315, with ills.); Turner & al. (1995: 43–44); all as A. aurea; Webb & Starr (2015: 68–70, 97–98, with ills.). Distr: Mexico (Baja California Sur: Sierra de la Giganta and Cape district); lava fields and granitic or volcanic slopes, mostly 300–1070 m; flowers September to April. I: Heller (2006: 20); Richter (2011: 74); Pilbeam (2013: 39); Hochstätter (2015: V: 4); all as A. aurea. Incl. Agave campaniflora Trelease (1912). [2e] Stems short; Ros rather open, 1–1.2
1–2 m, solitary; L linear to long-lanceolate, widely arching, pliable, guttered, rounded below, thickly fleshy towards the base, 60–110
7–12 cm, green to yellow-green, somewhat glaucous, margins straight to wavy; marginal teeth moderate, mostly 4–7 mm from low angular bases, regularly spaced, 10–20 mm apart, dark to light brown; terminal Sp subulate, 25–35 mm, dark brown or greyish-red, shortly decurrent or decurrent as dark horny margin through the uppermost 8–10 tooth bases; Inf 2.5–5 m, paniculate, peduncle with remote lanceolate Bra, reddish, fertile part broad, part-Inf broad, congested, 15–25, in the upper of the inflorescence, red to purplish in bud and fruit; Fl campanulate, 43–70 mm; Tep red to purplish in bud, opening yellow to orangeyellow, tube evenly grooved from the lobe sinuses, 8–14
14–18 mm, lobes lanceolate, acute, incurved, 16–19 mm, outer larger, with tufts of white papillae below the distinctly hooded tip, inner ones with a distinct long keel and yellow, the thin margin involute; Fil stout, 35–45 mm, those in front of the outer tepals inserted 5–10 mm above the tube base, slightly higher than those in front of the inner tepals; Anth 19–23 mm; Ov slender, angular-cylindrical, 25–35 mm, reddish, neck constricted, furrowed, 6–10 mm; Sty somewhat longer than the tepals; Sti capitate; Fr oblong, 35–45
15–17 mm, reddish, drying light brown, rounded-apiculate, not stipitate; Se irregularly crescent-shaped, 7–8
5–5.5 mm, dull. The most widespread and common taxon in Sect. Campaniflorae (as Group), easily

Agave AGAVACEAE

recognized by its long, narrow, green leaves arching out to form an open, spreading rosette, its broad, rather diffuse, reddish inflorescences, and its bright yellow flowers from reddish buds and ovaries (Gentry 1982: 313, Webb & Starr 2015). N of Todos Santos, it intergrades with both ssp. promontorii and var. capensis in the colour and shape of the leaves (Gentry 1982: 313 (as A. promontorii and A. capensis), Webb & Starr 2015: 69), and lower elevation populations of ssp. aurea in the Sierra de la Giganta are very similar to ssp. promontorii (Webb & Starr 2015: 69). A. aurea ssp. promontorii (Trelease) R. H. Webb & G. D. Starr (Haseltonia 20: 70, ill. (p. 71), 2015). Type: Mexico, Baja California (Nelson & Goldman 7437 [US]). — Lit: Trelease (1912b: 50–51, tt. 35–37); Berger (1915: 169–170); Gentry (1978: 71, 74, 80–83, t. 3); Gentry (1982: 319–321, with ill.); all as A. promontorii; Webb & Starr (2015: 70–71, 98, with ills.). Distr: Mexico (S Baja California Sur: Sierra La Laguna, Cape District); slopes and summits of granitic mountains, 720–1800 m; flowers January and April to May. I: Goldman (1916: t. 111B); Richter (2011: 74); Pilbeam (2013: 178); Hochstätter (2015: V: 6); all as A. promontorii.  Agave promontorii Trelease (1912a). [2e] Differs from ssp. aurea: Ros larger, 1–2
2–2.5 m; L larger, 100–150
11–17 cm, margins straight; marginal teeth curved, 5–10 mm apart, reddish-brown; terminal Sp longer, 30–50 mm, dark brown; Inf larger, 5–9 m, peduncle with deltoid Bra, fertile part with 25–30 part-Inf; Fl 60–75 mm, larger in nearly all parts. This subspecies is larger and occurs at higher elevations than ssp. aurea. Since the 2 taxa intergrade, subspecies status (Webb & Starr 2015) is more approprate than species status (Gentry 1978, Gentry 1982). A. aurea var. capensis (Gentry) R. H. Webb & G. D. Starr (Haseltonia 20: 70, ill., 2015). Type: Mexico, Baja California Sur (Gentry & Fox 11247 [US, ARIZ, MEXU]). — Lit: Gentry (1978: 72–78); Gentry (1982: 310, 316–318); both with

65

ills. and as A. capensis; Webb & Starr (2015: 70–71, 98, with ills.). Distr: Mexico (Baja California Sur: Cape District); arid slopes,  sea-level to >300 m. I: Heller (2006: 71); Richter (2011: 74); Pilbeam (2013: 49); Hochstätter (2015: V: 5); all as A. capensis.  Agave capensis Gentry (1978). [2e] Differs from ssp. aurea: Ros smaller, 0.6–0.8
0.8–1.2 m, caespitose by axillary budding, eventually in large clusters; L smaller, mostly 35–60
4–7 cm, grey-green with a light glaucous covering, margins wavy; marginal teeth reddishbrown to greyish, on short mamillate prominences; terminal Sp short, 15–30 mm, dark brown; Inf mostly 2.5–3.5 m, fertile part relatively narrow; Fl 50–65 mm; Fil with an apical gland at the point of anther attachment. — Cytology: 2n = 60 (Pinkava & Baker 1985: as A. capensis). Differs from the other 2 taxa primarily in its prolifically offsetting growth leading to large clusters of rosettes (Webb & Starr 2015: 70). The varietal status is warranted because of its distribution, which overlaps with that of ssp. aurea. A. avellanidens Trelease (Annual Rep. Missouri Bot. Gard. 22: 60, tt. 61–62, 1912). Type: Mexico, Baja California (Brandegee 6 [UC]). — Lit: Berger (1915: 267); Gentry (1978: 61–64, with ills.); Gentry (1982: 361–363, with ills.); Heim (2014: 8–11, with ills.); Webb & Starr (2015: 71–73, 98, with ills.). Distr: Mexico (S Baja California: Sierra La Libertad); desert scrub, to 325 m; flowers April to May. I: Irish & Irish (2000: t. 6); Richter (2011: 104); Pilbeam (2013: 40); Hochstätter (2015: V: 24). – Fig. 8. [2j] Stems to 0.5 m; Ros 0.6–1.2
1–1.5 m, solitary; L many, broadly linear-lanceolate to ovate, thickly fleshy, rigid, little or not narrowed at the base, shortly acuminate, smooth, 40–70
9–14 cm, yellow-green, green or blue-green, margins straight, frequently horny; marginal teeth straight or variously curved, variable in size and curvature, flattened, 5–15 mm, grey to brown, mostly 10–30 mm apart, rather regularly spaced, on small to large, broadened bases; terminal Sp conical, strong, broadly grooved above, 25–45 mm, brown to greyish, strongly decurrent as horny margin; Inf 4–6 m,

66

J. Thiede

Fig. 8 Agave avellanidens (cult.: USA; Arizona, Desert Botanical Garden Phoenix). (Copyright: J. Thiede)

paniculate, peduncle with small Bra, fertile part in the upper of the inflorescence, conical, with 25–35 dense large globose part-Inf; Fl slender, 40–70 mm; Tep pale greenish-yellow, drying orange-yellow, tube flaring, ridged within and grooved on the outside from overlapping lobe sinuses, 4–6
13–15 mm, lobes  equal, 16–24 mm; Ov fusiform, tapering to the base, 20–40 mm, pale yellow, neck sometimes constricted; Sty somewhat shorter than the filaments; Sti capitate; Fr broadly oblong, 35
20 mm, not stipitate, scarcely beaked, dark brown; Se unknown. — Cytology: 2n = 60 (Bennett & Leitch 1995, Simpson & al. 2011). According to Gentry (1982: 354, 394), A. avellanidens, A. gigantensis and A. moranii form a separate group within Sect. Deserticolae (as Group), differing from other members in non-surculose rosettes with broader and greener leaves. Webb & Starr (2015) formalized this group as Sect. Conicae, named after its characteristic conical inflorescences, and added the greyish-leaved A. turneri to it. Within Sect. Conicae, A. avellanidens is differentiated by its short green broadly linear-lanceolate leaves not or scarcely narrowed towards the base (Webb & Starr 2015). — A. avellanidens resembles A. shawii ssp. goldmaniana from Sect. Umbelliflorae (as Group) in habit, and they appear to hybridize near Rosarito where their ranges overlap (Gentry 1982, Webb & Starr 2015).

A. azurea R. H. Webb & G. D. Starr (Haseltonia 19: 90–92, ills., 2014). Type: Mexico, Baja California Sur (Webb & Starr 20130418-1 [HCIB]). — Lit: Webb & Starr (2015: 88–89, with ills.). Distr: Mexico (Baja California Sur: Picachos de Santa Clara); rocky colluvial hillslopes and alluvial fans, 295–360 m, in desert scrub; flowers in April, but appearing opportunistically. [2k] Acaulescent, Ros 0.8–0.9
1–1.5 m, solitary; L 13–32, obovate to oblanceolate, flat to slightly guttered, attenuate at the tip, slightly rough and waxy above, smooth to slightly rough below, 55–76
14–22 cm, widest just above the middle, 10–17 cm broad at the narrowest point below the middle, glaucous blue-green, faintly to moderately banded, with light imprints from the central bud, margins straight to rarely repand; marginal teeth deflexed, 10–15 mm, orangeyellow, reddish-purple or brownish-red, aging to whitish-grey, 40–50 mm apart, sessile or on small prominences; terminal Sp 40–70 mm, decurrent to the first 1–2 pair(s) of teeth; Inf 2–4.5 m, paniculate, peduncle 0.7–3 m; primary Bra triangular, 14–17
2–3 (at the base) cm, fertile part 1.3–1.5 m, Bra 7
4 cm, without terminal Sp, with 6–9 partInf with 6–8 stalked ‘umbels’ with 8–14 flowers each; Ped 6–30 mm; Fl 79–85 mm; Tep yelloworange, tube 9–22
15–22 mm, lobes slightly unequal, outer 22–30
9–13 mm, inner 21–29
10–13 mm; Fil 43–65 mm, inserted on the rim of the tube or 14–15 mm above the base of the tube; Anth 16–28 mm; Ov 30–40 mm, neck 6–10 mm;

Agave AGAVACEAE

Sty 85–90 mm when fully exserted; Fr oblong, 35–60
16–19 mm, with transverse lineations on the locules, beaked for 1–5 mm; Se crescentshaped to ovate, 5–7
4–5 mm, shiny black. Plants from the Picachos de Santa Clara were included in, but recognized as different from, A. vizcainoensis by Gentry (1982: 407) and were also identified by him as A. sebastiana or A. gigantensis according to the protologue, but have now been named A. azurea. A. azurea differs from A. vizcainoensis in its mostly solitary habit, larger size of most characters, leaves with significantly different shape and margins, and smaller fruits. It also resembles A. turneri of Sect. Conicae in habit, but the latter has a different inflorescence. Webb & Starr (2015) placed A. azurea and A. vizcainoensis — both characterized by long tepal tubes — in a separate Sect. Intermediae, which is considered intermediate between Sect. Deserticolae and Sect. Umbelliflorae (see under A. vizcainoensis). A. bahamana Trelease (Mem. Nation. Acad. Sci. 11: 40, tt. 84–86, 1913). Type: Bahamas (Britton & Millspaugh 2340 [MO, NY, F]). — Lit: Berger (1915: 203); Britton & Millspaugh (1920: 74–75); Gillis (1976); Correll & Correll (1982: 310). Distr: Bahamas (Andros, Berry Islands, Cat, Cave Cay, Eleuthera, Exuma, Grand Bahamas, San Salvador [Watling] Island); open rocky plains and ridges, open coppices and pinelands, and sandy beach dunes, in dry broad-leaf evergreen wood- or scrubland; flowers mostly January to May. I: Hochstätter (2015: VII: 79, 82). Incl. Agave sobolifera Hitchcock (1893) (nom. illeg., ICN Art. 53.1); incl. Agave rigida Northrop (1902) (nom. illeg., ICN Art. 53.1). [2r] Acaulescent; Ros solitary; L rather narrowly lanceolate, fleshy, curved, concave, occasionally somewhat conduplicate, 200–300
15–20 cm, dull greyish to greyish-green, margins nearly straight; marginal teeth straight or the longer teeth appressed-recurved, triangular, scarcely lenticular at the base, 3–5 mm, usually 5–10 mm apart, reduced near base and tip of the leaf or rarely greatly reduced throughout or nearly lacking, sometimes on small green tubercles;

67

terminal Sp slightly recurved, stoutly conical, usually becoming involutely grooved below the middle, smooth, 10–15
4–5 mm, brownish becoming grey, dull, decurrent; Inf to 10 m, paniculate, peduncle with deltoid non-overlapping Bra, fertile part ovoid, part-Inf on slightly ascending Br,  in the upper of the Inf; Ped 10 mm; Fl 50–60 mm; Tep 15
4 mm, golden-yellow, tube conical, 7 mm, lobes 15
4 mm; Fil 30–35 mm, inserted almost in the tube throat; Ov oblong-fusiform, 30–35 mm; Sti  as long as the filaments; Fr oblong, 40–50
20–25 mm, shortly stipitate and beaked; Se 8
6 mm. The most widespread Agave species on the Bahamas. Its area covers all larger islands of the N Bahamas. — Britton & Millspaugh (1920) suggested that A. acklinicola, A. cacozela, A. indagatorum and A. millspaughii “may be but races of A. bahamana”. Gillis (1976) included these four species as well as A. braceana (i.e., all other members of Sect. Bahamanae) in the synonymy of A. bahamana, but this concept was not followed by subsequent authors. The recent accounts by Álvarez de Zayas (1995), AcevedoRodríguez & Strong (2012), and Freid & al. (2014) recognize all six species as distinct, pending a detailed study of the group. A.
bedinghausii Rebut (Suppl. Cat. Cact., 3, 1893). Type: not typified. — Lit: Koch (1865: 112); Ladenburg (1893: 74). Distr: Cultivated only. [1g
3a] Acccording to the protologue (not seen) reported as the artificial hybrid between A. ghiesbreghtii (of Subgen. Littaea) and A. maculata (of Subgen. Manfreda) (=
Mangave). A.
bedinghausii was first mentioned by Koch (1865: 112) as mere name in the catalogue of “van Sweet in Gent”, most probably an error for L. de Smet, Gent, Belgium. The plant was exhibited by the private nursery of M. Bedinghaus, Gent (Ladenburg 1893: 74). Trelease (1920: 123) listed an A. bedinghausii as a mere form in the synonymy of the strong-toothed A. verschaffeltii (= A. potatorum), which thus must have been a different plant. A.
beguinii hort. ex A. Berger (Agaven, 61, 1915). Type: not typified. — Lit: Ullrich

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(1993b: 27, 31–32, as A. beguinii). Distr: Cultivated only. [1] This is a garden hybrid of unknown parentage that was probably raised in the “Jardin Botanique au Parc de la Tête d’Or” at Lyon (France) and named and distributed by P. Rébut at Chazey-d’Azerognes (France). It was given to La Mórtola via M. le Chanoine Béguin (Brignoles, France) in 1899 and flowered there in 1905 (Berger 1912: 357, Berger 1915: 61). Apparently lost from cultivation. A. bicolor (Solano & García-Mendoza) Thiede & Eggli (Kakt. and. Sukk. 52(6): 166, 2001). Type: Mexico, Oaxaca (García-Mendoza & al. 2403 [MEXU, BRIT, FEZA, SMU]). — Lit: Solano Camacho & García-Mendoza (1998); García-Mendoza (2011a: 83–86); both as Polianthes and with ills. Distr: Mexico (NW Oaxaca); grasslands and in pine and pine-oak forests, 2300–2500 m, flowers July to August. I: Castro-Castro & al. (2015: 145); Hochstätter (2016: II: 6); Castro-Castro & al. (2016: 721); all as Polianthes.  Polianthes bicolor Solano & García-Mendoza (1998). [3b2] Herbaceous; corm cylindrical, 4–6
2–2.5 cm, bulb ovoid, (2–) 3–5
(1–) 1.5–2.5 cm, covered with dry leaf bases; R contractile, thickened; basal L (3–) 4–6 (–12), lanceolate, semi-succulent, generally undulate, smooth to papillose, apex acute, (5–) 8–15
(0.6–) 1–4 cm, shiny green, margins papillose, hyaline, usually repand; Inf erect, 24–40 (–54) cm, peduncle with lanceolate Bra, lower Bra 3–5 (–7.5)
0.2–0.5 cm, upper Bra progressively smaller, fertile part with 3–5 (–9) flowering nodes; Ped 6–13 mm, reddish; Fl geminate, (20–) 23–29 mm, fragrant; Tep yellowish-green, tube funnel-shaped, abruptly widened at the base, basal part erect, abruptly curved outwards just above the ovary, horizontal or down-curved at anthesis, (1.6–) 2.4–3 (–5.5) mm ∅ at the mouth; Tep lobes orbicular to transversally elliptic, apiculate, subequal, outer 2–3 (–4)
(1.7–) 2–3 (–4) mm, inner with cucullate apex and pilose tips; Fil filiform, inserted somewhat above mid-tube 11–18 (–19) mm above the ovary, free part 4 (–15) mm,

J. Thiede

included, green; Anth oblong, 4–5 (–9)
1–1.5 (–2.5) mm, yellowish-green; Ov cylindrical, 10
3–4 mm; Sty (10–) 19–26 (–32) mm; Sti 3-lobate; Fr globose, 11–13
10–11 mm, with persistent perianth remains; Se semicircular, one face straight, 4.5
2.4 mm, black. According to the protologue characterized by its semisucculent undulate leaves with repand margins, flowers with the tube abruptly widened above its base, green tepal lobes, and included stamens. The species belongs to Ser. Bravoa of Sect. Polianthes and is morphologically closest to A. coetocapnia and A. graminifolia. A. bicolor, A. alboaustralis and A. oaxacana represent the S-most species of Sect. Polianthes and its sole representatives in Oaxaca (key in Solano & Ríos-Gómez (2014)). A.
blissii (Worsley) Thiede & Eggli (Kakt. and. Sukk. 50(5): 111, 1999). — Lit: Worsley (1911: with ill.); Howard (1977); both as Polianthes. Distr: Cultivated only. I: Howard (2001: 9).  Polianthes
blissii Worsley (1911). [3b1
3b2] This is the garden hybrid A. coetocapnia (as Bravoa geminiflora)
A. amica (as Polianthes tuberosa) first raised in 1911, but still extant in cultivation (Howard 2001). A. boldinghiana Trelease (Mem. Nation. Acad. Sci. 11: 21, tt. 11–13, 1913). Type [syn]: Curaçao (Boldingh A2 [MO [6 sheets]]). — Lit: Berger (1915: 225–226); Hummelinck (1936: 239–240, 248–249, tt. VI, VIIIc, figs. 19–20); Hummelinck (1938: 20–21); Hummelinck (1993: 107–109, 219, with ills.). Distr: Leeward Antilles (Aruba, Bonaire, Curaçao), NE Colombia (?); coral rocks and coral debris, 2–50 m; flowers in May. I: Heller (2006: 68); Pilbeam (2013: 42–43); Hochstätter (2015: VIII: 87). [2u] Stems almost none; Ros 2 m ∅, suckering; L rather many to rather few, narrowly oblanceolate, subacuminate, rather slightly S-curved, apex slightly curved upwards, openly concave, 90–125
15 cm, green, passing into somewhat glaucous, margins rather straight; marginal teeth often irregularly upcurved above and recurved below, heavily triangular or from crescent-shaped

Agave AGAVACEAE

bases, mostly 4–5 (–7) mm, scarlet becoming chestnut-brown, mostly 10–15 mm apart, on rather well to strongly developed hardening or partly green prominences; terminal Sp acicular, (very) slightly upcurved at the tip, somewhat flexuous, grooved and usually involute towards the base, smooth, polished towards the tip, 15–30
2–4 mm, red-brown, shortly or not decurrent, dorsally rather slightly intruding into the leaf tissue; Inf 5 m, paniculate, peduncle with rather distant, narrowly triangular, appressed Bra, fertile part narrowly oblong, part-Inf few, distant, on ascending Br in the upper or less of the Inf, freely bulbilliferous; Ped 4–7 mm; Fl 45–46 mm; Tep golden-yellow, tube conical, 5–7 mm, lobes 15–20
4 mm; Fil 34–35 mm, inserted 0–1 mm below the tube throat; Anth 18–21 mm; Ov broadly fusiform or oblong, 20–25 mm; Sty length unknown; Fr 33–36 mm, beaked, stipitate; Se unknown. Always distinguishable from all other species in its geographical range, but less well defined compared with certain forms of A. cocui and A. vicina (as A. vivipara) from other regions (Hummelinck 1938). The species appears not to belong to the indigenous flora (Hummelinck 1993) and was designated as “cultivated” already by Boldingh (1914: 17). Records from coastal NE Colombia (Bernal & al. 2015: 794) need verification against A. cocui. Trelease’s “type” at MO consists of 6 specimens that represent syntypes (Smith & Figueiredo 2014e: 229). Fig. 9 Agave bovicornuta (Etter & Kristen 561: Mexico; Sonora, near San Bernardo, Río Mayo, 1700 m). (Copyright: J. Etter & M. Kristen)

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A. bovicornuta Gentry (Publ. Carnegie Inst. Washington 527: 92, 1942). Type [syn]: Mexico, Sonora (Gentry 3672 [CAS [2 sheets], ARIZ [2 sheets]]). — Lit: Gentry (1982: 328–332); González-Elizondo & al. (2009: 61–64); Starr (2012: 22, 59–63); all with ills. Distr: Mexico (S Sonora, SW Chihuahua, S Durango, N Sinaloa); rocky open slopes in oak woodlands and pine-oak forests, 930–1850 m; flowers December to May. I: Irish & Irish (2000: t. 7); Heller (2006: 69); Richter (2011: 73, 108); Pilbeam (2013: 44–45); Greulich (2014a: 56); Hochstätter (2015: IV: 19); Moore (2016: 270). – Figs. 9 and 10.  Agave maximiliana var. bovicornuta (Gentry) González-Elizondo & al. (2009) (nom. inval., ICN Art. 36.1b). [2d] Acaulescent, Ros 0.6–1
1.5–2 m but often smaller, solitary; L lanceolate to spatulate, much narrowed towards the base, widest at or above the middle, smooth, 45–80
10–17 cm, yellowish-green to light or deep green, younger L frequently shining glaucous, with conspicuous imprints from the central bud, margins crenate; marginal teeth dimorphic, larger teeth mostly 8–12 mm, flexuous and slender above a broad base, mostly 20–40 mm apart, on prominent prominences, smaller teeth mostly 2–5 mm, 1 to several between the larger teeth, all chestnutbrown or dark brown to greyish-brown with age; terminal Sp strong, 30–50 mm; Inf 2.5–7 m, paniculate, peduncle short, with triangular-

70

J. Thiede

Fig. 10 Agave bovicornuta (Cultivated material without known wild origin). (Copyright: U. Eggli)

acuminate reflexed Bra 20–30 cm long, fertile part narrow, part-Inf short, compact, 20–30 in the upper of the inflorescence; Ped 5–10 mm; Fl 55–65 mm; Tep greenish-yellow, tube 6–9
12–14 mm, lobes linear-lanceolate, ascendingspreading, conduplicate, involute, sinuses broadly overlapping, 15–21
4–7 mm, ITep broader and 2-costate within, yellow; Fil 30–45 mm, yellow, inserted (3–) 4–5 mm above the tube base; Anth 18–24 mm, yellow; Ov 26–29 mm, pale green, neck constricted, 4–6 mm; Sty hardly longer than the tepals; Sti capitate, 3-lobate; Fr oblong, 40–50
15–20 mm, stipitate, valves thin; Se finely punctate, 15
7 mm, curved side with a flange or wing, hilar notch shallow. Gentry’s protologue does not indicate a holotype so that all specimens represent syntypes. The designation of “Holotype DS” (Gentry 1982: 329) is a lectotypification which is ineffective under ICN Art. 8.1, since the 2 sheets at DS (now at CAS) are not cross-labelled (ICN Art. 8.3). Distinguished within Sect. Crenatae by light to yellowish-green leaves with narrow bases, relatively small flowers, and the low insertion of the filaments in the middle of the tube. A. bovicornuta is common and widespread in the N Sierra Madre Occidental, but with usually local populations with a limited number of individuals. Its nearest relative appear to be the populations of A. maximiliana from Durango and Nayarit, but these have more glaucous, palecoloured rosettes with more pronouncedly heteromorphic teeth and a mostly shorter terminal spine (Gentry 1982). González-Elizondo & al. (2009)

provide a new record from Durango and provisionally suggest to treat the species as variety of A. maximiliana. Etter & Kristen (2003a: 9) mention and depict a putative natural hybrid with A. angustifolia (as A. vivipara). The flowers, washed to remove the bitterness, are eaten cooked or in the form of tortillas, and the “hearts” are baked for preparing a strong alcoholic beverage “suguí” (Bye & al. 1975: 94). — ‘Reggae Time’ is a cultivar with bluish-green leaves, and ‘Holstein’ and ‘Heifer’s Cream’ are variegated cultivars (Starr 2012). A. braceana Trelease (Mem. Nation. Acad. Sci. 11: 40, t. 83, 1913). Type: Bahamas (Brace 1982 [MO, F, NY]). — Lit: Berger (1915: 203); Britton & Millspaugh (1920: 74); Correll & Correll (1982: 310–311, with ills.); Freid & al. (2014: 217). Distr: Bahamas (Abaco, Andros, Great Bahama); rocky or sandy soils in open pinelands or coastal coppices; flowers mostly October to February. I: Hochstätter (2015: VII: 83). Incl. Agave mexicana Dolley & al. (1889) (nom. illeg., ICN Art. 53.1). [2r] Acaulescent; Ros solitary; L broadly oblanceolate, fleshy, curved, nearly flat, 65–70
19–20 cm,  3
as long as broad, grey, margins between the teeth straight or concave when the teeth are raised on low green prominences; marginal teeth straight or the lower teeth gently recurved, triangular, 2–3 mm, scarcely lenticular at the base, usually 5–10 mm apart; terminal Sp conical, straight or gently curved, flat or roundly grooved to about the middle or becoming

Agave AGAVACEAE

involute, smooth, 10–15
3 mm, brownish becoming grey, dull, slightly decurrent; Inf 4–7 m, paniculate, not known to be bulbilliferous; Ped 10 mm; Fl 40–45 mm; Tep golden-yellow, tube conical, 7 mm, lobes 15–17
3–4 mm; Fil 35 mm, inserted almost at the tube throat; Ov oblong-fusiform, 20 mm; Sty longer than the filaments; Sti capitate; Fr broadly oblong, 20–35
20 mm, beaked, shortly stipitate; Se 8
6 mm. The flowers are visited at day by birds (hummingbirds, warblers (Dendroica spp.) and others). Wintering warblers show temporal territorial aggression against conspecifics and hummingbirds (Emlen 1973). Documented from the islands Abaco and Great Bahama in the protologue; Freid & al. (2014: 217) add Andros. — See also the note under A. bahamana. A. bracteosa S. Watson ex Engelmann (Gard. Chron., ser. nov. 18: 776, ills., 1882). Type: [lecto — icono]: l.c. figs. 138–139. — Lit: Gentry

Fig. 11 Agave bracteosa (Etter & Kristen 696: Mexico; Coahuila, between Saltillo and Monterrey, 1370 m). (Copyright: J. Etter & M. Kristen)

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(1982: 90–93); Ullrich (1990b); García-Mendoza (2003); Starr (2012: 64–68); all with ills. Distr: Mexico (SE Coahuila, W-C Nuevo León); scattered on limestone cliffs and rocky slopes of the N Sierra Madre Oriental, 900–1750 m; flowers May to July. I: Curtis’s Bot. Mag. 140: t. 8581, 1914; Irish & Irish (2000: t. 8); Heller (2006: 70, 154); Richter (2011: 62, 116); Pilbeam (2013: 46–47); Hochstätter (2015: VII: 35). – Fig. 11. [1c] Ros open, small to medium-sized, forming caespitose mounds by above-ground axillary budding; L relatively few, 30, longlanceolate, arching and recurving, with weak fibres, widest near the base, convex in the basal , flat above, asperous, (40–) 50–70
3–5 cm, yellowgreen, margins minutely serrulate, without teeth; terminal Sp absent, leaf tip drying early, friable, wind-scuffed, yellowish; Inf ascending to erect, 1.2–1.7 m, ‘spicate’, peduncle with erect triangular-acuminate Bra, fertile part densely flowered in the upper of the inflorescence; Fl geminate, 22–26 mm; Tep white to pale yellow, tube virtually none, reduced to a short receptacle, lobes ovate, spreading, hyaline, distinct, 11 mm, the outer overlapping the inner, lanceolate, acute, the inner broadly ovate, obtuse, all floccose at the tips; Fil long-exserted, 50–60 mm, white, inserted on the receptacle, persisting, elongating after anthesis; Anth sagittate, 7–8 mm, yellow; Ov fusiform, 12–14 mm, virtually without neck; Sty eventually exceeding the stamens; Fr ovoidoblong, obtuse, 18–20
8–10 mm, stipitate; Se deltoid to semicircular, 3–3.5
2–2.3 mm. Very distinctive on account of its unarmed curling leaves and white flowers, so that is has never been confused with other species (Gentry 1982). Its distinctiveness prompted Ullrich (1990b) to place it within a reconsidered monotypic Group Serrulatae Baker. Gentry (1982) reports the inflorescences to emerge laterally from upper leaf axils so one and the same rosette may flower repeatedly, but this was dismissed as erroneous by Zona (2018). — The neotypification by Gentry (1982: 91) was superseded with a lectotype by Ullrich (1990b) as given above. ‘Monterrey Frost’ (Starr 2012: 68, with ill.), ‘Stingray’ (Starr 2014: 221, with ill.), and ‘Medio-Picta Alba’ (Greulich

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2017c: 161, with ill.) are variegated cultivars. A plant cultivated outdoors on a protected site in Switzerland withstood temperatures of 12  C to 18  C (Bolliger 2014). A. brevipetala Trelease (Repert. Spec. Nov. Regni Veg. 23: 362, 1927). Type: Hispaniola, Haiti (Ekman 1604 [US [2 sheets], B, G, GH, ILL, S]). — Distr: Hispaniola (Haiti: Massif de La Selle, top of Morne Cabaio), 2350 m; flowers in August. [2p] Acaulescent; Ros solitary; L broadly lanceolate,  100 cm, green, rather dull; marginal teeth broadly triangular, variously curved, with lenticular bases, 5–10 mm but apical and basal teeth smaller, glossy chestnut-brown, 10–15 mm apart, teeth at mid-leaf clasping the green marginal prominences below them, intervening margin straight; terminal Sp smooth, curved, subterete, involutely narrowly grooved, 20–25
6 mm, rather glossy chestnut-brown, decurrent for some 10 cm and connecting to the small upper teeth; Inf paniculate, part-Inf with densely clustered flowers at their tips, not known to be bulbilliferous; Ped 5 mm; Fl 35 mm; Tep colour unknown, drying dark, tube 5 mm, lobes 10 mm; Fil 25 mm, inserted towards the tube throat; Ov thick, oblong, 20 mm; Fr and Se unknown. Described on the base of the dried type material only. According to Álvarez de Zayas (1995) possibly conspecific with A. antillarum (see there). See also the note for A. brevispina. A. brevispina Trelease (Repert. Spec. Nov. Regni Veg. 23: 363, 1927). Type: Hispaniola, Haiti (Ekman 5371 [US [2 sheets], B, GH, ILL, S]). — Distr: Hispaniola (Haiti: Croix-des-Bouquets, Plaine Cul de Sac); limestone, 150 m; flowers in December. [2p] Acaulescent; Ros solitary; L broadly lanceolate,  100
10 cm, dark green, rather dull; marginal teeth straight or some of the lower teeth recurved, rather narrowly triangular, with lenticular base, 1–3 mm, 5–15 mm apart, intervening margin almost straight; terminal Sp straight, somewhat flattened, involutely narrowly grooved, slightly granular, 10
3 mm, rather dull hazel-

J. Thiede

brown, tip darker, decurrent for 2
its length; Inf 4 m, ‘paniculate’, part-Inf rather slender, recurving, shortly few-flowered at the tips and flowers densely clustered, not known to be bulbilliferous; Ped slender, 10–15 mm; Fl 40 mm; Tep yellow, tube 5 mm, lobes 12
5 mm; Fil 25 mm, inserted almost at the tube throat; Ov oblong-fusiform, 25 mm; Fr and Se unknown. — Cytology: 2n = 60 (Granick 1944: identity doubtful). Described on the base of the dried type material only. The sample studied cytologically by Granick (1944) is of unknown origin. Trelease left this species and A. brevipetala unplaced, which was followed in the first edition of this handbook. According to Álvarez de Zayas (1995), both are possibly conspecific with A. antillarum (see there) and belong to Sect. Antillanae. A. brittoniana Trelease (Mem. Nation. Acad. Sci. 11: 44–45, tt. 98–99, 1913). Type: Cuba, Santa Clara (Britton & al. 4776 [MO [3 sheets, cross-labelled later], NY [4 sheets], MEXU]). — Lit: Berger (1915: 273–274); Álvarez de Zayas (1996: 121–125, with ills.). Distr: C Cuba. A polymorphic species, which occurs abundantly and prolifically at anthropogenic sites. It differs from A. acicularis esp. in its broadly lanceolate leaves with decurrent terminal spine, and from A. grisea in its shorter leaves. Álvarez de Zayas (1995: 41) and Álvarez de Zayas (1996), and the first edition of this handbook recognize 2 heterotypic subspecies, while AcevedoRodríguez & Strong (2012: 85) and Govaerts (2014+) place both in the synonymy of the species. A. brittoniana ssp. brachypus (Trelease) A. Álvarez (Fontqueria 44: 121, 1996). Type: Cuba (Britton & al. 6183 [MO, NY [2 sheets]]). — Lit: Trelease (1913: 45, t. 99: 1, as var.); Álvarez de Zayas (1996: 121–125, with ills.). Distr: C Cuba (Alturas de Santa Clara); on skeletal soils and serpentine rocks in xeromorphic shrublands and derived secondary formations.  Agave brittoniana var. brachypus Trelease (1913). [2q] Differs from ssp. brittoniana: L tip canaliculate, normally with small denticles at the

Agave AGAVACEAE

inner margin; Inf somewhat laxer; Fl smaller; Fr more cylindrical. The shorter pedicels given as diagnostic in the protologue are not a constant feature (Álvarez de Zayas 1996). — The basionym could be regarded as a provisional and hence invalid (ICN Art. 36.1b) name, but Trelease’s illustration and the adjacent caption leaves no doubt that he fully accepted the taxon. The lectotypification proposed by Álvarez de Zayas (1996: 121) is unnecessary and moreover erroneous, since an element not originally included by Trelease is selected. A. brittoniana ssp. brittoniana — Lit: Trelease (1913: 44–45, tt. 98–99); Berger (1915: 273–274); Álvarez de Zayas (1996: 121–124, with ills.). Distr: C Cuba; on limestone, shale and slate, in evergreen forests and montane rain forests, primarily abundant in secondary formations and heavily disturbed sites, 100 to >1000 m; flowers January to March. I: Richter (2011: 92, 137); Hochstätter (2015: VII: 90); both as A. brittoniana. [2q] Acaulescent, Ros solitary; L broadly lanceolate, abruptly acute towards the tip, (70–) 80–100 (–110)
(13–) 15–20 (–24) cm, green, sometimes somewhat greyish, slightly glossy, margins often concave between the teeth; marginal teeth variously curved, 4–6 (–8)
3–6 (–7) mm at mid-leaf, (6–) 8–10 (–15) mm apart, slender-tipped from lenticular or voluminous bases, bases sometimes on retrorse green prominences in the lower of the lamina; terminal Sp unguiculately curved, subconical or involutely much thickened in the basal parts, openly grooved to the middle or involute, smooth, somewhat polished, (10–) 15–20 (–25)
4–6 mm, brown, dotted with white,  decurrent; Inf (4–) 5–8 m, paniculate, peduncle very short or nearly none, partInf ascending, (11–) 15–30 cm, not bulbilliferous; Ped slender, 5–10 mm; Fl 25–35 (–45) mm; Tep yellow, tube conical, sometimes very open, 3–6
5–8 (–10) mm, lobes 9–14 (–16)
3–5 mm, dorsally greenish; Fil 18–25 mm, inserted nearly in the tube throat; Anth 9–15 mm; Ov fusiform, 15–20 (–25) mm; Sty longer than the filaments; Sti capitate; Fr oblong, sometimes nearly cylindrical, 23–40 (–45)
11–15 (–17)

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mm, dark chestnut-brown, tip slightly beaked, basally strongly stipitate; Se semicircular to subtriangular, 5–6
3–4 mm. Occurs at least sometimes in regions with a predominance of ophiolitic soils (López Almiral 2013). A. brittoniana ssp. sancti-spirituensis A. Álvarez (Fontqueria 44: 125, ill. (p. 122), 1996). Type: Cuba, Sancti Spíritus (Jiménez & al. 69532 [HAJB]). — Distr: C Cuba (Montañas de Sancti Spíritus); on calcareous and shale rocks in montane rain forests and derived secondary formations; flowers in February. [2q] Differs from ssp. brittoniana: L much shorter, more broadly oblong and less lanceolate; Fl 36–46 mm, with longer tepals and anthers. A. brunnea S. Watson (Proc. Amer. Acad. Arts 26: 156, 1891). Type: Mexico, Coahuila (“Tamaulipas”) (Pringle 2218 [GH, US [photo]]). — Lit: Rose (1903c: 19); Berger (1915: 31–32); Verhoek-Williams (1975: 187–191); Piña Luján (1985: 28–29, 59–61, with ills.); GarcíaMendoza (2003: with ill.); Castillejos-Cruz (2009: [109]–[114], with ills.); all as Manfreda except Berger. Distr: Mexico (SE Chihuahua, N, NW, W & C Coahuila, N, NE & E Durango); dry hills or desert plains, volcanic or alluvial alkaline soils (sandy or gravelly clay), in desert scrub, grasslands, and low pine forests, 1000–1700 m; flowers late June to July. I: Hochstätter (2016: I: 15, as Manfreda).  Manfreda brunnea (S. Watson) Rose (1903)  Polianthes brunnea (S. Watson) Shinners (1966). [3a1] Herbaceous, robust; corm (1–) 2–5.5
0.9–2.5 cm, reproducing vegetatively by buds below the leaf bases; bulb cylindrical to ovoid, 3–7.5
1.5–4 (at the base) cm, completely covered with membranous dry leaf bases fraying into fine fibres at the tip, (4.5–) 5.5–9.5 cm; R fleshy and fibrous, contractile, with filiform ramifications; L evergreen, succulent, 4–8, linear-lanceolate to broadly lanceolate, erect to semi-erect, recurved to undulate, channelled above, tip acute, with a long point, smooth, 10–33
1.2–2.9 (–4) cm, light green to glaucous, intensely mottled with

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purple to red above, both faces verrucose, margins with scattered, broad, hooked, cartilaginous teeth, usually large, deltoid or truncate-bipartite, 2.5 mm, 3–11 mm apart, with narrow pale hyaline band on the leaf margin between the teeth; Inf 0.7–1.3 m, ‘spicate’, green with purplish tinge, peduncle with lanceolate, narrowly attenuate Bra shorter than the internodes, fertile part (5–) 14–25 (–29) cm, dense, with 9–30 solitary flowers; Fl 30–60 (–75) mm, sessile, ascending to appressed to the axis; Tep membranous to semisucculent, tube narrowly funnel-shaped, straight, gradually constricted above the ovary, 30–45
5–6 mm, outer face yellowish-green, inner face brown, lobes oblong to narrowly oblong, spreading to revolute, tip obtuse, cucullate, with a tuft of short hairs, 8–18
4–6 mm, greenish-red to brown with purple tinge; St much exserted; Fil 45–65 (–70) mm, exceeding the tube for 40–60 mm, inserted at near the mouth of the tube, greenishred to cherry-red; Anth 11–17 mm, reddish to yellowish-green; Ov cylindrical to long-ellipsoid, 10–20 (–23)
4–6 mm; Sty (50–) 70–100 mm, exserted for 35–53 (–65) mm; Sti clavate, 3-lobate, deeply furrowed, yellowish-green with small purple dots; Fr ellipsoid to subcylindrical, 18–30 (–36)
12–16 mm, without scar; Se deltoid, plano-concave, 4–5
3–4 mm, shiny, somewhat rough. Easily distinguished from the other species in Ser. Brunneae of Sect. Polianthes by its long narrow tepal tube with exserted stamens and styles and by the coarse teeth on the leaf margins (Verhoek-Williams 1975: 190). The corm is used as soap and shampoo, and the leaves, corm and bulb were formerly used as antidote against snakebites (Castillejos-Cruz 2009). “One of the most attractive manfredas” which “definitely requires a dry winter—early spring rest” (Hannon 2002: 251). A. bulbulifera (Castillejos & E. Solano) Thiede (Haseltonia 17: 94, 2012). Type: Mexico, Guerrero (Castillejos & al. 1807 [MEXU, CHAPA, FEZA, IEB, MICH, NY, US]). — Lit: Castillejos-Cruz & Solano (2008); CastillejosCruz (2009: [115]–[120]); both with ills. as Manfreda. Distr: Mexico (Guerrero); slightly

J. Thiede

inclined, somewhat rocky slopes rich in organic material, in pine-oak forests, 1200–1260 m; flowers June to July; only known from the type locality. I: Hochstätter (2016: I: 16, as Manfreda).  Manfreda bulbulifera Castillejos & E. Solano (2008). [3a3] Herbaceous; corm compressed, 0.5–0.7 cm, forming bulbils 0.5–1.5 cm long at its distal part, bulb ovoid, 3–5
2.5–3 cm, covered with dry leaf bases 4.5–6 cm long, membranous below and fibrous above; R fleshy, fibrous, contractile; L drought-deciduous?, 2–4, prostrate or semierect, linear to lanceolate, attenuate, channelled, glabrous, tip acute, 12–25
0.3–0.6 cm, greenyellowish to dark green, surface often slightly rough, margins hyaline, narrow, entire, partly microscopically papillose; Inf erect, 0.7–0.8 (–0.86) m, ‘spicate’, peduncle 0.63–0.74 m, with 5–7 Bra similar to the leaves, fertile part 7–8.5 (–12) cm, with 7–10 solitary, laxly arranged flowers; Ped 1–3 mm; Fl 25–32 mm; Tep greenyellowish to green-whitish, tube minute, 0.5–1
2–3 mm, lobes linear, erect or revolute, 25–30 (–35)
15–30 mm, apex thickened, cucullate, with a bundle of whitish hairs; St 50–60 mm, reddish; Ov obclavate, 12
2.3 mm, greenyellowish; Sty 50–68 mm; Sti 3-lobate, greenyellowish; Fr subglobose, triquetrous, 10–12
11–12 mm, apex without scar; Se 3–4
3 mm, black, dull. Belonging to Ser. Guttatae of Sect. Manfreda and most similar to A. guttata, A. littoralis and A. pratensis according to the protologue (as Manfreda). It differs from these in its compressed corms forming bulblets at their distal portion, its very short tepal tube, filiform tepal lobes, stamens and style by far exceeding the perianth in length, and an obclavate ovary. A. bulliana (Baker) Thiede & Eggli (Kakt. and. Sukk. 50(5): 112, 1999). Type: Mexico (Karwinsky s.n. [not preserved]). — Lit: Rose (1903b: as Prochnyanthes); McVaugh (1989: 260–263, as P. mexicana); Castro-Castro & al. (2010: as P. mexicana); Thiede (2012a); all with ills. Distr: Mexico (Durango, Zacatecas, Nayarit, Aguascalientes, Jalisco, Guanajuato, Querétaro, Michoacán); dry rocky slopes or roadcuts, in

Agave AGAVACEAE

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Fig. 12 Agave bulliana (Etter & Kristen 1353: Mexico; Aguascalientes,  20 km SE of Aguascalientes, 2270 m). (Copyright: J. Etter & M. Kristen)

Fig. 13 Agave bulliana (Etter & Kristen 2735: Mexico; Jalisco,  20 km SE of Concepción Buenos Aires, 1860 m). (Copyright: J. Etter & M. Kristen)

pine-oak grasslands, or in shaded moist ravines, 1150–3100 m; flowers late June to early September. I: Curtis’s Bot. Mag. 121: t. 7427, 1895, as Prochnyanthes; Howard (2001: 13, as P. mexicana); Hannon (2002: as P. mexicana); Etter & Kristen (2012: 85); Hochstätter (2016: II: 31–32, as P. mexicana); Castro-Castro (2017: 139, as P. mexicana). – Figs. 12 and 13.  Bravoa bulliana Baker (1884)  Prochnyanthes bulliana (Baker) Baker (1895); incl. Polianthes mexicana Zuccarini (1837)  Prochnyanthes mexicana (Zuccarini) Rose (1903); incl. Prochnyanthes viridescens S. Watson (1887). [3b2] Herbaceous, large (for Subgen. Manfreda), usually single; corm cylindrical, 1–3
1.5–2.5 cm, bulb oblong, covered with fibrous

dried leaf bases 7–10 (–12.5) cm long; R fusiform, semifleshy with a wiry core, contractile; L few, deciduous, (1–) 2–3 (–7), the larger in a basal Ros, upwards diminishing up to the small Bra, chartaceous, thin, fibrous, erect or occasionally curved backwards from about the middle, often twisted, with a distinct midrib, shallowly channelled over the midrib, lamina flat, broadly undulate, or revolute, linear-lanceolate to oblanceolate, narrowed towards the base, (8–) 20–47 (–65)
(0.7–) 1.3–5.2 (–8) cm, light or dark green, dull, often speckled with magenta towards the base, spotted or not, veins slightly prominent on both surfaces, papillate, margins very narrow, hyaline, papillate to erose-papillate or papillatedenticulate, tip a soft Sp; Inf (0.6–) 0.9–2 (–2.8)

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m and more, ‘spicate’, fertile part elongate, (9–) 17.5–47 (–83.5) cm, with (2–) 4–22 (–25) flowering nodes with geminate flowers; Ped (0.2–) 25–40 (–90) mm; Fl functionally pendent by an abrupt curve in the tepal tube; Tep white tinged with grey-green or dull green and red, white or creamy within, tube curved near the middle or at from the ovary, narrow below, 5–17.5
3–8 mm, abruptly widened above the bend, upper part (11–) 15–27
5–20 mm, lobes flaring, broadly deltoid, (3–) 4–9 (–15)
3–12 mm, tips pubescent; Fil 8–20 mm, inserted near the base of the bend of the tube; Anth linear, 3–15 mm; Ov ellipsoid, 4–8 (–12) mm; Sty filiform, 12–32 mm, finally equalling the tube or longer, white; Sti 0.7–4 mm, 3-lobate; Fr globose to oblong, 8–25
10–17 mm; Se deltoid to semicircular, 3.5–6
2.5–3, black. For some time recognized as monotypic genus Prochnyanthes, separated from Polianthes on account of the downward-curved perianth tube widening above the bend. The detailed morphometric studies of Castro-Castro & al. (2010) confirmed that Prochnyanthes consists of a single species only with a considerable variability in floral and other features. When treating Prochnyanthes mexicana as Agave, a change of name was necessary to avoid homonymy with the earlier A. mexicana Lamarck 1783. Therefore, the second-oldest epithet (Prochnyanthes bulliana) had to be chosen (Thiede & Eggli 1999). In the first edition of this handbook, the former genera Prochnyanthes and Polianthes were placed as distinct informal groups within Subgen. Manfreda, but the recently described Agave (Polianthes) oaxacana is morphologically intermediate, and Prochnyanthes was also included in Polianthes by García-Mendoza & Solano (2007: 115–116). The history, introduction, horticulture, biology and taxonomy of A. bulliana was summarized by Thiede (2012a). Guillot Ortiz & Meer (2006b) erroneously stated that they introduced the species for the first time into European horticulture, but it was in fact cultivated much earlier at Munich in 1837 and at Kew in 1884 (Thiede 2012b). The ethnobotany and uses of the species are dealt with by Verhoek-Williams (1978b).

J. Thiede

A.
bundrantii (T. M. Howard) Thiede & Eggli (Kakt. and. Sukk. 50(5): 111, 1999). Type: not typified. — Lit: Howard (1978: with ill.). Distr: Cultivated only. I: Hochstätter (2016: II: 27).  Polianthes
bundrantii T. M. Howard (1978). [3b1
3b2] This is the garden hybrid A. amica (as Polianthes tuberosa)
A. howardii (as Polianthes howardii), still extant in cultivation. Solano Camacho (2002: 55, as Polianthes) erroneously lists the hybrid (as “P. brundrantii”) in the synonymy of P. howardii. ‘Chirp’, ‘Opal Eyes’ and ‘Mexican Firecracker’ are commercially offered cultivars (Guillot Ortiz & al. 2006, Hochstätter 2016: II: 27, both with ills.). A. cacozela Trelease (Mem. Nation. Acad. Sci. 11: 41, tt. 89–91, 1913). Type: Bahamas, New Providence (Cunningham s.n. [not located]). — Lit: Berger (1915: 204); Britton & Millspaugh (1920: 75); Correll & Correll (1982: 312). Distr: Bahamas (New Providence); rocky margins of salt marshes; flowers mostly January to March. I: Hochstätter (2015: VII: 83). [2r] Acaulescent; Ros solitary; L lanceolate, fleshy, curved, deeply guttered, typically roughish, 150–200
20 cm, yellowish-green, somewhat overcast with grey; marginal teeth nearly straight or the larger teeth appressed-recurved, narrowly triangular, rarely lenticular at the base, 2–5 mm (middle of the lamina), usually 10–15 mm apart, intervening margin straight or somewhat concave; terminal Sp triquetrously conical, straight or the tip slightly curved, openly grooved below the middle, smooth, 15–20
3–5 mm, brownish becoming grey, dull, decurrent; Inf 6–7 m, paniculate, peduncle with broadly triangular non-imbricate Bra, fertile part dense, ovoid, part-Inf on horizontal or slightly ascending Br, in the upper of the inflorescence, reportedly sometimes bulbilliferous; Ped 10 mm; Fl 50–60 mm; Tep golden-yellow, tube rather open, 7 mm, lobes 20
4–5 mm; Fil 40–45 mm, inserted nearly in the tube throat; Ov oblong-fusiform, 35–40 mm; Sty longer than the filaments; Fr narrowly oblong, 35–45
15 mm, beaked, shortly conical-stipitate; Se 6–7
4 mm.

Agave AGAVACEAE

According to the protologue, the leaves of seedlings of the type collection were decidedly papillate-roughened on both faces. See also the note under A. bahamana. A. cajalbanensis A. Álvarez (Revista Jard. Bot. Nac. Univ. Habana 1(2–3): 33–39, ill., 1981). Type: Cuba, Pinar del Río (Bisse & Álvarez 32466 [HAJB [4 sheets]]). — Lit: Álvarez de Zayas (1985: 9, 13); Urquiola Cruz & al. (2010: 35, with ill.); González-Oliva & al. (2014: 11). Distr: W Cuba (Pinar del Río: Sierra de Cajálbana); denuded slopes of ultrabasic rocks, in pine forests and xeromorphic thorn scrubs; 100–400 m. I: Pilbeam (2013: 48); Hochstätter (2015: VII: 68). [2p] Stems short; Ros solitary, semiglobose, to 1.2 m ∅; L many, oblanceolate in the lower , straight, fleshy, coriaceous, only slightly concave, 50–100
8–10 cm, grey-green, somewhat dull, margins with asymmetrical slightly recurved prominences, these 3
4–5 mm, intervening margin nearly straight; marginal teeth basally slightly recurving, 2–4 mm, dark chestnut-brown to nearly black, 10–20 mm apart; terminal Sp conical, straight, basally flattened, 10–15 mm, dark chestnut-brown, not lustrous, not decurrent; Inf 3–6 m, paniculate, part-Inf 3-parted, broad, slightly incurved, 30–40 cm; Ped 18–25 (–35) mm; Fl 40–45 mm; Tep orange or yellow, tube open, 5–6 mm, lobes with broad margins, 12–15 mm, inner membranous, outer dorsally sulcate; Fil 15–20 mm, filiform, inserted near the base of the tube; Anth 10–12 mm; Ov fusiform, trigonous, basally constricted, 15–20
5–7 mm; Fr oblong, apically acute or apiculate, 20–25
15 mm; Se unknown. Easily identifiable by its somewhat lobed leaf margins, the recurved marginal teeth, its orange (or yellow) flowers and small fruits. It is closest to A. grisea, but the leaves and inflorescences are only as large according to the protologue. It differs from A. legrelliana (another species with broad panicles and orange flowers from W Cuba) by its greyish leaves 15 mm; Inf with 20–35 partInf; Fl shorter, 45–63 mm; Ov shorter, 25–30 mm, neck short; Tep green, lobes subequal, 15–21 mm, light greenish-yellow. Gentry (1982: 569) identified a cultivar from El Salvador as Trelease’s A. acuispina and placed it here. A. cantala var. cantala — Lit: Berger (1915: 236–237); Trelease (1920: 119); Gentry (1982: 568–570, with ills.); McVaugh (1989: 146). Distr: Cultivated worldwide, esp. in SE Asia; not known from the wild. I: Todaro (1876–92: t. 15, as A. candelabrum); Morris (1889: as A.

J. Thiede

candelabrum); Richter (2011: 77); Hochstätter (2015: VII: 34). Incl. Agave bulbifera Salm-Dyck (1834); incl. Agave laxa Karwinsky ex Otto (1842) (nom. illeg., ICN Art. 53.1); incl. Agave rumphii Hasskarl (1843); incl. Agave vivipara Dalzell & A. Gibson (1861) (nom. illeg., ICN Art. 53.1); incl. Agave candelabrum Todaro (1878). [2g] Stems 30–60 cm; Ros tall, slender, laxly leafy, 2–2.5 m ∅, surculose; L linear, longacuminate, thin, frequently reflexing, guttered, roundly keeled below towards the base, rough below, smooth above, 150–200
7–9 cm, light or dark green, margins straight; marginal teeth antrorsely curved, larger teeth 3–4 mm, brown, mostly 20–30 mm apart, reduced or lacking towards the leaf tip; terminal Sp very small, 5–15 mm; Inf 6–8 m, paniculate, peduncle slender, with narrow chartaceous Bra soon becoming reflexed, part-Inf ascending to recurved, lax, 20, on slender stalks, in the upper of the inflorescence, freely bulbilliferous; Fl slender, 70–85 mm; Tep greenish tinged purple or reddish, tube cylindrical-funnelform, grooved, 14–17
15 mm, lobes subequal, linear-spatulate, rapidly involute-conduplicate, 25–28 mm, apex roundedcucullate, inner with narrow keel; Fil stout, abruptly outcurving from the tube wall, 50–55 mm, inserted 11–14 mm above the tube bottom; Anth excentric, slender, slightly curved, slatey purple; Ov fusiform, tapering below to a basal rim, roundly 6-ribbed, 32–42 mm, shortly constricted above but virtually neckless; Sty stout, shorter than the filaments; Sti capitate; Fr and Se unknown. — Cytology: 2n = 77–97 (3
) (Lv & al. 2009, Simpson & al. 2011). Recognizable by the thin, long, narrow, weak leaves which are frequently reflexed above the middle, and by small teeth and large purplishgreen flowers in broad, diffuse, slender-peduncled panicles which are only rarely produced (Gentry 1982). Roxburg published it from India whereto it was most certainly brought from North America by European traders. It has apparently been cultivated since long for fibre in SE Asia from the Philippines to India and is still hardly known in the Americas, where it remained unrecognized for long (Gentry 1982). Gentry (1982) collected

Agave AGAVACEAE

offsets from a plant at Villa Guerrero, Jalisco, very much looking like A. cantala and said to be spontaneous there. No plants from W-C Mexico were known to McVaugh (1989), and Hernández-Vera & al. (2007a) could not find A. cantala near Villa Guerrero. The triploid chromosome number explains the sterility of the species. According to Trelease (1920) it is related to A. tequilana (which originated from A. angustifolia) and according to Valenzuela-Zapata & Nabhan (2003) it is merely a domesticated taxon obtained from A. angustifolia, for both of which triploid cultivars are known. A.
cavanillesii D. Guillot & P. Van der Meer (Flora Montiber. 28: 73–74, ill., 2004). Type: Spain, Valencia (Guillot s.n. [VAL]). — Lit: Guillot Ortiz & al. (2008: 81, with ill.). Distr: Cultivated only. I: Hochstätter (2015: VII: 101). [2g
2g] A spontaneous hybrid A. decipiens
A. fourcroydes found among its naturalized parental species in Spain. A. caymanensis Proctor (Fl. Cayman Islands, ed. 2, 183, fig. 71, t. 7, 2012). Type: Cayman Islands (Proctor 52171 [IJ]). — Lit: Proctor (1984: 241–242); Hummelinck (1984: 208–212, 226–228); both with ills., as A. sobolifera. Distr: Cayman Islands (Grand Cayman, Little Cayman, Cayman Brac); dominant in dry rocky situations, semi-deciduous forests and deciduous shrublands with Bursera and Pilosocereus. I: Burton (2005: 168, 170, 174); Hochstätter (2015: VII: 69, 78). [2p] Acaulescent when young, at maturity commonly with woody stems to 1 m clothed by dead leaves; Ros solitary; L 30, elliptic-oblanceolate to (narrowly) lanceolate, fleshy, massive, somewhat S-shaped, guttered, sometimes folded, narrowed at the base, acuminate at the apex, to 125–150
20–25 cm (above the middle),  5
as long as broad, medium to dark green; marginal teeth curved or reflexed-triangular (rarely straight), 1–7 mm, glossy dark brown, 5–15 mm apart, usually 6–9 per 10 cm at mid-leaf, often from green prominences of the margin 1.5–3 mm high, very small below the terminal spine, intervening margin concave; terminal Sp slenderly conical to subulate, lower usually clearly grooved between the

79

converging leaf margins, 17–23
2.5–4 (at the base) mm, decurrent only in outer leaves; Inf to 6 m or more, paniculate, peduncle 3.5 m, with 30 Bra, fertile part densely paniculate towards the apex, 2 m, 2.5
as long as broad, with up to 15 part-Inf, longest 40 cm, main one 24 cm, freely bulbilliferous; Ped 5–10 mm; Fl 41–51 mm; Tep bright yellow, tube 3–5.5 mm, lobes expanded at the base, narrowing to an elongate apex, 12–20 mm; Fil 30–41 mm, inserted 0.5–2.5 mm below the mouth of the tube; Ov narrowly fusiform, 25–40
7–8 mm; Sty 30–35 mm, almost as long as the filaments; Sti expanded; Fr ovoid, 28
17 mm, stipitate for 6 mm, very shortly apiculate; Se crescent-shaped, 6–7
4–4.5 mm. Plants now placed here were included in the Jamaican A. sobolifera by earlier authors such as Adams (1972), Proctor (1984) and Hummelinck (1984). A. caymanensis differs, according to the protologue, by forming woody stems at maturity, broader leaves in relation to their length, and by flowers with a much longer ovary (30–40 mm vs. 15–25 mm), shorter tepal lobes, and a longer, exserted style. Fruits and seeds were not described in the protologue, and data is here added from the published figure. Hummelinck (1984) carried out a detailed field study of Agave in the Cayman Islands. His detailed descriptions and many photographs supplement the short protologue and leave the purported differences less clear: The ovary is given as 25–32 mm (vs. 25–35 mm in Jamaican A. sobolifera), the lobes as 13–20 mm (vs. 20–25 mm), and the style as 30–35 mm (vs. 39–55 mm). Thus, the distinction of A. caymanensis is in need of re-study against more ample Jamaican material of A. sobolifera. Burton (2005) published 3 habitat photographs and used the name A. caymanensis 7 years before it was formally published. The Vitelline Warbler (Setophaga vitellina) feeds on nectar from the flowers (Bradley & Rey-Millet 2013: 236). A. cerulata Trelease (Annual Rep. Missouri Bot. Gard. 22: 55, 1912). Type: Mexico, Baja California (Nelson & Goldman 7180 [US]). — Lit: Berger (1915: 262, 264); Gentry (1978: 35–48, with ills.); Gentry (1982: 363–375, with

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ills.); Turner & al. (1995: 44–45, with ills.); Navarro-Quezada & al. (2003); Starr (2012: 69–73, with ills.); Webb & Starr (2015: 77–81, with ills.). Distr: Mexico (Baja California, Baja California Sur). Molecular population studies including 5 samples of all 4 subspecies of A. cerulata showed a clade that also includes one sample of A. deserti, and which is nested within further clades with A. deserti and A. subsimplex, thus questioning the separation of these species (Navarro-Quezada & al. 2003). — The leaf shape of A. cerulata changes geographically with climate, and the leaf size decreases with increasing aridity (Burgess 1986). A. cerulata ssp. cerulata — Lit: Trelease (1912b: 45, 55, tt. 45–47); Berger (1915: 264); Gentry (1978: 35–43, with ills.); Gentry (1982: 363–369, with ills.); Turner & al. (1995: 44–45); Starr (2012: 69–73, with ills.); Webb & Starr (2015: 77–78, with ill.). Distr: Mexico (C & S Baja California, N-most Baja California Sur); on hills and rocky mountain sides, in desert scrub with frequent fog and sea breezes, 25–640 m; flowers April to June. I: Irish & Irish (2000: t. 10); Heller (2006: 72); Richter (2011: 102); Pilbeam (2013: 51); Hochstätter (2015: V: 1, 14). [2i] Ros 0.2–0.5
0.4–0.75 m, abundantly surculose or rarely solitary; L few, narrowly lanceolate to triangular-lanceolate, long-acuminate, mostly 25–50
4–7 cm and 5–12
as long as broad, yellow to light green, sometimes crosszoned, rarely light glaucous-grey, margins nearly straight to shallowly repand; marginal teeth weakly attached, 1–4 mm, greyish-brown, bordered with a brown ring at the base, irregularly spaced, on low tubercles, sometimes lacking through much of the leaf length; terminal Sp acicular, 30–60 mm, light to dark grey, decurrent only to the uppermost teeth or less; Inf 2–3.5 m, paniculate, peduncle with small scarious triangular Bra, fertile part slender, narrow, part-Inf small, mostly 6–12, all white waxy-glaucous at bud stage; Fl mostly 45–60 mm; Tep in bud white waxy-glaucous, at anthesis pale yellow, tube broadly funnel-shaped or discoid, with thick Nec and bulges opposite the filament insertions, 3–5
11–14 mm, lobes

J. Thiede

ascending to spreading, elliptic, equal, 16–22 mm, the inner wide with low broad keel and broadly overlapped at the base by the outer lobes; Fil usually incurved at anthesis, variable in length, 30–40 mm, inserted at the base of the lobes on the rim of the nectary; Anth centric to excentric, 15–20 mm; Ov fusiform, narrowly tapering towards the base, 22–32 mm; Sty shorter than the filaments; Sti capitate; Fr narrowly oblong, 30–50
12–13 mm, waxy light grey, bluntly apiculate, narrowly stipitate; Se crescent-shaped, hilar notch open or obscure, marginal wings pronounced on both sides around the curvature, 5
3 mm, sooty black. Mostly characterized by slender, yellow, long-acuminate, lanceolate leaves with a brown ring at the weakly attached, moderate to small teeth, small narrow panicles with a conspicuous waxy bloom on peduncle, pedicels and fruits, and light-yellow, spreading tepal lobes. A. cerulata may be confused with the closely related A. deserti, but the latter is more robust, has grey-green (rather than yellow-green) leaves 4–7
longer than broad (vs. 5–12
longer than broad) and generally without the distinctive brown rings at the base of the teeth, and the capsules are generally broader and without waxy white bloom (Gentry 1982) (see A. deserti for further notes). Breslin (2015) provides a new record from Isla Cedros. A. cerulata ssp. dentiens (Trelease) Gentry (Occas. Pap. Calif. Acad. Sci. 130: 43, 1978). Type: Mexico, Baja California (Rose 16819 [MO, NY, US]). — Lit: Trelease (1912b: 45, 51, tt. 38–40, as A. dentiens); Berger (1915: 264, as A. dentiens); Gentry (1978: 43–44, tt. 2–3); Gentry (1982: 369–371, with ills.); Turner & al. (1995: 45, with ill.); Wilder & al. (2008: 135–136, ill. p. 140); Guillot Ortiz & Meer (2008b: 24, ill. p. 30); Starr (2012: 70, 73); Webb & Starr (2015: 77–80, with ill.). Distr: Mexico (Baja California: Isla San Esteban); hillsides and arroyos; budding in April. I: Heller (2006: 73); Pilbeam (2013: 52); Hochstätter (2015: V: 15);  Agave dentiens Trelease (1912). [2i] Differs from ssp. cerulata: Ros 0.5–0.7
0.8–1.5 m; L long-acuminate, 40–55 (–70) cm,

Agave AGAVACEAE

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Fig. 14 Agave cerulata ssp. subcerulata (Etter & Kristen 383: Mexico; Baja California Sur, Vizcaino Desert, 170 m). (Copyright: J. Etter & M. Kristen)

light glaucous-grey, margins straight; marginal teeth friable, 1–2 mm, or nearly toothless; terminal Sp acicular, 30–50 mm, brown to grey; Inf broad, part-Inf 8–18 in the upper of the inflorescence, on 30–40 cm long Br. An isolated island population closely related to typical A. cerulata. With its broad inflorescences, it could bridge the gap between Sect. Deserticolae and Sect. Conicae (Webb & Starr 2015: 80). — Naturalized at one locality in Spain (Guillot Ortiz & Meer 2008b). A. cerulata ssp. subcerulata Gentry (Occas. Pap. Calif. Acad. Sci. 130: 44–48, ills. (pp. 46–47, 49), 1978). Type: Mexico, Baja California Sur (Gentry 10330 [US, ARIZ, MEXU]). — Lit: Trelease (1912b: 45, 51, 55, tt. 30–40, 45–47); Gentry (1978: 44–48, with ills.); Gentry (1982: 371–374, with ills.); Turner & al. (1995: 45); Starr (2012: 70); Webb & Starr (2015: 81, with ills.). Distr: Mexico (S-most Baja California, N Baja California Sur); volcanic or gypsum slopes in desert scrub; flowers April to May. I: Heller (2006: 72); Richter (2011: 80, 141); Pilbeam (2013: 54); Hochstätter (2015: V: 17). – Fig. 14. [2i] Differs from ssp. cerulata: Ros 0.15–0.3
0.3–0.5 m; L short-acuminate, 15–30
2.5–7 cm, mostly 3–6
as long as broad, mostly light grey to bluish-glaucous over green, margins conspicuously crenate with prominent tubercles; marginal teeth well developed, weakly attached, larger,

3–8 mm in the middle of the lamina; terminal Sp subulate, usually sinuous, 20–40 mm. The S-most element in the A. cerulata complex, resembling A. subsimplex from the opposite Sonoran coast in habit (Gentry 1982). It might represent a genetic connection to the more southern A. sobria (Webb & Starr 2015). A. cerulata var. nelsonii (Trelease) R. H. Webb & G. D. Starr (Haseltonia 20: 80, ill., 2015). Type: Mexico, Baja California (Nelson & Goldman 7111 [MO]). — Lit: Trelease (1912b: 45, 61, tt. 65–67, as A. nelsonii); Gentry (1978: 44–46, with ills.); Gentry (1982: 371–373, with ills.); Turner & al. (1995: 45, with ill.); Starr (2012: 70–71, ill.); the last 4 as ssp. nelsonii; Webb & Starr (2015: 78–81, with ill.). Distr: Mexico (C Baja California); igneous highlands in desert scrub or chapparal vegetation, 140–1120 m; flowers June to September. I: Goldman (1916: t. 111D, as A. nelsonii); Heller (2006: 73); Richter (2011: 102, 103); Pilbeam (2013: 53); Hochstätter (2015: V: 16). – Fig. 15.  Agave nelsonii Trelease (1912)  Agave cerulata ssp. nelsonii (Trelease) Gentry (1978); incl. Agave shawii E. C. Nelson (1911) (nom. illeg., ICN Art. 53.1). [2i] Differs from ssp. cerulata: Ros 0.5–0.75 m ∅; L short-acuminate, 20–35
6–8 cm, mostly 3–6
as long as broad, mostly light grey to bluish-glaucous over green, margins either repand and with small prominences, or nearly straight;

82

J. Thiede

Fig. 15 Agave cerulata var. nelsonii (cult.: USA; Arizona, Arizona-Sonora Desert Museum Tucson). (Copyright: J. Thiede)

marginal teeth firmly attached, larger, 3–9 mm, frequently on small tubercles; terminal Sp strongly subulate, 20–40 mm; Inf with 15–20 part-Inf. The N-most element in the A. cerulata complex. Recognized at subspecies rank by Gentry, but reduced to varietal status by Webb & Starr (2015) since its distribution shows substantial geographical overlap with ssp. cerulata at its S end. Webb & Starr speculate that the taxon might represent a stabilized hybrid between A. cerulata and A. shawii ssp. goldmaniana. Hochstätter (2015: V: 16) depicts a natural hybrid with the latter. A. chamelensis (E. J. Lott & VerhoekWilliams) Thiede & Eggli (Kakt. and. Sukk. 50 (5): 110, 1999). Type: Mexico, Jalisco (Lott & Wendt 1663 [MICH, BH, CAS, MEXU]). — Lit: Lott & Verhoek-Williams (1991); Castillejos-Cruz (2009: [121]–[126]); both with ills., both as Manfreda. Distr: Mexico (Jalisco, Oaxaca); uncommon along arroyos in tropical (semi-) deciduous and oak forests, 25–75 (–1170) m; flowers December to February. I: Hochstätter (2016: I: 17); Castro-Castro & al. (2017: 62); both as Manfreda.  Manfreda chamelensis E. J. Lott & VerhoekWilliams (1991). [3a2] Herbaceous; corm 3–15
2–3 cm, reproducing vegetatively by buds from the corm, bulb shortly cylindrical, 2–3.5
1.5–3 cm,

covered with membranous dry leaf bases 2.5–3.5 cm long and not separating into fibres; R fleshy and fibrous, with filiform ramifications, contractile; L drought-deciduous, 9–12, linearlanceolate, spreading, brittle, herbaceous to somewhat fleshy, narrowly channelled, nearly conduplicate near the narrow base, tip acute, veins papillate on both faces, 37–77 (–91)
(1–) 1.6–4.8 (–6.5) cm, margins with a narrow yellowish cartilaginous band, minutely denticulate, teeth regular; Inf 0.75–1.2 (–2) m, ‘spicate’, fertile part 10–20 cm, with 10–25 (–35) sessile flowers; Fl nearly erect at anthesis; Tep green, tube funnelshaped, 6–13
4–5 mm, lobes oblong, reflexed to tightly revolute, tip obtuse, with a tuft of short hairs, 8–11
4–5 mm; Fil erect, filiform, 21–32 mm, inserted in the middle of the tube, greenish-yellow with many purple dots; Ov oblong to ovoid, not constricted, not protruding into the tepal tube, 5–10 mm; Sty 25–35 mm, exserted; Sti clavate, trigonous; Fr globose, 12–16
10–15 mm; Se deltoid, plane-concave, 5–6
4–5 mm, shiny. This species appears to be closest to A. scabra and A. jaliscana. It differs from both by shorter floral bracts, filaments curved near the tip at bud opening, and by the absence of coarse fibrous remains of old leaves, from the first-named by its shorter floral tube and globose beakless capsules, and from the last-named by its wider leaves and shorter styles and filaments (Lott & Verhoek-

Agave AGAVACEAE

Williams 1991, as Manfreda). The tropical lowland habitat is untypical and apparently not known for other species of Subgen. Manfreda except A. littoralis, A. petskinil and A. paniculata. A. chazaroi A. Vázquez & O. Valencia (in Vázquez-García & al. (eds.), Agaves Occid. Méx., 48–49, ills. (tt. CC-DD, pp. lxx-lxxiii), 2007). Type: Mexico, Jalisco (Cházaro Basán˜ ez & al. 8161 [IBUG]). — Distr: Mexico (N Jalisco); basaltic cliffs in tropical deciduous forests, 900–1200 m; flowers January to February. I: Richter (2011: 56); Etter & Kristen (2012: 91–92); Pilbeam (2013: 55); Hochstätter (2015: IX: 26). [1b] Acaulescent; Ros 1.45
0.8 m, solitary, not surculose; L up to 20, rigid, convex below, concave to flat above, to 80
to 20 (mid-leaf) cm, yellowish-green, margins entire and horny dark brown all along; marginal teeth none; terminal Sp thin, large, canaliculate, 30–50 mm, dark brown; Inf erect, to 2.5 m, ‘spicate’, peduncle with filiform Bra, these 20–30
0.5 mm; Ped 10–15 mm; Fl 20–30 mm, geminate, densely arranged in the upper of the inflorescence, funnel-shaped; Tep ivory-white, tube, lobes, Fil and Anth not described; Ov cylindrical, 10 mm incl. neck, cream-white; Fr oblong, 20 mm, glaucous when unripe, turning green (?) when ripe; Se unknown. Based on the inadequately short protologue appearing close to A. pelona from Sonora, which differs esp. by the dark reddish flowers. It differs from other species of Sect. Heteracanthae (as Group Marginatae) in having few and broad leaves, and is better placed in Sect. Inermes (as Group Amolae) (Cuevas Guzmán & al. 2012). Here, it appears to be closest to A. gilbertii from which it differs by its stemless habit, and broader leaves without paler median stripe and with a much longer terminal spine. The tube length is not given in the protologue, but it appears to be very short (cf. t. CC-3), in contrast to 11–12 mm in A. gilbertii and 8–9 mm in A. pelona. A. chiapensis Jacobi (Hamburg. Gart.- & Blumenzeit. 22: 213, 1866). Type [neo]:

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Mexico, Chiapas (Gentry 12178 [US, ARIZ, MEXU, MICH]). — Lit: Jacobi (1871: 80–82); Berger (1915: 44–46, with ill.); Gentry (1982: 224–226, with ills.); Lott & García-Mendoza (1994: online version with ills.); García-Mendoza (2003: with ill.); García-Mendoza (2011a: 22–24). Distr: Mexico (N Oaxaca, Chiapas), Guatemala; limestone cliffs in the Montañas del Oriente, in humid oak, pine, pine-oak or pinejuniper forests or desert scrub-oak forest ecotones, 1400–3000 m; flowers March to April. I: Irish & Irish (2000: t. 11); Hils (2001); García-Mendoza (2002: 179); Heller (2006: 74); Lochner (2010); Richter (2011: 114); Pilbeam (2013: 56); Hochstätter (2015: IX: 4). Incl. Agave chiapensis var. major hort. ex A. Berger (1915); incl. Agave teopiscana Matuda (1974). [1f] Stems short or none; Ros openly spreading, robust, 0.5–0.8
1.3–1.5 m, mostly caespitose, rarely solitary; L 25–36, ovate, rarely lanceolate, soft, ascending, narrowed near the base, shortly acuminate, rounded below, flat to slightly hollowed above and upcurving, smooth, (20–) 30–50 (–65)
7–17 cm, light shiny greygreen or grey-green to glaucous, lower face with imprints from the central bud, margins slightly repand to crenate, sometimes reddish or glaucous; marginal teeth deltoid, straight or upcurved, (1–) 2–4
2–4 mm, closely spaced and (1–) 4–5 (–10) mm apart, blackish or dark brown to greying, sometimes reddish; terminal Sp subulate, straight to sinuous, strong, rigid,  openly grooved above, rounded to broadly keeled below, (15–) 25–40
3–4 mm, blackish or reddish, decurrent for (20–) 25–30 (–40) cm; Inf 2–3.5 m, ‘spicate’, peduncle long, green, closely bracteate with Bra to 13–22
2.5–4 cm, greenish-red or purple, fertile part dense, in the terminal ; Ped 1–2 mm; Fl geminate, trigonous, fleshy, funnel-shaped, (40–) 60–70 (–85) mm, obscured in large tufts of broad-based bracts, the longer bracts equalling the flowers, reddish; Tep yellowish, whitish or green, sometimes tinged reddish or purple, tube funnel-shaped, thick-walled, 3-angled, grooved, 8–12
9–10 mm, lobes oblong or

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linear-elongate, divaricate to recurved, fleshy, thick, unequal, 25–32
4–8 mm, the outer slightly longer, the inner with a prominent keel and involute, all cucullate and glandular at the tip, copiously papillate on and below the hood; Fil very broad at the base, (45–) 65–80 mm, reddish to dark purple-brown, inserted at the tube mouth; Anth (15–) 20–25 (–30) mm, yellow; Ov cylindrical to trigonous, truncate at base and apex, grooved to the base, (10–) 20–25 (–30)
3–4 mm, neck 4 mm, sulcate; Sty very robust, 85–90 (–110) mm, dark purple-brown, thickened below the stigma; Sti weakly 3-lobed; Fr oblong to obovoid, rounded-truncate at the base, shortly apiculate at the apex, 30–40
10–15 mm, thin-walled; Se hemispherical, with regular complete marginal wing, 4–5
3–4 mm, black. Gentry (1982: 226) designated the neotype cited above. The species is characterized by forming large caespitose clones and its ovate leaves with small teeth to 4 mm long, its dense inflorescences with large greenish-red or purple bracts, and its yellowish, whitish or green flowers with reddish to purple filaments (García-Mendoza 2011a). It appears to be closely related to A. warelliana, which differs in its brown (Gentry in error “red”) leaf margins with denticles 1–3 mm apart (Gentry 1982: 226). García-Mendoza (2011a) provides a new record from Oaxaca. A. chrysantha Peebles (Proc. Biol. Soc. Wash. 48(4): 139, 1935). Type: USA, Arizona (Peebles & Harrison 5543 [US]). — Lit: Gentry (1982: 426–431, with ills.); Ullrich (1992i); Turner & al. (1995: 45–47); Hodgson (1999a: 4–5); Reveal & Hodgson (2002); Klopper & al. (2010: 56, 57, 59, with ills.); Parker (2017: 256, 258, with ills.). Distr: USA (C & S-C Arizona); very common on sandy to gravelly places on granitic or volcanic soils in desert scrub, grasslands, pinyon-juniper and oak woodlands, 700–2100 m; flowers June to July (to August). I: Irish & Irish (2000: t. 12); Heller (2006: 65); Richter (2011: 127); Pilbeam (2013: 57); Hochstätter (2015: IV: 6).  Agave palmeri var. chrysantha (Peebles) Little ex L. D. Benson (1943)  Agave palmeri ssp. chrysantha (Peebles) B. Ullrich (1992).

J. Thiede

[2h] Acaulescent, Ros rather open to somewhat dense, 0.5–1.2
0.8–1 m, usually solitary, infrequently caespitose; L linear-lanceolate to lanceolate, spreading to ascending, straight, rigid, usually only a little narrowed below the middle, deeply guttered, 40–75 (–82)
(4–) 8–10 (–11) cm, pale to glaucousgreen or yellowish-green to green, not cross-zoned, margins repand to wavy; larger marginal teeth straight or flexed, 4–10 mm, (8–) 10–30 (–40) mm apart, smaller towards the leaf base, intermittent teeth lacking or 1–3 (–5), mostly in the distal of the leaves; terminal Sp slender, acicular, openly grooved above, 25–45 mm, reddish-brown or brown to grey, decurrent for 5–15 cm to the upper teeth; Inf 4–7 m, narrowly to broadly paniculate, open, peduncle with persistent triangular Bra 1–5 cm long, fertile Inf part narrowly to broadly paniculate, open, part-Inf slightly ascending, small, congested, (6–) 8–18 in the upper of the inflorescence, >10 cm, 12- to 21-flowered; Fl erect, 35–65 (–67) mm; Tep golden-yellow, rarely red-tipped, tube campanulate, 6–18
(7–) 10–21 (–26) mm, lobes erect, persistent and often leathery during and after anthesis, strongly unequal, (6–) 9–15 (–18) mm; St long-exserted; Fil erect, (32–) 35–50 mm, pale yellow, inserted at 2 levels  at mid-tube; Anth 7–20 mm, yellow; Ov slender, (14–) 22–30 (–33) mm, neck slightly constricted, (0.5–) 4–6 (–8) mm; Sty slightly longer than the tepals; Sti capitate; Fr narrowly oblong to obovoid, 35–50 mm, sessile or shortstipitate, apex shortly beaked; Se 6–7 mm. — Cytology: 2n = 60 (Pinkava & Baker 1985). The closest relative is clearly the largely allopatric A. palmeri with often broader leaves with shorter and less distant teeth, and a very similar flower structure, but the latter differs consistently in its clear yellow flowers with tepal lobes conspicuously red to brownish at the tips, its more congested ‘umbels’, and its shorter panicles (Gentry 1982, Klopper & al. 2010). Benson (1943) and Ullrich (1992i) both suggest infraspecific ranks under A. palmeri, with which it shows introgression where they meet (Gentry 1982: 429, 446). A. chrysantha hybridizes with A. delamateri, A. mckelveyana, A. murpheyi, A. palmeri, A. parryi var. couesii, A. schottii, and A. toumeyana (Hodgson 1999a, Reveal & Hodgson 2002, Parker 2017).

Agave AGAVACEAE

Fig. 16 Agave chrysoglossa (Etter & Kristen 548: Mexico; Sonora, Nacapule Canyon, 280 m). (Copyright: J. Etter & M. Kristen)

A. chrysantha appears to have developed minor shifts in several floral characters that enhance diurnal pollination; bats were not recorded as important pollinators (Slauson 2000). A. chrysoglossa I. M. Johnston (Proc. Calif. Acad. Sci., ser. 4, 12: 998–999, 1924). Type: Mexico, Sonora (Johnston 3123 [CAS, ARIZ, GH, US]). — Lit: Gentry (1982: 71–75); Ullrich (1994a); Turner & al. (1995); Wilder & al. (2008); Felger & al. (2011); all with ills. Distr: Mexico (C & S Sonora: coastal and San Pedro Nolasco & Tiburón islands); slopes or often bare rocks in hot coastal and lowland regions, in desert and thorn scrub, 300–760 m; flowers mid-March to mid-May. I: Etter & Kristen (2001: 172–173); Richter (2011: 60); Pilbeam (2013: 58–59); Hochstätter (2015: VIII: 20). – Fig. 16. [1b] Stems short; Ros openly spreading, 1–1.3
2–2.4 m, mostly solitary, sometimes suckering

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profusely; L few, linear-lanceolate, straight or slightly curved, deflexed at maturity, convex below, flat above, smooth, 70–120
4–7 cm, widened at the base, light green, margins fragile, 1 mm wide, brown; marginal teeth none; terminal Sp acicular, with a short fine groove at the base above, 20–40 mm, brown, aging greyish; Inf mostly 2–4 m, ‘spicate’, peduncle with dry chartaceous Bra, lower Bra longer than the flowers, fertile part densely flowered, in the upper of the inflorescence; Ped bifurcate, 10–15 mm; Fl geminate, 35–45 mm; Tep yellow, tube shallow, 4–4.5
9 mm, lobes ovate, flat, outcurving at anthesis, clasping the filaments after anthesis, thin,  equal, outer 14–16
6 mm, keeled, the inner wider (8 mm) and unkeeled; Fil 40–45 mm, inserted on the rim of the tube at the base of the lobes; Anth 14–18 mm; Ov slender, scarcely grooved, 16–20 mm incl. a 3–5 mm long neck; Fr mostly oblong, 20
10 mm, shortly apiculate, parallel-veined and slightly cross-ridged; Se crescent-shaped, with 1 to several wrinkles on the faces, marginal wing low, firm, the hilar notch sharply angled, mostly 4–4.3
2.5–2.8 mm. Closely related to A. vilmoriniana, but distinguished by straight narrow flat leaves (vs. deeply guttered and arching in A. vilmoriniana) and without bulbils in the inflorescence. Gentry (1982) considers the inland populations at Río Yaqui as showing introgression from A. vilmoriniana; Turner & al. (1995) map these as A. chrysoglossa. The Sierra Seri populations of A. chrysoglossa appear to bridge a morphological gap to A. pelona (Turner & al. 1995). Specimens by R. Felger cited by Gentry (1982: 87) seem closer to A. pelona and are placed there by Turner & al. (1995). See also under A. ocahui. For the habitats on the Gulf Islands, see Wilder & al. (2008) and Felger & al. (2011). The seed production is estimated as 500,000–750,000 seeds per plant. The flowers are visited by numerous insects, hummingbirds, and other birds (Gentry 1982). A. cocui Trelease (Mem. Nation. Acad. Sci. 11: 19, tt. 5–7, 1913). Type: Venezuela (Zuloaga s.n. [MO [syn, 4 sheets]]). — Lit: Berger (1915: 224–225); Hummelinck (1936: 244–247, 249, tt. VII, VIIIa); Hummelinck (1938: 21–23, tt. Ib,

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IVb); Figueredo (2010: with ill.). Distr: Venezuela (incl. Isla Margarita), Colombia; mainly coastal, but also extending into inter-Andean valleys, 400–1000 m; flowers January to May. I: Hoyos F. (1985: fig. 83); Mostul (2005: 70); Figueredo (2010); Richter (2011: 96); Pilbeam (2013: 60); Hochstätter (2015: VII: 36). Incl. Agave americana Humboldt (1808) (nom. illeg., ICN Art. 53.1); incl. Agave cocui var. cucutensis Hummelinck (1936); incl. Agave cocui var. laguayrensis Hummelinck (1936). [2u] Acaulescent or sometimes subacaulescent; Ros to 2.5 m ∅, suckering; L rather many, lanceolate to elliptic, sometimes suboblong, rarely oblanceolate, guttered, often slightly S-curved, tip often slightly upcurved, sharply acute or subacuminate, (80–) 100–120 (–140)
20–30 cm, (3.5–) 4–5 (–6)
as long as broad, glaucous, soon green and glossy; marginal teeth mostly upcurved above and recurved below, acuminately triangular or from crescentshaped bases on green or later hardening prominences, (2–) 2.5–6 (7) mm, reddish chestnutbrown, (6–) 10–14 (–22) per 10 cm, intervening margin concave; terminal Sp conical or rarely acicular, sometimes triquetrous, rather often distinctly laterally flattened, very rarely flexuous, often recurved near the tip, shallowly grooved below the middle and involute below, smooth, sometimes rough, (10–) 12–22 (–30)
(2–) 3–5 (–6) mm, red-brown, slightly or not decurrent, dorsally very slightly immersed into the leaf tissue; Inf (5–) 6–8 (–10) m, paniculate, peduncle with 15–25 Bra, Bra serrate towards the tips, narrowly triangular, at length recurved, fertile part in the upper of the inflorescence, narrowly oblong, part-Inf on nearly horizontal branches, freely bulbilliferous; Ped (3–) 4–8 (–12) mm; Fl (40–) 45–55 (–65) mm; Tep yellow, tube openly conical, (3–) 4–6 (–7) mm, lobes (18–) 20–24 (–25)
5–7 mm; Fil (30–) 35–45 (–55) mm, inserted 1 mm above to 0–1 (–1.5) mm below the throat; Anth (15–) 18–23 (–25) mm; Ov 25–40
7–8 mm; Sty (38–) 44–50 (–55) mm; Fr oblong, (35–) 40–50
20–25 mm, usually very slightly beaked, shortly stipitate; Se 7–9
5–6 mm. — Cytology: 2n = 60 (Mata-Sucre & al. 2017). The type at MO consists of 4 sheets which are not cross-labelled and thus represent syntypes

J. Thiede

(Smith & Figueiredo 2014e: 231). Not always clearly separated from A. vicina (Hummelinck 1938: as A. vivipara). Hummelinck’s concept of A. cocui included plants from Curaçao and neighbouring islands here separated as A. vivipara (see there). The species is not restricted to coastal regions and extends into inter-Andean valleys as far as Táchira (Venezuela) and Santander (Colombia; Hummelinck (1938: 21)). A. cocui is an important crop in Venezuela, used from wild plants in traditional production systems esp. for liquor, soap, fibres, and preserves (Salazar & al. 2009, Figueredo 2010). Flowering starts mainly with the onset of the dry period in January or later and is highly synchronized among the populations (Figueredo & Nassar 2011). Molecular populational studies showed that A. cocui has one of the lowest estimates of genetic diversity as yet reported, which is esp. low in its centre of economic use, suggesting that human impact represents a significant threat to its gene pool (Figueredo & Nassar 2011), but 3 populations could be separated with AFLP data (Yépez & al. 2008). For the in vitro propagation, see Salazar & al. (2009). See also under A. boldinghiana. A. coetocapnia (M. Roemer) Govaerts & Thiede (Willdenowia 43(2): 331, 2013). Type: Mexico (Deppe s.n. [†; [lecto — icono]: Link & Otto, Icon. Pl. Rar., t. 18, 1828, as ‘Zetocapnia’]). — Lit: McVaugh (1989: 248–252); Solano & García-Mendoza (2013: with ills.); both as Polianthes geminiflora. Distr: N and C Mexico.  Bravoa coetocapnia M. Roemer (1847); incl. Bravoa geminiflora Lexarza (1824)  Polianthes geminiflora (Lexarza) Rose (1903); incl. Coetocapnia geminiflora Link & Otto (1828); incl. Robynsia geminiflora Drapiez (1841); incl. Bravoa graminiflora Hemsley (1884) (nom. inval., Art. 61.1); incl. Polianthes americana Sessé & Moçiño (1888); incl. Polianthes tubulata Sessé & Moçiño (1894); incl. Agave duplicata Thiede & Eggli (1999) (nom. illeg., ICN Art. 52.1). This is the most widespread and well-known species of Sect. Polianthes. It is rather variable in the size of the plant, leaves, bracts, inflorescences, and the number of flowering nodes (McVaugh 1989, Solano & García-Mendoza 2013).

Agave AGAVACEAE

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When transferring Polianthes geminiflora to Agave, Thiede & Eggli (1999) published the new name A. duplicata due to the earlier A. geminiflora (Tagliabue) Ker Gawler (1817), but were unaware of the prioritable epithet available as Bravoa coetocapnia (Govaerts & Thiede 2013). Solano & García-Mendoza (2013) provided a taxonomic treatment (as Polianthes geminiflora) and neotypification. The hybrid of this species with A. bulliana is known as A.
kewensis (see there). A. coetocapnia ssp. clivicola (McVaugh) Govaerts & Thiede (Willdenowia 43(2): 332, 2013). Type: Mexico, Jalisco (Wilbur & Wilbur 2133 [MICH, IEB]). — Lit: McVaugh (1989: 250); Solano & García-Mendoza (2013); CastroCastro & al. (2015: with ill.); all as Polianthes geminiflora var. Distr: Mexico (NW, SW, C & S Jalisco, W Michoacán); mostly on steep shaded slopes, barrancas, gullies, in oak, pine, and pine-oak forests, grasslands, or ecotones with tropical subdeciduous forests, (900–) 1200– 2560 m; flowers June to September (to October). I: Guillot Ortiz & al. (2006: 67); Castro-Castro & al. (2015); Hochstätter (2016: II: 12); CastroCastro & al. (2016: 721); all as Polianthes geminiflora var./ssp.  Polianthes geminiflora var. clivicola McVaugh (1989)  Agave duplicata ssp. clivicola (McVaugh) Thiede & Eggli (1999) (incorrect name, ICN Art. 11.4)  Polianthes geminiflora ssp. clivicola (McVaugh) Hochstätter (2016). [3b2] Differs from ssp. coetocapnia: L margins usually very narrowly revolute, with very thin pale scarious or hyaline edge, smooth and entire, rarely obscurely roughened. This is the W element within A. coetocapnia, differing in leaf margin characters. A. coetocapnia ssp. coetocapnia — Lit: McVaugh (1989: 248–252); Solano & GarcíaMendoza (2013: with ills.); Guillot Ortiz & Meer (2016a: 72–74); all as Polianthes geminiflora. Distr: Mexico (Durango, Nayarit, Jalisco, Guanajuato, Querétaro, Hidalgo, Michoacán, México, Tlaxcala, Distrito Federal, Morelos, Guerrero, Puebla); mainly on rocky slopes in oak, pine or cloud forests, mainly in the Trans-Mexican

Fig. 17 Agave coetocapnia ssp. coetocapnia (Etter & Kristen 2835: Mexico; Jalisco,  20 km S of San Cristóbal de la Barranca, 1470 m). (Copyright: J. Etter & M. Kristen)

Volcanic Belt, 900–2800 m; flowers June to August (to November). I: Guillot Ortiz & al. (2006: 67, as P. geminiflora); García-Mendoza & Esparza Alvarado (2010: 62–63, as P. geminiflora); Etter & Kristen (2012: 85, as A. duplicata); Castro-Castro & al. (2015: 145, as P. geminiflora var.); Hochstätter (2016: II: 11, as P. geminiflora var.); Castro-Castro & al. (2016: 721, as P. geminiflora var.). – Fig. 17.  Agave duplicata ssp. duplicata. [3b2] Herbaceous; corm cylindrical, 1.5–2.7
1–2 cm, bulb ovoid, 1.8–4
1.3–3 cm, covered with dry leaf bases; R contractile, fleshy and fibrous; L 2–6 (–8), linear-lanceolate to broadly attenuate towards the acute tip, (8–) 12–35 (–50)
0.1–2.5 (–3.7) cm, glaucous, plain green or spotted with small or large and confluent green or dark brown spots, lower face slightly papillose on the veins, margins entire, denticulate, papillose to erose; Inf 35–100 (–154) cm, ‘racemose’,

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peduncle with linear to lanceolate Bra 4.5–31
0.1–2.3 (–2.8) cm, fertile part lax, denser towards the tip, 10–40 (–60) cm; Ped strongly ascending, (3–) 10–18 mm; Fl geminate, but often only 1 developing, pendent, unscented; Tep red, orange or coral-red, tube slender and terete at the base, broadened above, 14–21
1–3 (–5) (at the tip) mm, throat straight or slightly oblique, lobes erect, ovate, (sub-) orbicular to transversally elliptic, subequal, 2–3 (–5)
1–2.5 mm, cucullate, apex obtuse to rounded, with a tuft of short white hairs; St included; Fil filiform, (5–) 10–21 (–24) mm, white, inserted 2–5 mm above the tip of the ovary; Anth linear, 3–6 mm, yellow; Ov erect, ellipsoid; Sty filiform, 10–25 mm, sometimes as long as the tepal tube, white; Sti 3-lobate; Fr shortly oblong or almost globose, 7–17
5–12 mm; Se semicircular, flat, 2–4 (–5)
2–3 mm, black, shiny or dull. Zamudio & Galván Villanueva (2011: 87) provide a new record for Guanajuato. The flowers attract hummingbirds and insects (Etter & Kristen 2012). A. coetocapnia ssp. pueblensis (E. Solano & García-Mendoza) Govaerts & Thiede (Willdenowia 43(2): 332, 2013). Type: Mexico, Puebla (Solano & al. 849 [MEXU, CHAPA, ENCB, FEZA, HUAP, HUMO, IEB, US]). — Lit: Solano & García-Mendoza (2013: as Polianthes geminiflora var.). Distr: Mexico (W & SW Puebla); open spots on calcareous soils, in oak and juniper forests, desert scrub, grasslands and corn fields, 2160–2220 m; flowers July to August. I: Hochstätter (2016: II: 13, as P. geminiflora ssp.).  Polianthes geminiflora var. pueblensis E. Solano & García-Mendoza (2007)  Agave duplicata ssp. pueblensis (E. Solano & GarcíaMendoza) Thiede (2012) (incorrect name, ICN Art. 11.4)  Polianthes geminiflora ssp. pueblensis (E. Solano & García-Mendoza) Hochstätter (2016). [3b2] Differs from ssp. coetocapnia: L very narrow, 1–2 mm broad, papillose on the lower face, margins regularly denticulate. This is the SE element within A. coetocapnia, differing in its leaf margin and very narrow leaves.

J. Thiede

A. colimana Gentry (Cact. Succ. J. (US) 40: 212–213, ills., 1968). Type: Mexico, Colima (Gentry 18325 [US, ARIZ, DES, MEXU]). — Lit: Gentry (1982: 102–107, with ills.); McVaugh (1989); Ullrich (1991a: with ills.); VázquezGarcía & al. (2007b: 49–50, t. G). Distr: Mexico (W, SW & S Jalisco, Colima, Michoacán); primarily coastal on rocky sites, or more inland in tropical deciduous forests, 0–1000 m; flowers November to March. I: Etter & Kristen (2012: 95); Pilbeam (2013: 61). Incl. Agave ortgiesiana Roezl (1871) (nom. inval., ICN Art. 36.1b?); incl. Agave schidigera var. ortgiesiana Baker (1877)  Agave ortgiesiana (Baker) Trelease (1914) (nom. illeg., ICN Art. 53.1)  Agave angustissima var. ortgiesiana (Baker) Trelease (1920); incl. Agave maritima Rose ex Hort. Kew (1897) (nom. inval., ICN Art. 38.1a); incl. Agave ortgiesiana var. brevifolia Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a). [1d] Shortly caulescent; Ros 0.4–0.6
1–1.2 m, solitary; L many, linear, straight, slightly narrowed above the base, widest near the middle, thin and flat above, smooth, 40–70
1–2.5 cm, green, with light green mid-stripe, margins narrow, brown, filiferous with fine, long, brown threads; marginal teeth none; terminal Sp weak, short, broadly rounded below, flattened or somewhat hollowed above, 5–8 mm, greyishbrown to dark brown, decurrent into the leaf margin; Inf 2–3 m, ‘spicate’, peduncle with linearacicular dark brown or purplish Bra, lower Bra longer than the flowers, the upper somewhat shorter, fertile part of the inflorescence slender, from 1 m above the base, part-Inf not crowded; Ped 10–15 mm; Fl geminate, 40–50 mm; Tep pale yellow or lavender, tube slightly 6-furrowed, narrow, 9–17
5–9 mm, lobes recurved at anthesis, subequal, 14–19
4–6 mm, outer 1 mm longer, thin, obtuse, with thickened papillate tip; Fil 30–40 mm, pale reddish, inserted 8–11 mm above the tube base and 1–2 mm below the lobe sinuses; Anth 17–21 mm, yellow; Ov round, 14–20 mm, greenish-yellow, neck 4–7 mm, slightly constricted; Sti contiguous with the style and not clavate, roundly trigonous, with papillae decurrent on the angles below the tip; Fr oblong,

Agave AGAVACEAE

20–25 mm, conspicuously constricted at the base, beaked, stipe stout, to 10 mm; Se small, hemispherical, rugulose, 2–3
3.5–4 mm. Typical plants are distinctive with their elongate leaves and deep narrow flower tube, but sometimes approach A. schidigera in flower tube length and A. multifilifera in leaf characters and are then hard to distinguish without complete material (Gentry 1982). There has been a long dispute about the correct name for this species: McVaugh (1989: 135) regards the earlier name A. ortgiesiana Roezl (1871) as invalid since it was published in an excerpt of a letter only. Ullrich (1991a), in contrast, accepts A. ortgiesiana Roezl as validly published and thus as the correct name for the taxon. Govaerts (WCSP, April 2014) does not cite the Roezl name, but instead accepts A. ortgiesiana (Baker) Trelease (1914). Here, the established name A. colimana is retained for the time being, pending a formal proposal to conserve it over the earlier A. ortgiesiana (Baker) Trelease. Tschapka & al. (2008) observed a highly specialized nectar-feeding bat on this species, Ceiba pentandra, Pachycereus pecten-aboriginum and Pseudobombax ellipticum; another species of migratory nectar-feeding bat was traced by Stoner & al. (2003). A. collina Greenman (Contr. Gray Herb. 11: 296, 1897). Type: Mexico, Morelos (Pringle 6349 [MO, AC, BR, CAS, CM, ENCB, F, G, GH, GOET, JE, K, MEXU, MICH, MIN, MSC, NDG, NY, P, PH, S, US, VT]). — Lit: Berger (1915: 253); Trelease (1920: 116); Gentry (1982: 586). Distr: Mexico (Morelos, Guerrero); 1500 m; flowers in June. [2g] Acaulescent, unknown whether Ros solitary or offsetting; L 30–40, linear-attenuate, gradually narrowing towards the apex, convex below, concave above, 60–80
5–8 cm (in the middle, narrowing to 2 cm just above the base), bluishgreen, somewhat green-banded below, margins narrowly cartilaginous; marginal teeth straight or upcurved, 3–5 mm, reddish-brown, on broad lenticular bases, 8–25 mm apart, intervening margin straight or very slightly emarginate, with a fine, brown or yellowish, horny line; terminal Sp conical, broadly and somewhat crookedly

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canaliculate, 20–30
3–5 mm, reddish- or purplish-brown, somewhat decurrent; Inf 3–4 m, paniculate, fertile part 60–80 cm, lower partInf 30 cm; Fl 60 mm (90 mm incl. the exserted stamens); Tep greenish-yellow, tube funnelshaped, 15–16 mm, lobes linear-oblong, obtuse, thickened at the apex, 19–20 mm; Fil flattened, glabrous, 50 mm; Anth 25–30 mm; Ov 25 mm, neck constricted; Sty slightly shorter or  as long as the filaments; Sti capitate; Fr 45–50
25–30 mm, shortly beaked; Se crescent-shaped to obliquely triangular, 8–10
6–8 mm. Listed by Gentry (1982: 586) under “incertae sedis”, but accepted as species by Espejo Serna & López Ferrari (1993), Govaerts (2014+), and Portal Datos Abiertos UNAM (2018+: accessed Jan. 2019). According to the protologue it is probably identical with A. serrulata (as “serratula”), a synonym of A. angustifolia. The measurements of the protologue fall well within the range of the variable A. angustifolia, and A. collina may belong to that complex as suggested by Gentry (1982: 586). Gentry deviates in part from the protologue, and describes the species as “a stout, short-stemmed plant with nearly white glaucous to pale green leaves with nearly black teeth and spine” and as “frequent in middle elevations from Morelos to Guerrero”. Fernández Nava & al. (1998: 2) list the species for the Balsas basin (Guerrero). A. colorata Gentry (Publ. Carnegie Inst. Washington 527: 93–94, t. 14: fig. 2, 1942). Type: Mexico, Sonora (Gentry 3050 [CAS, ARIZ]). — Lit: Gentry (1972: 117–119, ills.); Gentry (1982: 431–433, ills.); Ullrich (1993f: with ills.); Turner & al. (1995: 48–49); Starr (2012: 78–83, 320–321, ills.). Distr: Mexico (S Sonora, N Sinaloa); foothills or coastal regions, open rocky sites in thorn or pine-oak forests, on volcanic or white limestone rocks, 60–200 m; flowers March to June. I: Irish & Irish (2000: t. 13); Heller (2006: 76); Klopper & al. (2010); Richter (2011: 131); Pilbeam (2013: 62–63); Hochstätter (2015: IV: 7); Moore (2016: 271). – Fig. 18. [2h] Stems short; Ros compact, 0.45–0.9
0.45–1.2 m, tapered at the base or hemispherical, solitary or sparingly suckering; L few, ovate, shortly acuminate to lanceolate, thick, firm,

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J. Thiede

Fig. 18 Agave colorata (Etter & Kristen 562: Mexico; Sonora, near Aquihuiquichi, 200 m). (Copyright: J. Etter & M. Kristen)

convex below towards the base, flat to concave above, asperous, 25–60 (–100)
12–18 cm, light grey, glaucous, frequently cross-zoned and red-tinted, margins prominently crenate or with prominences; marginal teeth straight or flexuous, mostly 5–10 mm (in the middle of the lamina), brown to greyish, 15–30 mm apart, smaller below; terminal Sp subulate, straight or flexuous, narrowly grooved above in the lower , mostly 30–50 mm, brown to grey; Inf 2–3 m, paniculate, lower Bra leaf-like, the others soon drying, appressed at the base, reflexed above, 10–15 cm, reddish, fertile part of the inflorescence narrow, part-Inf densely flowered, 15–20 in the upper of the inflorescence; Fl 53–73 mm; Tep yellow, reddish in bud, apex usually remaining reddish, tube cylindrical, thick-walled, narrowly grooved, 15–20
12–14 mm, lobes linear, strictly erect, thickly fleshy, unequal, outer larger, overlapping, 12–16
5–7 mm, inner shorter, thickly keeled; Fil stout, flattened, tapered, 45–65 mm, inserted at 2 levels at mid-tube, yellow or pink; Anth 21–26 mm; Ov 25–40 mm, pale green, neck short, not constricted; Sty thick, sometimes pink, exceeding the stamens after anthesis; Sti capitate; Fr oblong to clavate, 45–55
15–17 mm, thickly and shortly stipitate or sessile; Se tear-shaped, 6
5 mm, hilar notch frequently obscure, marginal wing narrow. — Cytology: 2n = 60 (Simpson & al. 2011). The closest relative, both morphologically and geographically, is A. shrevei, from which

A. colorata differs in its broader cross-zoned leaves with deeply crenate to repand margins, more densely flowered panicles with closely set ‘umbels’, and its shorter, wider tepal tube and thicker ovary. The species is uncommon in nature. Its compact size and bright glaucous leaves with pinkish cross zones make it an attractive ornamental (Gentry 1982). Plants from near Guaymas and San Carlos Bay (Sonora) have longer and narrower leaves with smaller marginal teeth (Gentry 1982: 432–433, Starr 2012); these were first misplaced in A. fortiflora (Gentry 1972: 125). Bats and possibly also hummingbirds are the assumed pollinators (Gentry 1982). Flowers pollinated by the likely main pollinator, the bat Leptonycteris yerbabuenae, show much higher fruit set, higher numbers of viable seeds, and a higher germination rate compared to fruits and seeds obtained from self-pollination (Borbón-Palomares & al. 2018). A. confertiflora Thiede & Eggli (Kakt. and. Sukk. 50(5): 111, 1999). Type: Mexico, Chihuahua (Hartman 536 [US, MEXU, UC]). — Lit: Rose (1899: 155, t. 18, as Pseudobravoa densiflora); García-Mendoza (2003); Solano & Feria (2007); Feria-Arroyo & al. (2010: with ill.); all as Polianthes densiflora. Distr: Mexico (SW Chihuahua); open sites on calcareous soils, in pine or pine-oak forests, 2190–2500 m. I: Hochstätter (2016: II: 9).

Agave AGAVACEAE

 Bravoa densiflora B. L. Robinson & Fernald (1895)  Pseudobravoa densiflora (B. L. Robinson & Fernald) Rose (1899)  Polianthes densiflora (B. L. Robinson & Fernald) Shinners (1966). [3b1] Herbaceous, small, 7–15 cm; corm cylindrical, to 1
1 cm, bulb oblong, to 1
1.5 cm, covered with dry leaf bases; R many, contractile, thickened; L 4–10, linear, attenuate, apex acute, 7.5–11
0.2–0.3 cm, green, margins entire, or partly papillose; Inf ‘spicate’, short; peduncle with Bra 2.5–5 cm, with broad scarious and attenuate tips; fertile part dense, with (2–) 4–6 floral nodes, 7.5–13 cm; Ped none; Fl solitary, slenderly tubular, spreading, curved, 43–55 mm; Tep pulverulent-tomentose on the outer face, dull yellow, tube narrow, scarcely widened, 21–44
3–4 mm, throat oblique, lobes erect, ovate, obtuse, equal, 2.5–4 mm, tip with a tuft of short white hairs; St included; Fil filiform, inserted in the tube; Anth linear, 5–10
1–2 mm, yellow to green-yellowish, at anthesis surrounding the throat; Ov cylindrical; Sty filiform, as long as the tube, included at anthesis; Sti 3-lobate; Fr globose, 8–12
7–10 mm; Se deltoid or semicircular, flat, 3–4
2.4–3.3 mm, shiny or dull. A hardly known but seemingly very distinct species. Long known from the type collection only, but Solano & Feria (2007: 1888) mention 13 records from 5 localities. The geographical distribution, niche modelling and extinction risk of the species are dealt with by Solano & Feria (2007) and Feria-Arroyo & al. (2010). The type locality is incorrectly cited as “Varogachic” in the protologue, but is Norogachi, a Tarahumara community in the town of Guachochi (database note for GH 00030302!), and the type is “Hartman, 5 July, 1892 (no. 536)” (US, MEXU, UC) and not Hartman 546, 6 July 1892 (GH, K, MEXU), partly labelled as “Type”. — When Polianthes densiflora is transferred to Agave, a new name is necessary to avoid homonymy with A. densiflora Hooker. A. congesta Gentry (Agaves Cont. North Amer., 476–479, ills., 1982). Type: Mexico, Chiapas (Gentry 23651 [US [2 sheets], ARIZ, DES, MEXU, MICH]). — Lit: Lott & García-Mendoza

91

(1994: online version with ills.); García-Mendoza (2003: with ill.). Distr: Mexico (Chiapas); widely scattered in pine-oak forests and pine woodlands in the Montañas del Oriente, on calcareous rocks, (930–) 2100–2500 m; flowers December to February. I: García-Mendoza (2002: 187); Richter (2011: 114); Pilbeam (2013: 64); Hochstätter (2015: VI: 6). [2o] Stems short; Ros compact, 0.5–1
1–2 m, solitary; L many, lanceolate to lanceolate-spatulate, at first curved-ascending, then horizontally spreading, thick, narrowed at the base, (shortly) acuminate, flat, (40–) 50–120
10–22 cm, green to yellow-green, sometimes faintly glaucous or pruinose, margins straight in the lower , repand to crenate in the upper ; marginal teeth straight to variously curved, moderate to rather large, dark to greyish-brown, usually remote, 20–50 mm apart, mostly on variously shaped prominences, 5–10 mm, base broad and low; terminal Sp stout, base very broad, widely flatly grooved above, 30–70 mm, grey to chestnut-brown, sharply decurrent to the upper teeth; Inf 6–8 m, paniculate (“racemose” in the protologue), peduncle with prominent Bra, fertile part straight, slender, with 40–50 part-Inf as congested rounded clusters, subsessile or on short lateral branches 2–10 cm long, in the upper or more of the inflorescence; Ped closely subtended by small purplish bracts, 3–10 (–20) mm; Fl (45–) 55–70 mm; Tep orange to reddish or purplish in bud, yellow at anthesis, softly fleshy, tube deeply funnel-shaped to tubular, thickwalled, grooved, knobby, 10–13 mm, lobes linear-lanceolate to narrowly triangular, involute, thickly rounded-cucullate, unequal, 17–30 mm, the outer larger, dark-coloured apically, the inner shorter, with a thick prominent keel; Fil slender, 40–60 mm, purplish, inserted at 2 levels near the throat of the tube; Anth centric, irregularly curved to sinuous, 20–30 mm, bronze-coloured; Ov angular-fusiform, (25–) 30–40 mm, neck short; Sty stout, lengthening after anthesis, somewhat longer than the filaments; Sti capitate; Fr oblong, 50
20 mm; Se unknown. The species name alludes to the congested flowers in the dense globose ‘umbels’. It is further distinguished by the dimorphic dark-coloured

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tepals and the filaments inserted at 2 levels. The closely related A. hiemiflora has the same but less pronounced characteristics, and is distinguished by smaller rosettes with fewer leaves, and paler tepals (Gentry 1982: 479). Gentry’s “holotype” at US consists of 2 sheets which appear to have been cross-labelled later. A. convallis Trelease (Contr. US Nation. Herb. 23: 138, 1920). Type [lecto]: Mexico, Oaxaca (Trelease 4 [MO [1128755 + 1256267]]). — Lit: García-Mendoza (2010); García-Mendoza (2011a: 23–26); León Vázquez & al. (2013). Distr: Mexico (S Puebla, NW & C Oaxaca); desert scrub and pine forests, in soils of volcanic origin, 1610–2640 m; flowers August to January. Incl. Agave dissimulans Trelease (1920). [1g] Acaulescent; Ros dense, 1–1.2
1.5–2 m, solitary or rarely caespitose; L (40–) 50–80, broadly lanceolate to lanceolate, erect, stiff, succulent, leathery, fibrous, thick and flat at the base, flat to slightly guttered towards the apex, (50–) 70–100
10–16 (in the middle) cm, dark green, yellow-green or purple, sometimes with a weak yellowish mid-stripe, margins straight, horny, continuous, white; marginal teeth broadly triangular, sometimes bifid, present towards the base, absent in the uppermost 5–15 (–35) cm, straight to variously curved, 9–15 (–20)
(4–) 7–10 mm, (10–) 25–45 (–70) mm apart, grey-white or brown, sometimes with a lenticular broadening beneath each tooth, sometimes with small intermittent teeth; terminal Sp conical, deeply channelled above, 30–55
5–8 mm, greyishwhite, decurrent on the lower face for 4–15 mm; Inf 3–6 (–8) m, ‘spicate’, peduncle green, with linear chartaceous deciduous Bra 11–15 cm long, fertile part in the upper ; Ped 3–4 (–10) mm; Fl geminate, campanulate, (34–) 38–43 mm; Tep yellowish-green outside, dark reddish inside, tube 4–6
4–6 (–10) mm, lobes oblong, 17–20
4–6 (–10) mm, inner ones keeled, outer ones larger; Fil (30–) 35–40 mm, red, inserted in the mouth of the tube; Anth 15–20 mm, reddish; Ov cylindrical, 16–20
4–6 mm, neck 2–4 mm, constricted; Sty 40–50 mm, yellowish; Fr obovoid, 24–25
12–16 mm; Se 5–6.5
3–4 mm, with inconspicuous wing, black.

J. Thiede

Previously placed in the synonymy of A. kerchovei (Gentry 1982: 149), but recently re-established as a species of its own (GarcíaMendoza 2010, García-Mendoza 2011a, Govaerts 2014+). The species differs from A. kerchovei in its larger number of shorter, relatively broader, broadly lanceolate to lanceolate leaves per rosette, the lack of teeth in the uppermost 5–15 (–35) cm of the margins, its more closely placed teeth, its oblong tepals, and its obovoid capsules and larger seeds. Furthermore, A. convallis occurs at higher altitudes mainly on igneous soils (García-Mendoza 2011a). This author also provides a new record for Puebla. Smith & al. (2018b) designated the lectotype cited above. A. cundinamarcensis A. Berger (Agaven, 222, 1915). Type: Colombia, Cundinamarca (Wercklé s.n. [not indicated]). — Lit: Bernal & al. (2015: 794). Distr: Colombia (Cundinamarca: Río Cuja valley S of Bogotá); 1250 m. I: Hochstätter (2015: VII: 59). [2v] Ros solitary, not suckering; L very thick above the base but only 15 cm broad, straightly spreading, then curved upwards and rapidly becoming broader, above the middle 45 cm broad and then again curved outwards, but the last 15 cm again curved upwards, 200
45 cm, lead-grey on yellowish-green base colour; marginal teeth flat and broad, nearly blunt, very short, hardly pungent; terminal Sp rather short; Inf paniculate, sparingly bulbilliferous, not further described; Fl, Fr and Se not described. Possibly first mentioned by Wercklé (1907: 122) as Agave “vom westlichen Cundinamarca”, and still insufficiently known. Plants studied by J. Etter and M. Kristen in March 2019 near the type locality of A. cundinamarcensis largely match the protologue and appear to represent the true species. This and the second species from Colombia, A. wallisii, were both placed in his Reihe Columbianae by Berger (1915: 222), which he regarded as possibly closest to the Caribaeae. Neither of these 2 species appears close to the geographically adjacent ColombianVenezuelan A. cocui from Sect. Viviparae. Xhonneux (2004), Greulich (2012b), and Hochstätter (2015: VII: 59) show plants placed here from habitats

Agave AGAVACEAE

93

Fig. 19 Agave cupreata (Martinez & Eggli 188b: Mexico; Guerrero, between Mazatlán and Chilpancingo, 1350 m). (Copyright: U. Eggli)

W and N of Bogotá and variegated plants with broad yellow margins cultivated at Bogotá and in Europe, which differ clearly from the protologue in having smaller, less curved, green leaves much less broadened in the middle. The variegated plants were given the cultivar name ‘Condor’. According to P. van der Meer (pers. comm. Nov. 2018/Feb. 2019), these plants, including the variegated ones, are misnamed and belong to an undescribed species provisionally named “A. bogotensis”. The variegated plant may be the same as the medium-sized plant with dark green leaves with broad yellow margins cultivated at Bogotá reported earlier by Wercklé (Berger 1915: 222). Wercklé (in Berger 1915) and P. van der Meer (pers. comm. Feb. 2019) both report further undescribed species from Colombia. A. cupreata Trelease & A. Berger (Agaven, 197, 1915). Type [lecto]: Mexico, Michoacán/ Guerrero (Langlassé 867 [B, GH, K, MEXU, MPU, P, US]). — Lit: Gentry (1982: 335–337, ills.). Distr: Mexico (E Michoacán, Guerrero); mountain slopes, dry hills and rocky ledges, in tropical dry and oak forests and grasslands, 1220–2100 m; flowers November to March. I: García-Mendoza (2002: 183); Smith (2002: 225); Heller (2006: 77); Etter & Kristen (2007b: 178); Richter (2011: 48, 73); Pilbeam (2013: 65); Greulich (2014a: 56). – Fig. 19.

[2d] Caulescent, Ros openly spreading, 0.8–1.6 m ∅, solitary; L broadly lanceolate or ovate, thickly fleshy, strongly narrowed at the base, flat to slightly concave above, 40–80
18–20 cm, bright shiny green, with conspicuous patterns from the central bud, margins deeply crenate-mamillate; marginal teeth straight to curved, strongly flattened, dimorphic, larger teeth 10–15 mm on distinct prominences, 30–60 mm apart, with smaller intermittent teeth of varying sizes, copper-coloured to grey; terminal Sp slender, sinuous, openly grooved above, 30–50 mm, light brown to greyish, with a sharp border decurrent to the upper teeth; Inf 4–7 m, paniculate, fertile part rather broad, part-Inf lax diffuse ‘umbels’, 14–25 in the upper of the inflorescence, branches with dark small bracts; Fl 55–60 mm; Tep brown-red in bud, at anthesis orange-yellow, tube broadly funnel-shaped, 6–7
14–15 mm, lobes erect, linear-lanceolate, acute, subequal, outer 20–21 mm, apex rust-coloured, broader and thicker than the inner, inner with narrow keel and thin involute margins; Fil thickened towards the base, 35–40 mm, inserted at mid-tube; Anth excentric, curved, 23–24 mm, yellow; Ov fusiform, knobby, grooved, thickwalled, 30–35 mm, olive-green, neck constricted, doubly 3-grooved; Sty hardly longer than the tepals; Sti capitate; Fr and Se unknown. — Cytology: 2n = 60, 120, 150, 180 (Palomino & al. 2012, Gomez-Rodriguez & al. 2013).

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A beautiful plant from the Río Balsas basin, distinguished by the broad, shiny green leaves with tall marginal prominences and conspicuous patterns from the central bud, and brightly coppercoloured marginal teeth (Gentry 1982: 335). Material of the type collection is present in several herbaria, and Gentry (1982: 352) lists the specimen at B as “holotype”, which is interpreted as unintentional lectotypification. See also under A. potatorum. A morphological characterization of populations in Guerrero is provided by Avendaño-Arrazate & al. (2015), and a study of wild populations in Michoacán by Martínez Castro & al. (2015). A. cupreata regenerates successfully by seed in different habitats, but the population density is reduced by cattle trampling and grazing on seedlings and floral stalks (Martin & al. 2011). Martínez-Palacios & al. (2011) studied the genetic diversity and its implications for conservation. The continued extraction of reproductive individuals from wild populations does not produce a decline in genetic diversity in 5 studied populations (Aguirre-Dugua & Eguiarte 2013). The species (vernacular name “Agave papalote”) is locally semi-domesticated with only sexual reproduction, and used for mescal production (Illsley & al. 2007). Molecular populational data do not clearly separate A. cupreata from A. potatorum, and both are seen to represent one species only with two lineages: One in lower areas of the Balsas basin (A. cupreata), and another one in the highest forests and open areas (A. potatorum) (Eguiarte & al. 2013: 495–496). A. dasylirioides Jacobi & C. D. Bouché (Hamburg. Gart.- & Blumenzeit. 21: 344, 1865). Type [neo]: Cult. BG Berlin (Anonymous s.n. [B]). — Lit: Baker (1877b: 556–557, with ill.); Berger (1915: 83–85); Standley & Steyermark (1952: 110–111); Gentry (1982: 237–240, with ills.); García-Mendoza (2003: with ill.); Thiede (2014a); Thiede (2017a). Distr: Mexico (Morelos, México); igneous rock cliffs on mountain slopes, in mixed pine and hardwood forests, 1500–2200 m; flowers in December. I: Curtis’s Bot. Mag. 94: t. 5716, 1868; Kemble (2002); Heller (2006: 78); Richter (2011:

J. Thiede

36, 58); Pilbeam (2013: 66–67); Hochstätter (2015: VIII: 6). Incl. Agave dealbata E. Morren ex K. Koch (1862); incl. Agave dealbata Lemaire ex Jacobi (1865) (nom. illeg., ICN Art. 53.1)  Agave dasylirioides var. dealbata (Lemaire ex Jacobi) A. Terracciano (1885); incl. Agave dealbata var. nana K. Koch (1869) (nom. inval., ICN Art. 35.1); incl. Agave dealbata [?] angustifolia Hort. (1871) (nom. inval., ICN Art. 38.1a); incl. Agave dealbata [?] compacta Hort. (1871) (nom. inval., ICN Art. 38.1a); incl. Agave dealbata [?] compacta angustifolia Hort. (1871) (nom. inval., ICN Art. 24.2, 38.1a); incl. Agave intrepida Greenman (1899). [1a] Ros symmetrical, 0.3–0.5
0.6–1 m, generally solitary; L 70–100, linear-lanceolate, straightly spreading, pliable, relatively thin, scarcely succulent, flat above, mostly 40–60
2–3 cm, glaucous-green, smoothly striate below and above, margins 1 mm wide, pale yellowishwhite, minutely serrulate; marginal teeth none; terminal Sp acicular, rounded below, flat near the base above, 5–15 mm, reddish-brown; Inf 1.5–2 m, ‘spicate’, arching, peduncle greenish to pale reddish, with conspicuous greenish to pale reddish persistent Bra 10–15 cm long, fertile part in the upper of the inflorescence, part-Inf with 1–4 flowers; Fl 26–35 mm, persistent after anthesis; Tep pale green to greenish-yellow, tube funnel-shaped, 8–12
8–12 mm, shallowly grooved below the lobe sinuses, lobes spreading, ovate to oblong, mucronate, equal, 9–11
6–7 mm, sometimes with dark midstripe, the outer flat, the inner with a broad keel and convergent ribs within; Fil 35–50 mm, whitish-green to pink, inserted at mid-tube; Anth centrally affixed, 12–15 mm, yellow to brownish, persistent; Ov linear-tapered, 3-angulate, truncate at the base, 9–12 mm, neckless intruding the tepal tube; Sty whitish, sometimes pale reddish apically; Fr ovoid, 20
10 mm, without beak, pale grey to brown; Se 4
3 mm. The name A. dealbata is commonly ascribed to Jacobi (who used it in the same publication where A. dasylirioides was described 1865), but it was in fact first used by K. Koch in 1862. Because of priority, it would displace the universally used

Agave AGAVACEAE

name A. dasylirioides, and a conservation proposal is envisaged to avoid this (Thiede 2014a). A. dealbata var. nana is a long-forgotten name for a small-growing form. The species is best known from Morelos esp. around Cuernavaca, but was recently also collected in the State of México (GarcíaMendoza 2003, Thiede 2014a). Gentry (1982: 249) cited one collection from San Luis Potosí (Rzedowski 7128a), but this most probably belongs to A. gracielae (Galván & Zamudio 2013). Plants in habitat near Yosondua in Oaxaca depicted as “A. dasylirioides” by Ullrich (1990f) and Polka (2006) and a cultivated flowering plant depicted by Brand (2012) belong to the close relative A. kavandivi (see there for differences). See also the comment for A. petrophila. The type of A. dasylirioides was erroneously said to have been collected in Guatemala. Gentry (1982: 241) assumed this species to hold a very basal position in the genus based on its ‘primitive’ (plesiomorphic) features (leaves serrulate and scarcely succulent, inflorescences relatively simple, ovary incompletely inferior, tepals all equal, lobes nearly of equal length), and this assumption is principally confirmed by molecular data insofar as Sect. Juncineae (as Group Striatae) represents the sister group to the remainder of the genus (Bogler & Simpson 1996, Bogler & al. 2006, Gil-Vega & al. 2007; see also introduction to the genus). A. datylio Simon ex F. A. C. Weber (Bull. Mus. Hist. Nat. (Paris) 8: 224, 1902). Type [neo]: Mexico, Baja California (Gentry & Arguelles 11200 [US, ARIZ, MEXU, MICH]). — Lit: Trelease (1912b: 45, 61–62, tt. 68–71, also as A. vexans); Berger (1915: 259–260, also as A. vexans); Gentry (1978: 97–99, also as var. vexans); Gentry (1982: 571–572, also as var. vexans); Turner & al. (1995: 49–50, also as var. vexans); Webb & Starr (2015: 90–91); all with ills. except Berger. Distr: Mexico (C & S Baja California Sur); sandy plains and rocky granitic slopes to 730 m; flowers September to December. I: Goldman (1916: t. 111C, as A. vexans); Richter (2011); Pilbeam (2013: 68–69); Hochstätter (2015: VII: 37–38, also as ssp. vexans).

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Incl. Agave vexans Trelease (1912)  Agave datylio var. vexans (Trelease) I. M. Johnston (1924)  Agave datylio ssp. vexans (Trelease) Hochstätter (2015). [2g] Acaulescent; Ros 0.3–1
0.6–1.5 m, suckering freely, rhizomes frequently elongate; L radiately spreading, lanceolate-linear, nearly straight, rather rigid, rounded below, guttered above, 20–80
3–4 cm, green to yellow-green or glaucous-green, young somewhat glaucous, margins nearly straight; marginal teeth deltoid, mostly recurved, triangular, flattened, rather blunt, mostly 2–5 mm, dark brown, 15–60 mm apart, more closely spaced below, often on broad or lenticular bases; terminal Sp conical to subulate, scarcely or flatly grooved above, 20–40
3–6 mm, dark brown to greyish, shortly decurrent; Inf 1.2–5 m, paniculate, part-Inf small ‘umbels’, 8–15 in the upper of the inflorescence; Ped < 5 mm; Fl 40–55 mm; Tep greenish-yellow, tube funnel-shaped, 5–15
3–4 mm, lobes erect to ascending, 12–20
3–4 mm; Fil 25–45 mm, inserted 4–6 mm above the tube base; Anth 15–30 mm, yellow; Ov fusiform or flask-shaped, 20–30 mm, glaucous; Sty shorter than the filaments; Sti capitate; Fr (broadly) oblong to pearshaped, 35–45
15–20 mm, shortly stipitate, conspicuously beaked; Se 6–7
5–6 mm. — Cytology: 2n = 87, 90, 174 (3
, 6
; Cave (1964), Simpson & al. (2011: as A. vexans)). Widely scattered in the Cape region of Baja California Sur. This is the only species of Sect. Rigidae (as Group) in Baja California, and is without close relatives (Gentry 1982). It belongs to the “sword-leaved group” within Sect. Rigidae whose geographically nearest member is A. aktites from the opposite Sonoran-Sinaloan coast. A. aktites differs clearly by narrower bluish leaves with sharply cuspid teeth, and larger flowers with a deep bulging tube (Gentry 1982). Var. vexans was differentiated by Gentry (1978) and Berger (1915) by its smaller rosettes and leaves, but Webb & Starr (2015) consider the size differences as resulting from edaphic conditions and refer it to the synonymy. The triploid chromosome number of the former var. vexans suggests that parts of it may have arisen as hybrid between a di- and a tetraploid parent.

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A. debilis A. Berger (Agaven, 33, 1915). Type [lecto]: Mexico, Oaxaca (Pringle 4745 [US, BM, BR, CM, ENCB, G, GH, JE, LE, M, MEXU, MSC, NDG, P, S, VT]). — Lit: Rose (1903c: 19); Verhoek-Williams (1975: 248–254); Piña Luján (1985: 86–89); McVaugh (1989: 233); Castillejos-Cruz (2009: [238]–[247], with ills.); García-Mendoza (2011a: 79–79, 81); all as Manfreda pringlei. Distr: Mexico (Nayarit, Jalisco, Hidalgo, Michoacán, México, D.F., Tlaxcala, Morelos, Guerrero, Puebla, Veracruz, Oaxaca); moist forests or pine-oak ericaceous woods, in cloud, pine-oak, pine, oak, thorn and tropical deciduous forests, desert scrub and grasslands, (1000–) 1500–3000 m; flowers mid-July to November. I: Hochstätter (2016: I: 37, as Manfreda pringlei).  Manfreda pringlei Rose (1903)  Polianthes debilis (A. Berger) Shinners (1966); incl. Manfreda angustifolia Rose in sched. (s.a.) (nom. inval., ICN Art. 29.1). [3a3] Herbaceous, of moderate size (for Subgen. Manfreda), reproducing vegetatively by stoloniferous horizontal rhizomes with plantlets at the tips; corm 3–5
(1–) 1.5–2.5 cm, bulb cylindrical to ovoid, 3.5–7.5
1.8–3.2 (–6) cm, covered with membranous dry leaf bases fraying into fine fibres at the tip, (4–) 6–11 cm long; R halffleshy, fibrous, vertical; L drought-deciduous, (2–) 4–10, erect-spreading, linear to linearlanceolate, narrowed towards the base, somewhat succulent, slightly channelled, occasionally gently undulate, tip acute to acuminate, with mediumsized point, smooth, (12–) 24–45 (–80)
(0.5–) 1–2.7 cm, dark brown to shining green, sometimes with purple dots, at times red-speckled on the lower face near the base, without prominent veins, margins narrow, hyaline, sometimes streaked with purplish-red, papillate to erosedenticulate, usually rough to the touch; Inf 97–200 cm, ‘spicate’, peduncle greenish with purple tinge, peduncular Bra 6–12, similar to the leaves, fertile part (10–) 25–33 cm; Fl (5–) 10–25, sessile, diffuse to ascending, tubular, congested, nearly erect, 30–40 mm, slightly curved at the junction of ovary and tepal tube; Tep tube cylindrical, slightly widened towards the throat, tip cucullate, with a tuft of short hairs, 9–21
(3–) 5–6 (in the middle) mm, lobes oblong, revolute,

J. Thiede

8–16
2–4 (–7) mm, green-yellowish to reddish with purple tinge; Fil 26–45 (–54) mm, exceeding the tube for 22–42 (–50) mm, inserted slightly unequally in the middle of the tube, greenishyellow to greenish-red; Anth 11–14 mm, reddish to yellowish-green; Ov cylindrical, (8–) 9–11 (–12)
3–5 mm, not constricted, apex prolonged into the tepal tube for 1 mm; Sty 40–60 mm, exserted for 30–48 mm; Sti clavate, trigonous, shallowly furrowed, yellowish-green with small purple dots; Fr ellipsoid to subcylindrical, 15–20
12–17 mm; Se deltoid, plane-concave, with narrow margin, 3–4
4–5 mm. This species shares many floral and fruit characters with A. guttata, but is nevertheless easily distinguished by its longer, narrower and more pliable and herbaceous leaves and its distribution. It may represent the moist-forest counterpart of the more xeromorphic A. guttata (VerhoekWilliams 1975: 252–253, as Manfreda). A. debilis is morphologically also similar to the recently published A. verhoekiae (for differences see there). A. decipiens Baker (Bull. Misc. Inform. Kew 1892: 183, 1892). Type [lecto]: USA, Florida (Dodge s.n. [K, BUS, CICY?, US]). — Lit: Berger (1915: 242–243, with ill.); Gentry (1982: 573–574, with ills.); Zona (2002: with ills.); Reveal & Hodgson (2002); Franck (2012: 6–7, with ills.); Rubal Lobo & al. (2013: with ills.). Distr: USA (S Florida); coastal sands, sandy soil in hammocks, at sea-level; flowers November/ December to March. I: Curtis’s Bot. Mag. 122: t. 7477, 1896, as A. laxifolia; Heller (2006: 79, 80); Richter (2011: 128); Pilbeam (2013: 70); Hochstätter (2015: VII: 39); Moore (2016: 263). Incl. Agave laxifolia Baker (1896); incl. Agave spiralis Brandegee ex A. Berger (1912). [2g] Arborescent, trunk 1–3 (–4) m, very broad through bulging leaf bases, frequently suckering; Ros extending down for some distance from the stem tip, 1–1.5
1.5–2 m, not caespitose; L 36–72, linear-lanceolate, rigidly spreading to recurving, fleshy, narrowed at the thickened base, long-acuminate, convex below towards the base, concave above towards the apex, (24–) 70–100 (–200)
(1.3–) 7–10 cm, bright green or yellowish-green, without imprints from the central

Agave AGAVACEAE

bud, margins slightly wavy, repand; marginal teeth 2–3.6 mm at mid-leaf, curved upwards or downwards, dark brown, 10–20 mm apart, on low prominences, the slender tips upcurving, often with some smaller intermittent teeth, intervening margins not continuously horny; terminal Sp conical, without groove, 10–29 mm, brown or brownish-black, decurrent or not; Inf 3–5 m, paniculate, peduncle with caducous Bra; partInf ascending, 8–18 in the upper of the inflorescence, with 15–23 flowers, to 45 cm, often bulbilliferous; Ped 5–11 mm; Fl erect, 60–80 mm, foetid-scented; Tep greenish-yellow, tube funnelshaped, 6–13
4–6 mm, lobes incurved, tips cucullate, with a tuft of short white hairs, subequal, 17–26 mm, greenish-yellow, outer linear, drying reflexed on the tube; St long-exserted; Fil erect, terete, 27–47 mm, yellow, inserted at 2 levels at or slightly above mid-tube; Anth 13–25 mm, yellow; Ov large and thick, 22–43 mm, neckless; Fr ellipsoid to oblong, 35–50 mm, apex beaked, shortly stipitate; Se unknown. — Cytology: 2n = 120, 150, 160, 180, 190 (Banerjee & Sharma 1989, Reveal & Hodgson 2002, Simpson & al. 2011). Geographically isolated from the remainder of the genus (except A. neglecta). The taxon is most probably of cultivated origin. It was reported to occur at old Indian village sites in 1933 and may well represent an old pre-Columbian food or fibre plant, comparable to A. delamateri. The different chromosome numbers suggest a prolonged human propagation and probable hybrid origin (Reveal & Hodgson 2002). A. decipiens is possibly derived from A. angustifolia, from which it mainly differs in its obvious stem 1.5 m long or more (Gentry 1982, Zona 2002), but individuals of the latter with distinct stems are found on the Yucatán peninsula (Cruz-Ramos & al. 1985, Zona 2002). Plants from the Yucatán peninsula placed here belong to A. angustifolia (Gentry 1982, Zona 2002: as A. vivipara), Further differences from A. angustifolia are the longer ovaries and shorter anthers of A. decipiens (Zona 2002). Gentry’s “lectotypification” (1982: 573) represents a neotypification and is superfluous, since the type material is extant (Zona 2002). Franck (2012) designated a lectotype at K from the 3 extant specimens of the type collection. Rubal Lobo &

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al. (2013) and López-Pujol & al. (2015b): report the species as naturalized in Prov. Cádiz, Spain. The report of the species as naturalized in S Africa (Smith & Steyn 1999) actually refers to A. angustifolia (Smith & al. 2008: as A. vivipara). Introduced ants exploit the nectar of plants cultivated in Florida (Koptur & Truong 1998). A. delamateri W. C. Hodgson & Slauson (Haseltonia 3: 130–140, ills., 1995). Type: USA, Arizona (Hodgson 5478 [DES, ASU]). — Lit: Hodgson (1999a: 5, 14, 19, with ills.); Reveal & Hodgson (2002); Parker & al. (2007); Parker (2018a: 22, 24, with ills.). Distr: USA (C Arizona); gravelly places in desert scrub, rarely in chaparral or pinyon-juniper woodlands, 725–1555 m; flowers late June through July. I: Pilbeam (2013: 71). Incl. Agave repanda Trelease ex Gentry (1982) (nom. inval., ICN Art. 36.1c). [2h] Acaulescent; Ros 0.9–1
0.9–1 m, solitary to caespitose, freely suckering; L lanceolate or oblanceolate, rigid, erect or erect-ascending, broadest near or just below the middle, acuminate, guttered, apex conspicuously incurved, mostly 50–63 (–74)
7.5–9 cm, bluish-grey-glaucous with purple-maroon tinge and green cross-banding, margins straight or repand; marginal teeth variable, usually reflexed, becoming porrect near the leaf base, 3.5–5 mm, 10–30 mm apart, intermittent teeth 1–1.5 mm, (3–) 6–12 mm apart, mostly along the distal of the leaf; terminal Sp slender, 28–35 (–49) mm, brownish-grey, decurrent for of the leaf length; Inf 4.5–6 m, broadly paniculate, open, peduncle with persistent triangular Bra 1–5.5 cm long, part-Inf widely spaced, horizontal, 12–17 in the upper of the inflorescence, >10 cm, with 14–20 flowers each; Fl erect, long-lived, 47–67 (–70) mm; Tep pale cream tinged light green, tube campanulate, 11–16
11–16 mm, lobes spreading, unequal, 9–18 mm, apex tinged with maroon, persistent and often leathery during and after anthesis; St long-exserted; Fil erect, 30–53 mm, yellow, apex tinged with maroon, inserted at a single level  at mid-tube; Anth 11–20 mm, yellow; Ov 21–29 mm, neck slightly constricted, 1–3.5 mm; Sty creamcoloured, maroon-flecked, becoming more

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maroon towards the stigma; Sti capitate, 3-lobed, cream-coloured tinged with maroon; Fr aborting, never fully developed; Se inexistent. According to the protologue already collected around 1920 by S. D. McKelvey and recognized as distinct by Trelease, who in 1929 used the unpublished name A. repanda. It is most closely related to the allopatric A. fortiflora and A. palmeri, but distinguished esp. by its numerous rhizomatous offsets, easily cut leaves, and 1(instead of 2-) seriate filaments. A. delamateri was first known only from about 90 sites, always in association with Mogollon or Salado settlement features, and probably represents a cultivar derived from A. palmeri or a closely related taxon by pre-Columbian people. Later field studies yielded more than 200 sites so that A. delamateri is the most common one among the pre-Columbian cultivars from Arizona (Parker 2018a). It hybridizes with A. chrysantha (Hodgson 1999a, Reveal & Hodgson 2002). Parker & al. (2007) found a low level of genetic diversity, relating to its continued vegetative propagation from few clones only (see also under A. murpheyi). A. phillipsiana, another pre-Columbian cultivar species in Arizona, differs in having longer leaves and flowers, a longer tepal tube, and longer tepal lobes (Klopper & al. 2010: 63). For further pre-Columbian cultivar species in Arizona, see also under A. verdensis. A. deserti Engelmann (Trans. Acad. Sci. St. Louis 3: 310–311, 370, 1875). Type: not established. — Lit: Trelease (1912b: 45, 52–53, tt. 41–42); Berger (1915: 262–263); Gentry (1978: 15–25, t. 1, with ills.); Gentry (1982: 376–385, with ills.); Turner & al. (1995: 50–54, with ills.); Hodgson (1999a: 5–6); Reveal & Hodgson (2002); Starr (2012: 84–89, with ills.); Webb & Starr (2015: 81–82, with ills.). Distr: SW USA, NW Mexico. The protologue cites specimens from Hitchcock and Palmer (= syntypes; specimens located at F, GH, MO [2 sheets and printed material], NY, PH); Gentry (1978: 16) adds that both were collected on Rancho San Felipe, San Diego County. No attempt for a formal lectotypification seems to have been made so far.

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A large and variable complex with hard-todefine limits, difficult to separate from A. cerulata, and also close to A. subsimplex (Gentry 1982: 405) (see there for differences). Moreover, also the recognized subspecies are not always clearly separable on morphological grounds, and a molecular study of 14 samples embracing all 3 subspecies did not find a clear correlation with Gentry’s infraspecific concept as used here (Navarro-Quezada & al. 2003). Gentry’s (1982) ssp. pringlei is here recognized at species rank (as A. pringlei) following Webb & Starr (2015). A. deserti ssp. deserti — Lit: Trelease (1912b: 45, 52–53, tt. 41–42); Berger (1915: 262–263); Gentry (1978: 15–25, t. 1, with ills.); Gentry (1982: 376–380, with ills.); Hodgson (1999a: 5–6, as A. deserti); Reveal & Hodgson (2002:

Fig. 20 Agave deserti ssp. deserti (Eggli s.n.: USA; California, Anza Borrego State Park). (Copyright: U. Eggli)

Agave AGAVACEAE

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Fig. 21 Agave deserti ssp. deserti (Eggli s.n.: USA; California, Anza Borrego State Park). (Copyright: U. Eggli)

with ills., as var.); Starr (2012: 84–89, ills.); Webb & Starr (2015: 77, 81–82, with ill.). Distr: USA (S California), Mexico (N & C Baja California); sandy to gravelly or rocky places, in desert scrub and pinyon-juniper woodlands, 500–1500 m; flowers mainly April to July, also November to January and in September. I: Irish & Irish (2000: t. 14); Heller (2006: 30); Richter (2011: 80, 125); Pilbeam (2013: 72); Hochstätter (2015: V: 9). – Figs. 20 and 21. Incl. Agave deserti Orcutt (1883) (nom. illeg., ICN Art. 53.1); incl. Agave consociata Trelease (1912). [2i] Ros mostly 30–70
40–80 cm, rather open, sparingly or prolifically suckering; L variable, ascending, narrowly triangular-lanceolate to lanceolate to linear-lanceolate, thick, rigid, scarcely narrowed above the broad clasping convex base, acuminate to long-acuminate, deeply guttered towards the apex, mostly 25–70
4.5–10 cm, 4–7
as long as broad, greyish-white, grey, blueglaucous to (yellowish-) green, often lightly crosszoned, margins usually straight but sometimes undulate or crenate; marginal teeth usually regularly spaced, loosely to firmly attached, variously curved or reflexed, smaller teeth 2–3 mm, longer teeth 5–10 mm, grey, mostly 10–30 mm apart, slender-tipped, brown to pruinose-grey, sometimes with a brown ring at the base; terminal Sp subulate to acicular, openly grooved, strong, generally

20–40 mm, light brown to greyish, decurrent to the 1. or 2. tooth to mid-blade; Inf 2–6 m, paniculate, peduncle slender to thick, green to glaucous, with scarious triangular Bra 8–15 cm long, fertile part usually narrowly to somewhat broadly paniculate, part-Inf short, small, 6–15 in the upper of the inflorescence; Fl 30–60 mm; Tep yellow, tube shallow, angular, openly funnel-shaped, 3–10
9–15 mm, pale green to yellow, lined with a thick nectariferous disk, lobes (broadly) linear, erect to spreading, rounded and abruptly hooked inwards at the apex, equal, 13–20
4–6.5 mm, (light) yellow, wilting at anthesis; Fil 25–42 mm, unequally inserted 2.75–3.75 mm above the tube base, or subequally at the base of the lobes, pale yellow; Anth 13–21 mm, yellow; Ov 15–30 (–38) mm, greenish, neck slightly narrowed to constricted, 4–6 mm, yellow to pale green; Sty 24–29 mm, pale yellow; Fr ovoid to oblong or obovoid, mostly 35–55
12–20 mm, shortly stipitate, apex beaked or shortly rounded to rostrate; Se 5–6
4–4.5 mm, black. — Cytology: 2n = 59, 60, 118 (Cave 1964, Simpson & al. 2011). Field experiments (Jordan & Nobel 1979) showed that successful seedling establishment takes only place in 1 out of 17 years. The length of the first drought period is the limiting factor for establishment, with water-stress in the seedling stage as the single most important factor. For a

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J. Thiede

Fig. 22 Agave deserti ssp. simplex (Etter & Kristen 250: USA; Arizona, Maricopa County, Saddle Mountain, 350 m). (Copyright: J. Etter & M. Kristen)

summary of many further ecological and ecophysiological studies on A. deserti see Nobel (1988). A. deserti ssp. simplex Gentry (Occas. Pap. Calif. Acad. Sci. 130: 22, ills. (pp. 23–24), 1978). Type: USA, Arizona (Gentry 23404 [US, ARIZ, ASU, DES, MEXU]). — Lit: Gentry (1982: 382–385, with ills.); Hodgson (1999a: 6); Reveal & Hodgson (2002: as var.); Starr (2012: 84–89, with ills.); Webb & Starr (2015: 77, 81–82, with ill.); Parker (2017: 256, 260, with ills., as var.). Distr: USA (SW Arizona, SE California), Mexico (NW Sonora); sandy to gravelly or rocky places, in desert scrub and pinyon-juniper woodlands, 300–1500 m; flowers mainly May and June, also in November and February. I: Heller (2006: 80); Pilbeam (2013: 74); Ettelt (2014: 123); Hochstätter (2015: V: 11). – Fig. 22.  Agave deserti var. simplex (Gentry) W. C. Hodgson & Reveal (2001). [2i] Differs from ssp. deserti: Ros usually solitary, rarely with 1–3 offsets; L lanceolate, moderately acuminate, light green to light glaucous-grey, margins repand to crenate; marginal teeth friable; Inf 4–6 m; part-Inf in the upper of the inflorescence. — Cytology: 2n = 60 (Reveal & Hodgson 2002). Ssp. simplex refers to the N and E populations of the species from hot, dry and low elevations, characterized by a predominantly solitary habit (Gentry 1982, Parker 2017). It hybridizes with A. schottii ssp. schottii, and possibly with A.

mckelveyana (Hodgson 1999a, Reveal & Hodgson 2002). It is esp. well-suited for hot and dry gardens in the SW USA (Parker 2017). A. desmetiana Jacobi (Hamburg. Gart.- & Blumenzeit. 22: 217, fig. 32, 1866). Type [neo]: Mexico, Sinaloa (Gentry 11569 [US, ARIZ, MEXU]). — Lit: Berger (1915: 234, as A. miradorensis); Gentry (1972: 94–99, with ills.); Gentry (1982: 622–624, with ills.); Reveal & Hodgson (2002); Franck (2012: 7–8, with ills.). Distr: Cultivated only; flowers September to November. I: Irish & Irish (2000: tt. 15–16); Pilbeam (2013: 75); Hochstätter (2015: VII: 40, as A. demeesteriana); Moore (2016: 267, 268). Incl. Agave demeesteriana Jacobi (1866); incl. Agave regeliana Jacobi (1866)  Agave miradorensis var. regeliana (Jacobi) A. Terracciano (1885) (incorrect name, ICN Art. 11.4); incl. Agave ananassoides Jacobi (1868); incl. Agave miradorensis Jacobi (1868). [2c] Stems short, trunk 20 mm apart or few and irregularly spaced; terminal Sp subulate, shortly and broadly grooved above, 20–30 mm, dark brown to reddish-brown; Inf 2.5–3 m, paniculate, peduncle with persistent triangular Bra 0.5–2 cm long, fertile part long, narrow and compact, often bulbilliferous, part-Inf ascending, congested, 20–36 in the upper of the inflorescence, with 15–21 flowers, >10 cm long; Fl erect, 40–60 mm; Tep green in bud, pale yellow at anthesis, drying ferrugineous, tube salverform, 10–12
14–17 mm, distinctly flanged inside below the inner tepals, deeply grooved and ridged on the outside, lobes erect, nearly equal, 13–15 mm; OTep 1 mm longer; ITep sharply involute; St long-exserted; Fil erect, 30–40 mm, yellow, inserted near the mouth of the tube; Anth centric, 13–26 mm, yellow; Ov 15–25 mm, 6-grooved in the upper part, shortly stipitate, neck 1–3 mm, not constricted; Fr and Se unknown. — Cytology: 2n = 60 (Reveal & Hodgson 2002). Distinguished by its smooth unarmed arching leaves and the short compact inflorescences with small flowers with a very short ovary and a broad tube (Gentry 1982: 623). According to Ullrich (1990d), it was originally introduced from Cuba as A. anomala, where it may have originated (see also under that species). Trelease (1920: 121) assumed an origin from El Mirador (Huatusco, Veracruz), based on the epithet of the synonymous A. miradorensis. Gentry (1972) and Gentry (1982) found it cultivated in Sonora and Sinaloa. Its systematic position is unclear, since it does not fit well into any section; Gentry’s placement in Sect. Sisalanae (as Group) is regarded as artificial by Ullrich (1990d). Here, Reveal & Hodgson (2002) and Govaerts (2014+) are followed who regard the species as a sterile, possibly ancient cultivar possibly derived from A. sisalana/A. kewensis or A. kewensis, respectively. It is consequently placed in Sect. Sisalanae. Govaerts (2014+) considers the earlier A. demeesteriana as correct name for the species. A conservation proposal is thus necessary to retain the established name A. desmetiana. The epithet honours the Belgian nurseryman De Smet but is frequently misspelled ‘desmettiana’. The species is naturalized in Florida (Franck 2012). The plant depicted as naturalized in Spain (Guillot Ortiz &

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Meer 2009a) has larger, cross-banded leaves and may merely represent A. sisalana. ‘Joe Hoak’ is a variegated cultivar traded in the USA (Starr 2014: 222, with ill.). Guillot Ortiz & Meer (2016d) discuss variegated and normal-leaved cultivars cultivated in Spain. Variegated plants are also shown by Moore (2016). A. difformis A. Berger (Agaven, 95–96, 1915). Type: Ex cult. (Berger s.n. [US, K]). — Lit: Gentry (1982: 135–138); Eguiarte & Scheinvar (2008: 44–47); both with ills.; Trejo-Salazar & al. (2015: pollination); López-Pujol & al. (2016). Distr: Mexico (W-C San Luis Potosí, N-C Querétaro, Hidalgo); coarse limestone rocky soils, in desert and submontane scrub on the arid side of the Sierra Madre Oriental, 1560–2100 m; flowers May to August; neophyte in Spain. I: Curtis’s Bot. Mag. 139: t. 8481, 1912, as A. haynaldii; Heller (2006: 82); Richter (2011: 68); Pilbeam (2013: 76); Hochstätter (2015: IX: 82); Greulich (2016c: 151). Incl. Agave haynaldii Todaro (1878). [1g] Stems short; Ros open, rather vigorous, variable, 0.7–1
1–1.5 m, freely suckering; L 25–50, polymorphic, straight or falcate or sinuous, stiffly ascending, thickly convex below, concave above, 50–80
4–6 cm, (dark) green to yellow-green, margins straight or repand, firm or detachable, predominantly light grey; marginal teeth variable, generally 5–10 mm, dark brown to grey, 20–30 mm apart, rarely double, sometimes with smaller intermittent teeth, or reduced or entirely lacking; terminal Sp conical-subulate, stout, with a short open groove above, 15–30 mm, dark brown to grey; Inf 3.5–6 m, ‘spicate’, peduncle waxy-glaucous, with thin, narrow, chartaceous, deflected Bra, fertile part slender, in the upper of the inflorescence; fertile Bra mostly longer than the flowers; Fl 30–40 mm; Tep light green to yellow and pink, tube flaring, 2.5–3.5 mm, lobes mostly erect, equal, sharply hooded at the apex, 15–18
3–4 mm, pale yellow or tinged pink, clasping the filaments at anthesis; Fil 35–45 mm, pale green (Gentry: pale to pinkish), inserted on the rim of the tube; Anth excentric, 15–18 mm, yellow or bronze-coloured; Ov 15–21 mm, waxy green, neck short; Sty slender, pale green, somewhat darker below the

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stigma; Sti slightly thickened; Fr oblong, transversely rugose, 22–26
12–15 mm, dark brown, shortly beaked, sessile; Se crescent-shaped, 5
3 mm, somewhat wavy on the faces, wing low, hilar notch small. A robust species, within Sect. Heteracanthae (as Group Marginatae) characterized by its very polymorphic long-ensiform leaves. Some forms resemble A. lechuguilla (Gentry 1982: 137). Magallán Hernández & Hernández Sandoval (2001) provide a new record for Querétaro. — The 5 type specimens at US bear 4 different dates and thus represent different gatherings. A. haynaldii, if really conspecific, antedates the established name A. difformis, and a formal proposal to conserve the established name is necessary. From 740 flowers produced at average, only 47% develop into fruits. These may have 117 ovules of which only 37% form viable seeds. When the fruits of an inflorescence open, they release about 23,100 viable seeds dispersed by wind. Nocturnal nectar-feeding bats are the most important flower visitors. In the morning, bees, hummingbirds, hymenopterans and bumblebees visit the flowers, and at sunset various hawkmoths. The species shows relatively high levels of genetic diversity with little differentiation among the populations which is possibly generated by high gene flow provided by the pollinators (Eguiarte & Scheinvar 2008). The pollination biology of this and further sympatric species was studied by Rocha & al. (2005) and Trejo-Salazar & al. (2015), recording bats and sphingids as primary pollinators. LópezPujol & al. (2016) report the species as neophyte from Spain. A. doctorensis L. Hernández & Magallán (Brittonia 67(1): 1–3, ills., 2014). Type: Mexico, Querétaro (Hernández 6094 [QMEX, ENCB, IEB, MEXU, TEX, XAL]). — Distr: Mexico (E-C Querétaro: Sierra del Doctor); on gentle slopes and flatlands on a superficial calcareous substrate, in rosette scrub or in juniper forests, 2400–2600 m; flowers in November (?). I: Hochstätter (2015: IX: 28). [1g
2a?] Ros open, 1–1.2
0.8–1 m, surculose; L 15–25 (–30), erect, lanceolate to

J. Thiede

narrowly elliptic, rigid, smooth, 50–75 (–95)
4–9.5 cm in the middle, pale green with imprints from the next inner leaves, margins corneous, easily detached, straight, to 1 mm wide, reddish, covered with narrow lines of wax that can be lost with age; marginal teeth deltoid or hooked, bases wide, retrorse, brittle, flat, 1–2 (–8)
1 mm, 15–45 mm apart, abundant close to the base, absent near the leaf tip, with small interstitial teeth, greyish, or occasionally toothless; terminal Sp subulate, curved to straight, grooved above, 27–43 mm, slightly decurrent; Inf 4.2–4.5 m, ‘racemose-paniculate’, peduncle with broadly acuminate papyraceous to chartaceous Bra 17–28
2.5–6.5 cm, beige to greenish, spine-tipped, reflexed, fertile part only 12–25% of the total inflorescence height, part-Inf 70–80, compact, conical, dorsiventrally flattened, with 6–7 flowers, the lower 7 cm, diminishing to 1.8 cm towards the tip; Ped 5–30 mm; Fl 42–54 mm; Tep yellowish-green, tube funnelshaped, 4–10
5–8 mm, lobes oblong to linear, channelled, unequal, 13–23
16–25 mm, outer 3–4 mm longer than the inner, margins involute, apex dark brown to reddish, thick, inner lobes cucullate; Fil dorsiventrally flattened, 35–45 mm, the outer inserted 5–7 mm from base of the tube, the inner 4–6 mm from the base of the tube; Anth 18–24 mm, yellow; Ov broadly elliptic, (20–) 24–28 (–33) mm, neck constricted; Sty  as long as the filaments; Fr usually oblong, 27–35
13–15 mm, beaked, dark brown, perianth persisting almost to maturity; Se crescentshaped to deltoid, 5–6
1–2 mm, shiny black. Placed in Sect. Heteracanthae (as Group Marginatae) in the protologue and compared with A.
glomeruliflora, which is the name used for a series of natural intersubgeneric hybrids between Subgen. Littaea and Subgen. Agave with intermediate ‘racemose-paniculate’ inflorescences (see there), as well as with A. montiumsancticaroli with similar intermediate inflorescences (and thus supposedly another intersubgeneric hybrid; see there). A. doctorensis also has these intermediate ‘racemose-paniculate’ inflorescences and possibly represents the hybrid between the sympatric A. americana (Subgen. Agave) and A. funkiana (Subgen. Littaea).

Agave AGAVACEAE

A. dolichantha Thiede & Eggli (Kakt. and. Sukk. 50(5): 111, 1999). Type [syn]: Mexico, Jalisco (Rose & Hay 6290 [US, GH, NY, P]). — Lit: Rose (1903a: 10); McVaugh (1989: 253–254); Cedano & al. (1995); Cházaro Basáñez & Machuca Núñez (1995: with ill.); García-Mendoza (2003); Cházaro Basáñez & al. (2010b: with ills.); Machuca Núñez & Cházaro Basáñez (2015: with ills.); all as Polianthes longiflora. Distr: Mexico (Jalisco, Michoacán); wet meadows, in ill-drained clayish soils or at disturbed places, in pine/oak or tropical deciduous forests, 1400–2700 m; flowers July to August. I: Feria-Arroyo & al. (2010: 19); Castro-Castro & al. (2016: 721); Castro-Castro (2017: 137); all as Polianthes longiflora.  Polianthes longiflora Rose (1903); incl. Polianthes michoacana Cedano & al. (1995)  Agave michoacana (Cedano & al.) Thiede & Eggli (1999). [3b1] Herbaceous; corm cylindrical, to 5.5
2.5 cm, with many spirally arranged, deltoid, white leaf scars; bulb ovoid to oblong, (2–) 3–5
1.3–1.8 (–2.5) cm, covered with fibrous dry leaf bases lacerate at the apex; R fibrous, white when young, turning brownish-grey with age; L (2–) 4–6 (–10), recurved, narrowly oblanceolate to linear, semi-amplexicaul, glabrous, apex acute, (14–) 26–37.5
(0.2–) 0.4–1.6 cm, green, with purple dots near the base, upper ones appressed to ascending, linear, carinate, broadly acuminate, 12–18.5
0.6–1 cm, margins hyaline, papillose or denticulate, rarely entire; Inf erect, (14–) 30–70 cm, spike-like, fertile part with 2–5 (–8) floral nodes, internode length decreasing distally; Ped none; Fl geminate, fragrant,  (75–) 85–120 mm; Tep white, pink, sometimes red with age, buds whitish to rose, tube erect or nearly so in the basal part, narrowly tubular, 2 mm ∅, gradually widening in the distal or , there funnel-shaped, curved outwards at or above the middle, tube at the base or sometimes whole tube with purple to pink dots, (63–) 70–106
2–6 (–8) mm, mouth oblique, lobes (somewhat) extended or archedreflexed; OTep oblong to ovate, attenuate towards the rounded apex, 15–27
6 mm; ITep lanceolate, fleshy, margins somewhat wavy, apex cucullate, with short white hairs, 12–27
3–8 mm; Fil filiform, white, 5.6–10 mm, inserted near the

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mouth of the tube; Anth linear, (7–) 12–17 mm, apex scarcely surpassing the tube; Ov cylindrical, 7–10
2.5–5 mm; Sty filiform, 48–87 (–100) mm, included at anthesis, white; Sti 3-lobate, lobes oblong, internally ciliolate; Fr globose, (13–) 20–30
10–16 mm; Se semicircular, 4.4
3.1 mm, shiny or dull. Machuca Núñez & Cházaro Basáñez (2015) report 22 populations from 16 municipios in Jalisco, and in addition known from 3 populations in Michoacán. Since its description by Rose (1903a: 10), the species was long known only from incomplete specimens offered for sale as cut flowers (McVaugh 1989: 253) until its recent discovery in the wild at several localities in Jalisco (Cedano & al. 1995, Cházaro Basáñez & Machuca Núñez 1995). Cedano & al. (1995) published similar plants from Michoacán as Polianthes michoacana, but its differences fall within the range of variability in A. dolichantha so that the populations from Michoacán were included in the latter (García-Mendoza 2003, Cházaro Basáñez & al. 2010b, Feria-Arroyo & al. 2010, Castro-Castro & al. 2016: 722; all as Polianthes longiflora). A. dolichantha is locally harvested for the cut-flower trade; its habitats are threatened by grazing (Cházaro Basáñez & al. 2010b, Machuca Núñez & Cházaro Basáñez 2015, both as P. longiflora). The geographical distribution, niche modelling and extinction risk of the species are dealt with by Solano & Feria (2007) and FeriaArroyo & al. (2010). — The “holotype” at US consists of 2 sheets which are not cross-labelled and thus represent syntypes; US 396106 is the true species, whereas US 396105 is Polianthes sessiliflora (here treated as A. apedicellata) according to a label by E. Solano C. placed on the specimen in the year 2000 (see also McVaugh 1989: 254). — When placed in Agave, a new name was necessary to avoid homonymy with A. longiflora (Rose) G. D. Rowley (1977). A. durangensis Gentry (Agaves Cont. North Amer., 433–436, ills., 1982). Type: Mexico, Durango (Gentry & Gilly 10576 [US, ARIZ, MEXU, MICH]). — Lit: Cházaro Basáñez & Vázquez-García (2006: with ills.); VázquezGarcía & al. (2007b: 51–52, t. H); González-

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Elizondo & al. (2009: 65–68, with ills.); AlmarazAbarca & al. (2013b: genetic variability). Distr: Mexico (S Durango, N & C Zacatecas); rocky slopes and gravelly bajadas in grama grasslands, pine-oak or sometimes tropical deciduous forests, 1700–2600 m; flowers April to July, also in November. I: Etter & Kristen (2007b: 177); Klopper & al. (2010: 60); Richter (2011: 52); Pilbeam (2013: 77); Hochstätter (2015: IV: 8). [2h] Stems short; Ros (0.6–) 0.8–1.2 (–1.8)
(0.7–) 1.2–1.8 (–3.1) m, solitary or caespitose; L broadly lanceolate, narrowed above the broad base, widest in the middle, straight to outcurving, flat to concave esp. towards the apex, thick and convex towards the base, asperous, mostly 40–90 (–130)
14–22 cm, glaucous-grey, pruinose, margins heavily armed, deeply crenate-mamillate; marginal teeth variously curved, prominent, broadly flattened, 10–20 mm, generally 10–20 mm apart; terminal Sp strong, broadly channelled above, (30–) 40–60 mm, pruinose-grey over brown; Inf 7–8 m, paniculate, peduncle short, with scarious remote reflexed Bra 15–25 cm long, fertile part long, lax, axis flexuous or zig-zag, part-Inf 18–30, sinuously spreading, trifurcate, small, in the upper of the inflorescence; Fl 60–80 mm; Tep yellow, tube cylindrical, broad and fleshy, lightly grooved, 15–22
12–17 mm, lobes strictly appressed to the filaments, becoming leathery, unequal; OTep larger, 10–12 mm, thickly rounded on the back and overlapping the inner, apex conspicuously papillate, almost horny, reddish; ITep smaller, sharply keeled; Fil somewhat flattened, 48–60 mm, inserted at 2 levels 8–12 and 6–10 mm above the tube base; Anth 18–25 mm; Ov 30–45 mm (incl. the unconstricted neck); Sty somewhat longer than the tepals; Sti capitate; Fr oblong, 45–60
16–18 mm, rounded and shortly beaked, obscurely stipitate; Se small, crescent-shaped to obovate or tear-shaped, 4.5–6
3.5–5 mm, with a broad but little raised rim wing. Distinguished by the large, rigid, broadly lanceolate leaves heavily armed with flexuous teeth on deeply crenate margins, and the large, long, lax panicle with a flexuous or zigzag axis, a short peduncle, and without close relationship to other species of Sect. Ditepalae (as Group) according

J. Thiede

to Gentry (1982). Vegetatively, it may be confused with the sympatric A. scabra of Sect. Agave (Gentry 1982: 436, as Group Americanae). A. temacapulinensis, also of Sect. Ditepalae, is very similar (see there). A. durangensis is economically important for the mescal industry in Durango (ContrerasHernández & al. 2016). Molecular studies of 3 natural populations found evidence for genetic differences between the populations, some incongruence between molecular data and morphology, and advanced diversification in 1 of the 3 populations (Almaraz-Abarca & al. 2013b). Seed morphology, germinability and phenolic profiles varied among 3 natural populations (Barriada-Bernal & al. 2013). Pollen and foliar phenol profiles indicate the presence of several chemotypes and allow the separation of the 2 studied populations (AlmarazAbarca & al. 2009). The edible flowers of A. durangensis represent an important source of antioxidant flavonols (Barriada-Bernal & al. 2014). The arthropod communities associated with the species were studied by González-Castillo & al. (2011). A. eggersiana Trelease (Mem. Nation. Acad. Sci. 11: 28, tt. 31–33, 1913). Type: US Virgin Islands, St. Croix (Ricksecker 282 [MO, US]). — Lit: Berger (1915: 220); Britton & Wilson (1923: 156); Proctor & Acevedo-Rodríguez (2005: 118); Chamorro & al. (2016: with ills.). Distr: US Virgin Islands (St. Croix); coastal cliffs and dry coastal shrubland, in open-canopy and openunderstorey habitats exposed to full sun. I: Hochstätter (2015: VII: 60). [2s] Acaulescent, not surculose; L numerous, narrowly lanceolate, nearly straight and erect, gradually acute, concave, mostly 120–200
10–15 cm, dull green, transiently slightly glaucous; marginal teeth straight or upcurved, narrowly triangular, 1–1.5 mm, usually 10 mm apart, with broadly lenticular blackish-brown base, intervening margin nearly straight, at first reddish; terminal Sp conically subulate, usually somewhat upcurved, often laterally compressed, smooth, 15–20 mm, brown, rather glossy, with slender dull-brown involute basal thickening to 3
10–15 mm, decurrent for its length or more and

Agave AGAVACEAE

dorsally intruded into the green leaf tissue; Inf 5–7 m, ‘paniculate’, peduncle with distant, deltate, spreading or reflexed Bra, fertile part densely flowered, part-Inf (slightly) ascending, 25, freely bulbilliferous, in the upper of the inflorescence; Ped 20–25 mm; Fl 50–60 mm, pumpkin-scented; Tep deep yellow, tube broadly open, 5–7 mm, lobes 20–25
5–8 mm; Fil 45–50 mm, inserted nearly in the throat; Anth not described; Ov oblong-fusiform, 20–30 mm; Fr and Se not produced. For the typification, see Smith & Figueiredo (2014e: 233). Endemic to St. Croix in the US Virgin Islands where it now appears to be largely extinct in the wild, but persisting as relatively few cultivated plants on St. Croix as well as on St. Thomas (Proctor & Acevedo-Rodríguez 2005: 118). The species is threatened by the Agave Snout Weevil (Scyphophorus acupunctatus) recently reported from St. Croix; Chamorro & al. (2016) provide management recommendations. In the first edition of this handbook, A. eggersiana was placed in the synonymy of A. missionum (the only other native Agave species on the US Virgin Islands) based on Álvarez de Zayas (1995), but it is here recognized as a distinct species following Proctor & Acevedo-Rodríguez (2005). Agave missionum differs from A. eggersiana by broader leaves and longer marginal teeth (Proctor & Acevedo-Rodríguez 2005: 117). A. ellemeetiana K. Koch (Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 8: 103 [1. April], 1865). Type [neo]: Ex cult. (Anonymous s.n. [K 524804, K [524802 + 524803]]). — Lit: Gentry (1982: 94–97); Thiede (2014b); both with ills. Distr: Mexico (C Veracruz, N & W Oaxaca). Incl. Agave ellemeetiana Regel (1865) (nom. illeg., ICN Art. 53.1); incl. Agave ellemeetiana Jacobi (1865) (nom. illeg., ICN Art. 53.1); incl. Agave ellemeetiana [?] obovata Hort. Sutherland (1875) (nom. inval., ICN Art. 38.1a). A. ellemeetiana was previously ascribed to Jacobi (Hamburg. Gart.- & Blumenzeit. 21: 457, 1865), but was first published as cited above (Thiede 2014b). Ullrich’s (1991k) notion that the species might possibly represent hardly more than

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a garden clone of A. attenuata ssp. dentata (as A. pedunculifera) must be dismissed. A. ellemeetiana differs from A. attenuata esp. in its nearly stemless habit with even fewer, bright green leaves, and its eastern distribution. For the history, rediscovery, distribution and habitats of the species, see Thiede (2014b). A. ellemeetiana ssp. ellemeetiana — Lit: Berger (1915: 125–126); Gentry (1982: 94–97); Alsemgeest (2004); Cházaro Basáñez & al. (2012); Thiede (2014b); all with ills. Distr: Mexico (C Veracruz, N Oaxaca); on often near-vertical cliffs and rock-faces, in cloud or tropical subdeciduous forests, in the cloud belt, rarely epiphytic, 400–1460 m; flowers February to May. I: Refug. Bot. 3: t. 163, 1869; Baker (1888b); Heller (2006: 83); Richter (2011: 62); Hochstätter (2015: VIII: 36); Castillo-Hernández & Flores-Olvera (2017: 541, as A. gomezpompae). [1c] Stems nearly none; Ros open, 0.35–0.5
0.7–1 m, surculose; L rather few, ovate to oblong, somewhat recurved, reclining at maturity, thickly soft-succulent, widest in the middle, acuminate, flat beyond the thick base, concave to flat above, smooth, 50–70
12–20 cm, light bright green, margins thin, friable, smooth, unarmed, white, sometimes reddish, sometimes finely serrulate towards the leaf tip; terminal Sp none, but leaf tip shortly acuminate and slightly calloused; Inf erect, 3–4.5 m, ‘spicate’, peduncle with ovate caudate-acuminate Bra 8–10 cm long, fertile part densely flowered from near the base, part-Inf usually with 4 flowers; Ped united in pairs, 15–20 mm; Fl campanulate, 28–40 mm; Tep pale greenish-yellow, tube very short, 1–2 mm, lobes lanceolate, concave, somewhat hooded at the tip, 13–15
5 mm, inner somewhat wider; Fil long exserted, 50–60 mm, inserted on the rim of the tube together with the lobes; Anth 10–12 mm, yellow; Ov bottle-shaped, 13–20 mm, neck conspicuously elongate; Sty slender, as long as the filaments; Sti thickened; Fr trigonous, rounded at the base, 13–15
10 mm, light brown, beaked; Se numerous, 3 mm, black, shiny. Introduced into cultivation from Mexico in 1862 or before, and persisting in cultivation esp. in Europe up to the present. Its natural habitats

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Fig. 23 Agave ellemeetiana ssp. subdentata (Etter & Kristen 3905: Mexico; Oaxaca,  10 km S of Yosonicaje, 2320 m). (Copyright: J. Etter & M. Kristen)

remained unknown until the recent past. At present, ssp. ellemeetiana is known from 3 areas in the Sierra Madre Oriental of C Veracruz and NE Oaxaca (Thiede 2014b), namely from the Sierra de Zongolica in C Veracruz (Cházaro Basáñez & al. 2012), and in N Oaxaca from the Sierra Mazateca (= Sierra de Huautla) (Lorea Hernández & MunnEstrada 2005) and from near San Bartolomé Ayautla where it was first observed by A. B. Lau (Ullrich 2007: fig. 6, as “A. attenuata?”). A. ellemeetiana ssp. subdentata (Trelease) Thiede (Bradleya 32: 157, 2014). Type [neo]: Mexico, Oaxaca (Spath & Griffin s.n. [ZSS]). — Lit: Köhres (2008: with ills., as A. ellemeetiana). Distr: Mexico (W Oaxaca: Putla/Tlaxiaco/Santiago Nuyoó area); volcanic rocks in cloud or tropical subdeciduous forests, 2300–2600 m; flowers in January. I: Pilbeam (2013: 78, as A. ellemeetiana); Hochstätter (2015: VIII. 37). – Fig. 23.  Agave ellemeetiana var. subdentata Trelease (1914). [1c] Differs from ssp. ellemeetiana: L margins whitish or reddish, finely but clearly denticulate throughout; L of adult plants have a wax bloom absent from ssp. ellemeetiana and appear to be partly broader, but measurements are not available (Thiede 2014b). Köhres (2008) reported plants of A. ellemeetiana from the W slopes of the Sierra Madre del Sur in Oaxaca above Santiago Nuyoó, and

published the first habitat photographs of it. Since these plants differ from typical A. ellemeetiana by leaf margins being finely denticulate throughout, Thiede (2014b) re-established the forgotten name var. subdentata Trelease (described as having the “leaves very minutely denticulate”) at subspecific rank. Localities of the subspecies in the Putla/Tlaxiaco/Santiago Nuyoó area were also visited by other collectors, and plants belonging here are commercially offered in the USA and Europe. A preliminary investigation using AFLP data placed material from Putla probably belonging to ssp. subdentata separate from A. ellemeetiana (Gil-Vega & al. 2007: as A. aff. pedunculifera). Further studies are needed to establish the status of ssp. subdentata and its differences from ssp. ellemeetiana. A.
engelmannii Trelease pro sp. (Annual Rep. Missouri Bot. Gard. 3: 167, tt. 55–56, 1892). Type: Ex cult. Missouri BG (Anonymous s.n. [K [2 sheets, cross-labelled]]). — Lit: Anonymous (1892: 398); Berger (1915: 184–185); both as A. engelmannii. Distr: Cultivated only. Incl. Agave attenuata var. subdentata Hort. Hoopes ex Trelease (1892) (nom. inval., ICN Art. 38.1a). [1b?
2] Raised from seed by J. Hoopes and flowered and fruited at the Missouri Botanical Garden 1890–1891. Seedlings from Missouri flowered later at La Mortola and were described

Agave AGAVACEAE

by Berger (1915). Placed close to A. horrida ssp. perotensis (as “A. polyacantha”) by Trelease (1920: 135) and in the synonymy of that species by Gentry (1982: 228). With inflorescences intermediate between the subgenera, with distinctly stalked part-inflorescences with up to 7 flowers (protologue t. 55, sheet K 001096501!) and thus obviously of intersubgeneric hybrid origin involving a species of Subgen. Littaea (possibly A. attenuata) and one of Subgen. Agave. A. ensifera Jacobi (Nachtr. Versuch syst. Glied. Agaveen 1: 13–14, 1868). Type [neo]: Ex cult. La Mortola (Berger s.n. [US 1023791]). — Lit: Baker (1877a: 369, with ill., as A. heteracantha); Berger (1915: 99); Gentry (1982: 139); Meer & Puche (2014: as A. univittata var. ensifera, A. lophantha var. latifolia and A. heteracantha). Distr: Known from cultivation only. I: Roosbroeck (2014); Hochstätter (2015: VII: 29).  Agave univittata var. ensifera (Jacobi) P. Van der Meer & C. Puche (2014); incl. Agave heteracantha Baker (1877) (nom. illeg., ICN Art. 53.1); incl. Agave lophantha var. latifolia A. Berger (1915)  Agave univittata var. latifolia (A. Berger) Breitung (1959). [1g] Ros dense, caespitose; L linear-lanceolate, ensiform, leathery-fleshy, strongly convex below and above, upwards becoming concave to the apex, at the base 1.5–1.7 cm thick, smooth, 50–60
4–5 (3.8–4 near the base) cm, dark green with a distinct light longitudinal central stripe 5–7 mm wide up to the middle, margins with a narrow grey border 0.5–1 mm wide; marginal teeth mostly antrorsely curved, 4–6 mm, light grey, closely set, 10–20 mm apart, mostly with broad bases, interspersed with smaller teeth, altogether 30–40 teeth per leaf side; terminal Sp short, basal groove above short and opening broadly with decurrent border, 10–15 mm, brown to grey; Inf 2–2.5 m, ‘spicate’, peduncle with deflected subulate Bra 6–10 cm long, partInf mostly with geminate flowers; Ped 2–3 mm; Fl 35–42 mm; Tep light green, tube open, short, 2–3 mm, lobes linear, involute around the filaments at anthesis, subequal, 14–17 mm, light yellowish, outer overlapping the inner at the base; Fil slender, 40–45 mm, inserted on the tube rim;

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Anth 16 mm, yellow; Ov 20–24 mm, neck constricted, 3 mm; Sty whitish, equalling the anthers; Fr and Se unknown. Of unknown origin and commonly cultivated along the Mediterranean Riviera at Berger’s time; apparently related to the A. lechuguilla – A. difformis group and recognized specifically by its long narrow leaves with numerous teeth of 2 sizes regularly spaced along the narrow margins, and the short, mostly weak terminal spine (Gentry 1982: 139). Meer & Puche (2014) classified the plant as A. univittata var. ensifera, but the name A. univittata var. latifolia (A. Berger) Breitung would take priority. The plants depicted as A. lophantha var. latifolia by Guillot Ortiz & Meer (2012: 33) and as A. univittata var. ensifera, A. lophantha var. latifolia and A. heteracantha by Meer & Puche (2014: 59–62, 64) have lanceolate leaves with teeth on distinct prominences and are most probably misidentified. Gentry (1982: 139) selected two specimens of Berger (Ch^ateau Grimaldi, Berger s.n., 19. 6. 1909, US 1023791 & 1023763) as “lectotype”, but since Jacobi did not cite any material, this is interpreted as first-step neotypification (ICN Art. 9.17) as they are not cross-labelled. Meer & Puche (2014: 59) designated US 1023791 as “lectotypus” (but actually representing a neotype), and US 1023763 as well as a third specimen (Berger s. n., 19. 6. 1909, ARIZ 268297) as “isolectotype”. These specimens are both not cross-labelled and are thus interpreted to represent different gatherings and not isoneotypes. A. evadens Trelease (Mem. Nation. Acad. Sci. 11: 20–21, tt. 9–10, 116, 1913). Type: Trinidad (Crueger 1333 [B †, K 000524793, MO, NY, U, UPS, US]). — Lit: Berger (1915: 225); Britton & Wilson (1923: 157); Hummelinck (1938: 23–24). Distr: Trinidad (along the coast and on Isla de Chacachacare and Isla Monos), Venezuela (Isla de Patos); littoral deciduous seasonal forests; flowers in March. I: Richter (2011: 97); Hochstätter (2015: VII: 41). Incl. Agave polyacantha Baker (1888) (nom. illeg., ICN Art. 53.1); incl. Agave vivipara Hart (1890) (nom. illeg., ICN Art. 53.1); incl. Agave polyantha Dodge (1897).

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[2u] Ros shortly caulescent, not surculose; L narrowly oblanceolate, gradually acute, openly concave or somewhat conduplicate, or with inrolled margins above, 70–100 cm, green, margins almost straight; marginal teeth small, on low broadly conical prominences, 0.5–1.5 mm, 6–15 mm apart, margins between the teeth convex; terminal Sp conical, straight or somewhat recurved, slightly involute at the base, 10–14
2–3 mm, slightly or not decurrent; Inf paniculate, fertile part lax, slender, part-Inf few, ascending, tripartite, not known to be bulbilliferous; Ped rather slender, (6–) 8–10 (–12) mm; Fl 47–55 mm; Tep bright yellow, tube open, 2–3.5 mm, lobes 19–25
4 mm; Fil 35–42 mm, inserted 0–0.5 mm below the tube throat; Anth 17–25 mm; Ov oblong-fusiform, 22–25
3–4 mm; Sty 32–38 mm; Fr broadly ellipsoid, 30–34
20–24 mm, slightly beaked, distinctly stipitate; Se D-shaped, angled, 6–8
4–6 mm, shiny black. According to the protologue intermediate in foliage between A. cocui and A. boldinghiana and known to Trelease from photographs and dissociated flowers only. If the name A. polyantha Dodge really proves to be conspecific, it would have priority. The characters regarded in the protologue as most important were taken by Trelease from his plate 9: Shortly caulescent, leaves narrowly oblanceolate, inflorescence laxly panicled at the end with few ascending branches. Hummelinck (1938: 24) assumed that these data have no value: the short stem appears to result from the bases of the plants being washed free, the narrowly oblanceolate leaves may be due to external circumstances, and the laxly panicled inflorescence may result from the great reduction of the picture which made flowerless branches invisible. Apart from Trinidad (main island), A. evadens is also reported from Isla de Chacachacare (Hummelinck 1938: 23, Adams & Baksh-Comeau 2005: 9) and Isla Monos (Chalmers 1965) and from Isla de Patos (Williams 1924: 278), which since 1942 belongs to Venezuela. Preserved flowers from Isla Margarita (Venezuela) placed here with reservation by Trelease may belong to A. cocui or A. vicina (as A. vivipara; Hummelinck (1938)). Material from Puerto Santo (Venezuela)

J. Thiede

“may possibly belong to A. evadens” (Hummelinck 1938: 24). Several specimens from the Venezuelan mainland placed here (Pittier 10238, US 1187053; Steyermark 107981, US 2706466; Steyermark 108044, US 2706464) need verification. The species was (is?) cultivated on St. Thomas (US Virgin Islands; Britton & Wilson (1923: 157)). Baksh-Comeau & al. (2016: 33) assign the species to the IUCN category “Vulnerable”. A. felgeri Gentry (US Dept. Agric. Handb. 399: 60–62, ills., 1972). Type: Mexico, Sonora (Gentry 11343 [US, ARIZ, MEXU, MICH]). — Lit: Gentry (1982: 107–110, with ills.); Ullrich (1991j: with ills.); Turner & al. (1995); Smith & Figueiredo (2014b: with ills.). Distr: Mexico (Sonora, esp. Bahía San Carlos); arid desert lowlands and rocky hillsides near the coast, to 200 m; flowers May to August, also October and December. I: Richter (2011: 63); Pilbeam (2013: 79); Hochstätter (2015: VIII: 45). [1d] Ros small, rather open, 20–50 cm ∅, surculose, forming rather densely caespitose groups; L rather few, erect to rigidly spreading, linear to narrowly lanceolate, straight or falcate, widest at the amplexicaul base, convex below, flat above in the lower , concave in the upper , epidermis rugose or scabrous above, 25–35
0.7–1.5 cm, matt light green to yellow-green, turning light yellow during and after flowering, with faint imprints from the central bud, frequently with pale median longitudinal stripe, margins with weakly filiferous narrow brown border, smooth, threads  absent from older leaves; terminal Sp weak, small, slightly grooved at the base, 6–15 mm, grey or dark brown; Inf 1.2–2.5 m, ‘spicate’, peduncle light green, 7–15 mm ∅ near the base, with chartaceous laxly arranged Bra, the lower very narrow and 2–3 cm, fertile part in the upper , part-Inf with 1–2 flowers; Ped strong, 2–5 mm; Fl 25–30 mm; Tep light green, lateral sections creamy-white, the very margins purple, tube 2–4 mm, lobes linear,  equal, 10–12
3–4 mm, strongly recurved to rolled back during the male phase, less so and straight with the onset of the female phase; Fil 20–25 mm, light shiny pinkish-purple, inserted 2 mm above the tube

Agave AGAVACEAE

base; Anth 8–9 mm, dull yellow; Ov terete, 12–14
4–5 mm, abruptly tapering apically, neck distinctly constricted; Sty stout, terete, 30–35 mm, light green; Sti prominent, white; Fr oblong, cylindrical or obovoid to globose, narrowed towards the base, 15–20
9–14 mm, distinctly beaked, light green to yellowish-green; Se irregular, thick, D-shaped, angled, wrinkled, hilar notch narrow and deep, 4–5
3 mm. — Cytology: n = 30 (Baker & al. 2009). Very similar to A. schottii from Sect. Parviflorae in vegetative features, but placed in Sect. Littaea (as Group Filiferae) due to its open shallow flower tube and long lobes, in which it occupies the most arid habitats. The type locality was destroyed for recreational development (Gentry 1982). Additional data on the reproductive morphology and phenology were provided by Smith & Figueiredo (2014b). Geographically it is closest to the broadflowered form of A. schidigera which forms a more robust non-suckering colony on the Cerros del Fuerte in N Sinaloa (Gentry 1982). Ullrich (1991j) emphasizes vegetative and geographical criteria and suggests a placement in Sect. Parviflorae (as Group), but this is not followed here. A. filifera Salm-Dyck (Hort. Dyck., 309, 1834). Type: [neo — icono]: Ill. Hort. 7(4): t. 243, 1860. — Lit: Berger (1915: 73–74); Gentry (1982: 110–111); Vázquez-García & al. (2007a: 51–52, t. I); Starr (2012: 90–93, 317–319); all with ills. except Berger. Distr: Mexico (Aguascalientes, Jalisco, Guanajuato, Querétaro, Hidalgo, México); on rocks, in desert scrub, thorn, oak and oak-juniper forests, 2340–3100 m; flowers June to September. I: Heller (2006: 84); Richter (2011: 63); Pilbeam (2013: 80, as ssp. filifera); Hochstätter (2015: IX: 30). Incl. Agave filifera ssp. filifera; incl. Agave filamentosa Salm-Dyck (1859)  Bonapartea filamentosa (Salm-Dyck) Boucen (1868) (incorrect name, ICN Art. 11.4)  Agave filifera var. filamentosa (Salm-Dyck) Baker (1877); incl. Agave filifera var. superba J. Croucher ex R. Hogg (1874); incl. Agave pseudofilifera Ross & Lanza (1892); incl. Agave filifera var. candida superba Hort. Besaucèle ex A. Berger (1915)

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(nom. inval., ICN Art. 36.1c/38.1a); incl. Agave filifera var. elatior Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave filifera var. immaculata Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/ 38.1a); incl. Agave filifera var. longifolia Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave filifera var. mediopicta Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave filifera var. splendens Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave filifera var. viridis Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a). [1d] Ros small, green, dense, surculose, forming large clumps; L many, lanceolate, straight, thick, broadest in the middle, acuminate, thickened and convex above and below from the base to the middle, smooth, 15–30
(1–) 2–4 cm, green, with white imprints from the central bud, margins finely filiferous; terminal Sp flat above, rounded below, 10–20 mm, greyish; Inf 2–2.5 m, ‘spicate’, peduncle densely covered with long-caudate recurving Bra, fertile part tall, in the upper of the inflorescence, tapering, densely flowered; Ped short, thick; Fl mostly geminate, 30–35 mm, ascending-outcurving; Tep reddish, tube funnelshaped, furrowed, 5–6
8 mm, mouth bulging at the filament insertions, lobes recurving, lanceolate, apiculate, equal, 14
4 mm, inner a little wider, with prominent keel; Fil slender, 30–35 mm, reddish, inserted on the tube rim; Anth centric, 7–12 mm, reddish; Ov fusiform, angled, 13–15 mm, neck furrowed; Sty strong, longer than the filaments; Sti 3-lobate; Fr oblong-ellipsoid to oblong-obovoid, acute, 15–17
12 mm; Se semicrescent-shaped. — Cytology: 2n = 60 (Cave 1964, Simpson & al. 2011). Separable from the closely related A. schidigera by its caespitose habit, smaller rosettes, shorter and thicker leaves, and smaller reddish-purple flowers with a shorter tube (Gentry 1982, Starr 2012). Ullrich (1992h) reduced A. schidigera and A. multifilifera to subspecies of A. filifera, but was not followed by Vázquez-García & al. (2007b), Starr (2012) and Govaerts (2014+: accessed April 2014). The species was recently recorded from E Jalisco (Hernández-Vera & al. 2007a). A record

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from C Zacatecas (Ramírez Prieto 2014: 54) may refer to A. schidigera. A. flexispina Trelease (Contr. US Nation. Herb. 23: 133, 1920). Type [syn]: Mexico, Durango (Palmer 330 [MO, US, CM, GH, K, NY, UC]). — Lit: Gentry (1982: 436–438); GonzálezElizondo & al. (2009: 77–79); Klopper & al. (2010: 58, 60–61); Alsemgeest & Roosbroeck (2010: 68–70); all with ills. Distr: Mexico (S Chihuahua, N Durango, Zacatecas, N Nayarit); rocky slopes and hillsides, in grama or oak-juniper grasslands and oak woodlands, 1300–2440 m; flowers July, October and November. I: Breitung (1963: 121, right-hand figure); Pilbeam (2013: 84); Ettelt (2014: 124); Hochstätter (2015: IV: 9). [2h] Ros open, small, 25–35 (–45)
50–70 cm, solitary or caespitose; L 35–40, ovate, acuminate, nearly flat to concave above, 16–30
6–8 (–12.7) cm, glaucous- to yellowish-green, margins repand to crenate; marginal teeth mostly retrorse, larger teeth mostly 5–8 mm, brown to pruinose, 10–15 mm apart, on small prominences, sometimes with small intermittent teeth; terminal Sp acicular, usually flexuous, flat to openly grooved near the base, 25–35 mm, brown to pruinose-grey, decurrent to the upper teeth; Inf 2–3.5 m, paniculate, peduncle with small, dry, appressed, hardly persistent Bra, fertile part slender, rather open, frequently narrow, part-Inf small, 6–12 (–16), few-flowered; Ped nearly none; Fl 50–70 mm; Tep greenish-yellow, with red tinge on buds and lobes, tube cylindrical to urceolate, bulging in the middle, slightly or narrowly grooved, 10–18 mm, lobes lanceolate to linear, broad, broadly rounded at the apex, unequal, outer larger, strongly overlapping the inner, 10–18 mm, persisting erect and drying leathery, inner with high narrow keel; Fil sometimes broadly flattened, 40–50 mm, with red tinge, inserted at one level, sometimes irregularly so; Anth 17–23 mm, yellow or bronze-coloured; Ov cylindrical, slightly angled, 22–35 mm, neck short, obscure; Sty longer than the filaments; Sti capitate, 3-lobate; Fr oblong, strongly trigonous, round at the apex, tardily dehiscent, 35–45
15–17 mm, abruptly stipitate; Se obovate to crescent-shaped, 5.5–7
4–5 mm, rim wing low.

J. Thiede

In appearance like a small A. shrevei or A. palmeri, but differing in having the filaments inserted at a single level in the tube (vs. 2 distinct levels in the species mentioned, and indeed in most members of Sect. Ditepalae). Rosettes and inflorescences of A. flexispina are consistently smaller than normally found in other species of Sect. Ditepalae (as Group) (Gentry 1982). The holotype is at MO according to the protologue, but the 2 sheets (MO 3346761, leaves; MO 3346762, inflorescence) are not cross-labelled and represent syntypes (Smith & Figueiredo 2014e: 234). The specimen Flores F. & al. 2079 (MEXU 520551!) provides a new record from Nayarit. The left-hand figure in Breitung (1963: 121) is incorrectly labelled and represents A. parryi var. truncata (Alsemgeest & Roosbroeck 2010: 69). A. fortiflora Gentry (US Dept. Agric. Handb. 399: 122–126, ills., 1972). Type: Mexico, Sonora (Gentry 19808 [US, ARIZ, DES, MEXU]). — Lit: Gentry (1982: 421, 438–440, with ills.); Klopper & al. (2010: 61). Distr: Mexico (NW Sonora); on isolated mountain ranges, mostly on volcanic rocks or also on limestone ridges, in desert scrub; flowers in cultivation (California) June to July. I: Heller (2006: 85); Pilbeam (2013: 85). [2h] Ros open, to 1
1.8 m, mostly solitary, occasionally suckering; L straightly ascending or outcurving and conduplicate, long-acuminate, gradually narrowed above the dilated base, widest in the middle, finely tuberculate-rugose (incl. teeth and terminal spine), to 50–100
8–12 cm, light grey-glaucous, usually cross-zoned, margins straight or teeth in the middle of the lamina on small prominences; marginal teeth curved downwards or erect, 5–10 mm (middle of the lamina), 10–30 mm apart, smaller and variously flexed above and below the middle, with irregularly arranged smaller intermittent teeth; terminal Sp subulate, rounded below, narrowly grooved above, 30–50 mm, chestnut-brown to light grey, decurrent along the margin to the uppermost teeth; Inf 4–6 m, paniculate, peduncle with short, triangular, chartaceous Bra 5–6 cm long at the lower branches, fertile part open and long-ovoid, partInf nearly horizontal, 8–15, densely 12- to

Agave AGAVACEAE

18-flowered; Ped 4–20 mm; Fl long-lived, 72–82 mm; Tep yellow, tube broadly bulging, sinuses overlapping and deeply grooved below, 11–13
18–20 mm, pale yellow, lobes erect, 20–23
7–8 mm, yellow, outer rounded, linear-lanceolate, with an involute margin, obtuse, inner with high narrow keel and involute hyaline margins, broadly hooded at the tip; Fil strong, elliptic in cross section, 60 mm, persisting erect after anthesis; Anth centrally affixed, 20–22 mm, yellow; Ov 45–50 mm, pale green, conspicuously angled below the outer tepals; Sty finally to slightly shorter than the filaments; Sti capitate, 3-lobed; Fr and Se unknown. The species is highly distinct in its floral morphology: The large flowers are sturdy (hence the name), and long-lasting, persisting erect for long after anthesis, and the flattening of the filaments is extreme, making them almost strap-shaped and quite the broadest in the genus. With these features, A. fortiflora does not show relations to other species of Sect. Ditepalae (as Group), but the erect tepal lobes dry in the same way as found in other species of the section (Gentry 1982). Plants from near Bahia San Carlos (Sonora) placed here in the protologue (Gentry 1972: 124, 125: upper photo) (Gentry & Arguelles 19881, Felger & Russell 11455) belong to A. colorata (Gentry 1982: 432–433, Starr 2012: 79). — The species is very rare, known only from the Sierra Jojoba and the Sierrita de López, and has apparently not been recollected since Gentry, but Kristen & Etter (photograph in Pilbeam (2013)) recently visited the habitat. Photographs from Nacapule Canyon labelled A. aff. fortiflora in Greulich (2017b) appear to represent A. chrysoglossa (on p. 40) and A. colorata (on p. 46). A. fourcroydes Lemaire (Ill. Hort. 11(Misc.): 65, 1864). Type: not typified. — Lit: Berger (1915: 238–240, with ills.); Gentry (1972: 138–139); Gentry (1982: 573–576, with ills.); Lott & García-Mendoza (1994: online version with ills.). Distr: Cultivated only; mainly E Mexico; locally naturalized in the Caribbean, Costa Rica, and the Mediterranean. I: Curtis’s Bot. Mag. 144: t. 8746, 1918; Heller (2006: 86);

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Sánchez Gullón (2013: 51); Hochstätter (2015: VII: 42). Incl. Agave rigida var. longifolia Engelmann (1875); incl. Agave ixtli var. elongata Baker (1877)  Agave rigida var. elongata (Baker) Baker (1881)  Agave elongata (Baker) A. Berger (1912) (nom. illeg., ICN Art. 53.1); incl. Agave longifolia hort. ex A. Berger (1915); incl. Agave ixtli hort. ex A. Berger (1915) (nom. illeg., ICN Art. 53.1); incl. Agave sullivanii Trelease (1920); incl. Agave fourcroydes var. espiculata L. H. Dewey (1929). [2g] Stems thick, 1–2
0.25 m; Ros large, suckering; L straight, linear, rigid, sword-like, thickly rounded at the base, acuminate, guttered, 120–180 (–250)
7–12 cm, grey-green, margins straight; marginal teeth slender, 3–6 mm, dark brown, regularly spaced, 20–35 mm apart; terminal Sp conical, stout, openly short-grooved above at the base, mostly 20–30 mm, dark brown, somewhat decurrent; Inf 4–7 m, paniculate, part-Inf 10–18 in the upper , freely bulbilliferous; Fl (50–) 60–70 mm; Tep greenish-yellow, tube urceolate, thick-walled, grooved, 12–16
12–15 mm, lobes linear, at first erect then sharply reflexing, closely overlapping, subequal, 16–18
3–5 mm, outer with flat outer face, inner keeled; Fil stout, tapered towards the apex, 45–60 mm, inserted 6–8 mm above the tube bottom; Anth slender, excentric, 20–24 mm, pale yellow; Ov fusiform, roundly-trigonous, tapered towards the base, (20–) 30–40 mm, neck briefly constricted; Sty stout, clearly longer than the tepals; Sti capitate, 3-lobate; Fr and Se normally not produced, Fr elongate-clavate, obtusely triangular, 3-sulcate, shortly stipitate, acutely rostrate; Se nearly crescent-shaped, 9–10 mm. — Cytology: 2n = 60, 90, 138, 140, 150 (Doughty 1936, Castorena Sanchez & al. 1991, Lv & al. 2009, Simpson & al. 2011), in cultivars 2n = 90, 150 (Robert & al. 2008). Widely cultivated for fibre (“Henequén”) esp. in E Mexico and Yucatán (Gentry 1982), but also used to produce an alcoholic beverage similar to mescal marketed under the registered trademark “sisal” (Larqué Saavedra 2007). The origin of A. fourcroydes is in Yucatán according to Lott & García-Mendoza (1994). The different cultivars subsumed under A. fourcroydes show different

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degrees of similarity with its wild progenitor A. angustifolia. ‘Sac Ki’ and ‘Yaax Ki’, cultivated for cordage, show four syndromes of domestication: gigantism, greater fibrosity, less pronounced thorniness, and less reproductive capacity (ColungaGarcía Marín & al. 1996, Colunga-García Marín & May-Pat 1997). The major differences of A. fourcroydes from its wild progenitor A. angustifolia (thicker and longer stems, longer and broader leaves, and its sterility) reflect these domestication syndromes. Morphological data (Colunga-García Marín & al. 1999) and combined morphological/molecular isozyme data (Colunga-García Marín 2003) indicated A. fourcroydes to represent a polyphyletic assemblage of different cultivars independently derived from within the variable A. angustifolia. However, with molecular AFLP data, several cultivars as well as variants of the progenitor species A. angustifolia (see there) cluster independently, so that Infante (2006) suggest the need for a revised classification. A. fourcroydes normally does not produce seeds and is only asexually propagated via rhizomes for plantation use, according to most authors. The sole descriptions of fruit and seed are given by Berger (1915) and Berger (1918). Offsets of a given mother plant are genetically variable so that selection of special genotypes for the improvement of Henequén is possible (Infante & al. 2003). Locally naturalized in the Caribbean (Cayman Islands, Cuba, Dominican Republic, Haiti, Leeward Islands), Costa Rica, Italy (Govaerts 2014+), Spain (Sánchez Gullón 2013), the Balearic Islands (Sáez & al. 2016) and the Canary Islands (Acebes Ginovés & al. 2010). A. franzosinii (Sprenger) Sewell (Gard. Chron., ser. 3, 6: 639, 1889). Type: [neo — icono]: Curtis’s Bot. Mag. 136: t. 8317, 1910. — Lit: Baker (1892: with ills.); Berger (1915: 157–160, with ills.); Gentry (1982: 291–292, with ills.); Guillot Ortiz & al. (2008: 50–52); Thiede (2017b). Distr: Known from cultivation only; locally naturalized in Spain. I: Irish & Irish (2000: t. 17); Lyons (2002: 162); Heller (2006: 26, 87, 157, 159); Smith & Figueiredo (2008: 57); Richter (2011: 41, 70); Pilbeam (2013: 86);

J. Thiede

Hochstätter (2015: VII: 82, as A. beaulueriana); Mesquida & al. (2016: 48–49); Moore (2016: 209, 233, 234).  Agave americana var. franzosinii Sprenger (1885); incl. Agave beaulueriana Jacobi (1869). [2a] Ros very large, 2–2.7 (–3)
to 4.5 m, freely suckering; L to 40, lanceolate, widely spreading, recurved or sharply reflexed, narrowed at the base, thickened and convex below towards the base, guttered above, somewhat asperous, 180–220
22–35 cm, light glaucous-grey or bluish-glaucous variously marked with green below the middle of the lamina, margins straight to repand; larger marginal teeth (middle of the lamina) 8–10 mm, dark brown, remote, on fleshy prominences, sometimes with smaller intermittent teeth; terminal Sp with short open groove above, 30–60 mm, dark brown, decurrent along the inrolled leaf tip; Inf 8–11.4 m, paniculate, peduncle strong, fertile part broadly cylindrical, to 2.9 m broad, part-Inf broadly spreading, several times compound; Fl large, 83–100 mm; Tep yellow, soon withering, tube 18–22 mm, lobes 30–32 mm, outer linear, inner keeled on the back, grooved, with 2 ribs within; Fil stout, 65–80 mm, yellow, inserted at mid-tube; Anth 38–40 mm, yellow; Ov 35–45
10–13 mm, light bright green, neck slightly narrower; Sty stout, to 120 mm; Sti clavate, 3-lobed; Fr elongate-clavate, 55–70 mm; Se to 12
8–9 mm, black, shiny. A very distinctive species not easily confused with, but obviously related to, A. americana (Gentry 1982), although it might be viewed as a larger version of the latter with greyish to glaucous leaves. The name is commonly ascribed to Baker (Bull. Misc. Inform. Kew 1892: 3, 1892), but was first mentioned by della Valle di Casanova (Garden [London, 1871–1927] 11: 63 (1877)) at the eponymous Villa Franzosini at the Lago Maggiore (Italy). The epithet was first published at varietal rank by Sprenger (in Deutsche Gärtn.-Zeitung 9: 130, 1885), which has been overlooked so far. Howard (in Howard & Thompson-Mills (1979: 489)) first established that A. franzosinii is antedated by A. beaulueriana. To conserve the established A. franzosinii against A. beaulueriana, a formal proposal to conserve was published, together with notes on typification, by Thiede

Agave AGAVACEAE

(2017b). The species is naturalized in Spain (Guillot Ortiz & al. 2008, Sáez & al. 2014). In the USA, two different “forms” are cultivated under this name: The typical form (e.g., at Lotusland), which is also propagated by tissue culture (Gomez Rhine 2009), and a clearly different, much smaller plant with stiffer leaves with more closely spaced teeth and a different colour, which is referable to A.
winteriana (see there). A. funkiana K. Koch & C. D. Bouché (Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 3: 47, 1860). Type [neo]: Mexico, Hidalgo (Gentry 12273 [[neo — syntypes]: ARIZ, MEXU, US]). — Lit: Berger (1915: 91); Gentry (1982: 139–141, with ills.); Guillot Ortiz & al. (2008: 51, 54, with ills.). Distr: Mexico (C Nuevo León, W & SW Tamaulipas, San Luis Potosí, N Querétaro, Hidalgo, SW México); rocky hillsides and slopes, in oak forests and mixed woodlands, 250–1800 m; flowers in August. I: Heller (2006: 88); Pilbeam (2013: 87); Hochstätter (2015: VII: 82). [1g] Acaulescent; Ros open, 0.6–0.9
1.2–1.8 m, freely suckering; L linear, radiating, firm, straight or somewhat falcate, patulous, base broadly clasping, convexly thickened below, concave above, mostly 60–80
3.5–5.5 cm, yellowish-green to dark green, frequently with pale median stripe, margins horny, nearly straight, firm, thin, brown to grey; marginal teeth mostly directed downwards, regular, slender, 3–5 mm, 10–25 mm apart, with a few small irregularly arranged intermittent teeth; terminal Sp conicalsubulate, with a narrow to open groove above, 10–30 mm, brown to white; Inf slender, 3.5–4.5 m, ‘spicate’, peduncle glaucous-grey, the small Bra apparently caducous, fertile part laxly flowered in the upper of the inflorescence; Ped 10 mm; Fl geminate, 40–45 mm; Tep pale glaucous-green, tube conspicuously grooved and knobbly-angled, 3.5–4 mm, lobes linear, mucronate with a small hood, at first spreading to ascending, 18–19 mm, appressed and clasping the filaments after anthesis; Fil 30–35 mm, red or pink, inserted on the tube rim; Anth 20 mm, yellow with pink flush towards the tips; Ov oblong-fusiform, 20–24 mm, neck grooved,

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constricted; Sty somewhat longer than the tepals, red; Sti capitate; Fr transversely wavy-ridged, strong-walled, 25–30
15 mm, light pruinosegrey, abruptly apiculate; Se thick, shape very variable, 5–6
3.5–4.5 mm, wavy-lined on the faces, hilar notch large. The neotype cited above was designated by Gentry (1982: 140). Obviously related to A. lophantha, but differing in its larger size, regular and linear slightly concave leaves with nearly straight fine margins, and numerous regular fine teeth. The species is not well known in habitat, and the few individuals observed show little variation (Gentry 1982: 140). The specimens cited by Gentry (1982: 189) from Chiapas (Gentry 12195, DES, MEXU) belong to A. ghiesbreghtii (Lott & García-Mendoza 1994). The following selection of specimens represent new state records: G. B. Hinton & al. 18722 (MO) for Nuevo León, Hernández & Martínez 4298 (MEXU) for Querétaro, García Mendoza 4393 (MEXU) for México State, and Gentry & al. 20095 (MEXU) for San Luis Potosí (Portal Datos Abiertos UNAM 2018+). Guillot Ortiz & al. (2008) report the species as naturalized in Spain. — ‘Fatal Attraction’ and ‘Blue Haze’ are horticultural selections, and ‘Hakuro Shiro Fukurin’ is a variegated cultivar with leaves with white margins. A. fusca (Ravenna) Thiede & Eggli (Kakt. and. Sukk. 52(6): 166, 2001). Type: Guatemala (Ravenna 325 [Herb. Ravenna]). — Lit: Ravenna (1987: with ill.); Castañeda Rojas & al. (2005: 70–71); Castillejos-Cruz (2009: [133]–[138], with ills.); all as Manfreda. Distr: Guatemala (Chimaltenango, Comalapa); sandy plains and embankments, in pine-oak forests, 1920–2410 m; flowers June to July. I: Lott & García-Mendoza (1994: online version); Hochstätter (2016: I: 19); both as Manfreda.  Manfreda fusca Ravenna (1987). [3a3] Plants herbaceous; corm 3–9
2–5 cm, subcylindrical, bulb 3–4.5
1.5–3 cm, shortly cylindrical to ovoid, nearly completely covered with dry leaf bases 4–12 cm long, these membranous at the base and fibrous above; R fleshy and fibrous, contractile; L drought-deciduous, 5–15, 30–60
(1–) 2.2–3.2 cm, light green, prostrate or

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semi-erect, linear-lanceolate, narrowed towards the base, semisucculent, canaliculate, lower face carinate to slightly scabrous, upper face glabrous, apex acute-acuminate, margins entire, with a papillose-denticulate hyaline band; Inf 77–200 cm, ‘spicate’, peduncle stiff, Bra 8–12, rather distant, fertile part 5–25 cm, with 7–21 flowers; floral Bra 0.2–1.1
0.3–0.6 cm; Fl 37–47 mm, solitary, sessile, ascending, with foetid odour; Tep glaucous-green to reddish on the outer face, semisucculent, forming a tube 14–24
5–6 mm, lobes narrowly oblong to lanceolate, extended to revolute, 15–17
4–6 mm, tip apiculate, with a bundle of short white hairs; Fil sparsely glandularpilose, 39–42 mm, erect at anthesis, exceeding the tube for 10–20 mm, inserted at mid-tube all on the same level, yellowish-green with purple dots; Anth 10–11 mm, yellow; Ov oblong, 12–18
(5–) 6–6.6 mm, greenish, prolonging 1 mm into the tube, not constricted above; Sty 37–57 mm, yellowish-green with purple dots; Sti capitately 3-lobed; Fr subglobose, 15–21
9–15 mm, with a scar near the apex, tepals persistent; Se deltoid, plane-concave, with narrow margin, 4–5
3–4.5 mm, shiny black. In the protologue, no differential diagnosis is given nor is the relationship indicated. GarcíaMendoza & al. (2000) clearly place the species in Ser. Guttatae (as A. guttata Group) of Manfreda Sect. Manfreda, based on a study of additional specimens at MEXU. This placement is confirmed by Castañeda Rojas & al. (2005) and CastillejosCruz (2009) (both as Manfreda), who provide additional morphological data. A. fusca (presently known from the inaccessible holotype and 2 further gatherings only) is distinct esp. in its flower colour and foetid odour. The species was erroneously omitted from the treatment of the family for the ‘Flora Mesoamericana’ (Lott & García-Mendoza 1994), but the online-version includes photographs. A. galvaniae (Castañeda & al.) Etter & Kristen (Haseltonia 12: 70, 2007). Type: Mexico, México (Castan˜ eda Rojas & al. 108 [MEXU, ENCB, IEB, MEXU, MO]). — Lit: Castañeda Rojas & al. (2005); Castillejos-Cruz (2009: [139]–[144]); both with ills. and as Manfreda. Distr: Mexico

J. Thiede

(S México); in disturbed tropical deciduous forests and ecotone with pine forests, 1450–1900 m; flowers in July. I: Hochstätter (2016: I: 20, as Manfreda).  Manfreda galvaniae A. Castañeda & al. (2005). [3a3] Plants herbaceous; corm 3–5
2–4 cm, subglobose to subcylindrical, bulb 2.7–4
1.3–2.5 cm, cylindrical, nearly completely covered with dried leaf bases 4.5–7.3 cm long, these membranous at the base and fibrous above; R fleshy, with filiform ramifications, contractile; L drought-deciduous, 2–5 (–7), prostrate or semierect, linear-lanceolate, 22–62
0.6–2.6 cm, light green, narrowed towards the base to a pseudopetiole 3–4
1 cm, semisucculent, slightly canaliculate, glabrous on both faces, apex acuminate, margins entire, with a hyaline band not visible to the naked eye; Inf 77–154 cm, ‘spicate’, erect, somewhat arched, reddish to greenish with purple tinge, peduncular Bra 5–8, similar to the leaves but smaller, fertile part 23–67 cm, lax, with 7–21 flowers; floral Bra 0.8–3.1
0.3–1 cm; Fl 27–42 mm, solitary, sessile, ascending; Tep reddish with purple tinge, semisucculent, forming a tube 13–24
4–5 mm, lobes narrowly elliptic to ovate, extended, 9–14
3–7 mm, tip cucullate, with a bundle of short white trichomes; Fil 15–24 mm, erect at anthesis, exceeding the tube for 10–20 mm, inserted at the upper of the tube all on the same level, yellowish-green; Anth 8–12 mm, yellow; Ov cylindrical, (5–) 8–12
2–4 (–5) mm, reddish with purple tinge, prolonging 1 mm into the tube, not constricted above; Sty 28–37 mm, whitish with purple tinge to yellowish-green; Sti capitately 3-lobed; Fr subglobose, 13–20
9–15 mm, with a scar near the apex, tepals persistent; Se deltoid, plane-concave, with narrow margin, 3–5
2–4 mm, shiny black. In the protologue placed in Ser. Guttatae (as A. guttata Group) of Sect. Herbaceae, and characterized by its globose corm nearly always completely covered with dry leaf bases, linearlanceolate pseudopetiolate leaves, short and lax inflorescences with reddish to yellowish-green with purple tinge, narrowly elliptic to ovate perianth lobes, and a shortly projecting ovary. It is morphologically closest to A. fusca and A. debilis

Agave AGAVACEAE

(= Manfreda pringlei), but differs in its hyaline leaf margin, the lax flower arrangement, the semisucculence and colour of its flowers, the shape and orientation of its perianth lobes, and its short filaments. A. garciae-mendozae Galván & L. Hernández (Cact. Succ. J. (US) 74(4): 188, ills. (pp. 189–190), 2002). Type: Mexico, Hidalgo (Galván & Galván 4646 [ENCB, CHAPA, MEXU, MO, QMEX, UAMIZ]). — Lit: Eguiarte & Scheinvar (2008: 35–39, with ills.); Trejo-Salazar & al. (2015: pollination). Distr: Mexico (San Luis Potosí, Querétaro, Hidalgo); mountain slopes and rocky outcrops, often in canyon bottoms, in limestone soils, in sclerophyllous scrub or oak/ pine-oak forests, 1800–2650 m; flowers June to August. I: Pilbeam (2013: 88); Hochstätter (2015: IX: 31). [1g] Stems short; Ros openly spreading, 0.75–1.15 (–1.3)
1–1.75 m, single or somewhat caespitose; L (30–) 50–80, lanceolate, straight or incurved, fleshy, thin, flexible mainly in the upper , flat or slightly convex below, flat to somewhat concave above, (33–) 48–78
6–13 cm, dark green to glaucous-green, margins horny, continuous, 0.5–2 mm wide, grey to dark reddish-brown, easily separated; marginal teeth variable, straight to curved, narrowly acuminate, deltoid, flattened, 7–14 mm, grey with reddish-brown tip, (7–) 10–40 mm apart, often with intermittent smaller teeth; terminal Sp conical to subulate, sometimes somewhat reflexed, widely grooved on the upper face, keeled abaxially, 27–48 mm, grey, decurrent; Inf 3.2–5.5 m, ‘spicate’, peduncle reddish to glaucous-green, with long-deltoid, chartaceous to papery Bra 7.5–23 cm long, fertile part in the upper of the inflorescence; Ped 2–3 mm; Fl geminate, 28–39 mm; Tep light green to yellowishgreen and somewhat glaucous, tube 1.5–2.5
5–6 (–7) mm, lobes oblong, involute, acute, slightly cucullate and hairy, 12.5–16
4–5 mm, sometimes reddish, inner with a wide keel; Fil 38–45 mm, inserted at the base of the lobes; Anth 10–13 mm, yellow or greenish-yellow; Ov cylindrical, 8–12
3.5 mm, neck constricted, 2–3
1.2–1.8 mm; Sty slightly longer than the filaments; Fr oblong or oblong-ellipsoid, (15–)

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17–20
9–11 mm, shortly apiculate; Se semicircular, black, 3–4.9
2–2.7 mm, marginal wing 0.8–1 mm wide, hilar notch shallow. Showing affinities with A. horrida and A. kerchovei according to the protologue, and thus placed in Sect. Heteracanthae (as Group Marginatae). It differs from the former in its open rosettes, the colour and larger size of its flexible leaves with a grooved terminal spine, its shorter ovary and more shallow tube; and from the latter in its shorter rosettes, fewer leaves with a different colour and flexibility, its shorter ovary and shallower tube. The inflorescences of A. garciae-mendozae consist on average of 3623 flowers, of which 426 are producing pollen or nectar at any one day. Only 60% of the flowers have a high probability to develop into fruits. From the total seeds per infructescence (471,120 on average), only (142,725 on average) are viable. The plants are self-compatible, but cross-pollination is most effective for fruit set, indicating a strong inbreeding depression. Most flower visitors are nectarivorous bats, honey bees and bumblebees are infrequent, and hawkmoths rare. The species appears to have a generalist pollination system with bats as main pollinators, and bees and bumblebees ensuring reproductive success when bats are missing. Relatively high levels of genetic diversity and small genetic distances between different populations have been found (González 2004, Eguiarte & Scheinvar 2008, Eguiarte & al. 2013: 491, Trejo-Salazar & al. 2015). The unpublished A. hidalgensis (also as “hidalguensis”) (Eguiarte & Scheinvar 2008, Arzate García 2009, both with ills.), one of the largest representatives of Subgen. Littaea (rosettes to 2.5 m ∅, leaves 70–90 cm, inflorescences exceeding 7 m) from the N Barranca de Metztitlán, appears to belong here. A. garciaruizii A. Vázquez & al. (Phytotaxa [in press], ills., 2019). Type: Mexico, Michoacán (Vázquez-García & al. 10140 [IBUG, CIMI, MEXU]). — Distr: Mexico (SE Jalisco, N Michoacán); rocky cliffs of the ItzícuaroApupátaro watershed, in tropical deciduous forests, 900–1000 m; flowers in March. I: Vázquez-

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García & al. (2007b: 41–42, t. B1-B4, as A. angustiarum). [1g] Subcaulescent; Ros open, 0.9–1.1 m, solitary; L 20–34, linear to lanceolate, openly spreading, straight, thick, firm, long-acuminate, flat to concave above, convex below, 50–80
6–10.6 cm, green to dark green, pale green median stripe above inconspicuous or lacking, margins whitish-grey, continuous; marginal teeth straight or inclined in the lower part of the leaf, curved upwards or bent in the upper part, flattened, the largest somewhat scattered, 4–7 mm, dark brown to grey, (3–) 10–30 mm apart, 8–14 teeth per 10 cm, teeth absent from the topmost 5–10% of the leaf, intervening margin nearly straight; terminal Sp acicular, narrowly grooved above, 30–40 mm; Inf 3–4.2 m, ‘spicate’, flexible, peduncle with toothless triangular Bra 10.5–13.5 cm long, fertile part in the upper 33–40%, 15–18 cm ∅ (infructescence 10–14 cm ∅); Ped 8–10 mm (fruiting 18–20 mm); Fl 57–64 mm, geminate; Tep pale yellow to yellowish-green, tube very short, 2–3
6–7 mm, lobes 17–18
4–5 mm, pale yellow to yellow, glaucous-greenish in bud; Fil 38–44 mm, yellow; Anth 16–21 mm, yellow; Ov 16–17 mm, neck narrow; Sty slightly longer than the filaments; Sti capitate; Fr 24–26
10–12 mm, fruit valves curved outwards, apically acute; Se crescent-shaped, 2–3
3–5 mm. A local endemic, according to the protologue closest to A. angustiarum and differing esp. in having an inconspicuous or absent median stripe on the upper face of the leaves, denser marginal teeth, longer anthers, longer fruiting pedicels, a larger infructescence diameter, and larger capsules with acute fruit valves curved outwards. A. garciaruizii is also close to A. arcedianoensis and differs esp. in having an inconspicuous or absent median stripe on the upper leaf face, denser marginal teeth, a shorter fertile inflorescence part, pale yellow to yellowish-green flowers with larger anthers and longer pedicels, and smaller fruits with apically acute, spreading fruit valves. The records and specimens for A. angustiarum from SE Jalisco (Hernández-Vera & al. 2007a) and from SE Jalisco and N Michoacán (VázquezGarcía & al. 2007b) belong here. The fruit valves

J. Thiede

curved outwards are reported only for A. garciaruizii and A. impressa. A. geminiflora (Tagliabue) Ker Gawler (J. Sci. Arts (London) 2: 86–90, 1817). Type: [icono]: Bibliot. Ital. Giorn. Lett. 1: t. inter pp. 112, 1816. — Lit: Berger (1915: 69–70); Gentry (1968); Gentry (1982: 113–115); McVaugh (1989); Ullrich (1995a); Cházaro Basáñez & al. (2004b); Vázquez-García & al. (2007b: 52–53, t. J); Starr (2012: 94–97, 317–319); all with ills. except Berger. Distr: Mexico (C Nayarit, N-C Jalisco); on rocks along arroyos, in oak woodland, 1000–1400 m; flowers October to December. I: Bot. Reg. 14: t. 1145, 1828; Irish & Irish (2000: t. 18); Guillot Ortiz & Meer (2008c); Richter (2011: 28, 29); Etter & Kristen (2012: 94); Pilbeam (2013: 89–90).  Littaea geminiflora Tagliabue (1816); incl. Bonapartea juncea Haworth (1812) (nom. illeg., ICN Art. 53.1); incl. Yucca boscii Hort. Panorm. ex Hornemann (1813) (nomen rejiciendum, Art. 56.1)  Dracaena boscii (Hornemann) J. Zeyher (1818)  Agave boscii (Hornemann) Anonymous (2013) (nom. inval., ICN Art. 29.1, 34.1b); incl. Bonapartea juncea Willdenow (1814) (nom. illeg., ICN Art. 53.1); incl. Yucca boscii Desfontaines (1815) (nom. illeg., ICN Art. 53.1); incl. Dracaena filamentosa Scannagatta ex Ker-Gawler (1817) (nom. inval., ICN Art. 36.1c); incl. Bonapartea flagelliformis Henckel (1820) (incorrect name, ICN Art. 11.4); incl. Tillandsia juncea Willdenow ex Steudel (1841) (nom. illeg., ICN Art. 53.1); incl. Agave geminiflora var. filamentosa Hooker (1856); incl. Littaea juncea É. Morren (1866); incl. Agave angustissima Engelmann (1875); incl. Agave geminiflora var. filifera A. Terracciano (1885); incl. Agave knightiana J. R. Drummond (1909); incl. Dracaena boscii Hort. J. F. Cels ex A. Berger (1915) (nom. illeg., ICN Art. 53.1); incl. Agave geminiflora var. stricta-viridis Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a). [1d] Stems short; Ros dense, 0.7–1 m, somewhat broader than tall, solitary; L many, linear, eventually arching, flexible, narrow, pliable, abruptly acute, roundly convex below and above, smooth, 45–60
0.6–0.8 cm, green,

Agave AGAVACEAE

margins finely filiferous or rarely naked; terminal Sp shortly subulate, 5–7 mm, greyish; Inf 4–7 m, ‘spicate’, peduncle long-tapering, stout at the base (9–12 cm ∅), with narrow ascending-reflexed Bra, fertile part in the upper of the inflorescence, part-Inf with mostly geminate flowers; Ped slender, 5–8 mm; Fl 40–52 mm; Tep pale yellowish-greenish below, tinged above with red or purple, tube narrowly funnel-shaped, 6-grooved, 6–11 mm, lobes linear-lanceolate, recurving, slightly unequal, 18–21
4–5 mm, tinged with red, inner slightly shorter, distinctly keeled; Fil 35–46 mm, colourless or reddish, inserted on the tube rim; Anth excentric, 20–22 mm; Ov slender, fusiform-angled, 16–20 mm, neck grooved; Sty thickened above, slightly shorter than the filaments; Sti not thickened; Fr oblong, trigonous, tough, persistent, 18–20
9–10 mm, shortly stipitate, shortly beaked, reddish when young; Se crescent-shaped, thick, irregularly veined on the faces, 3–4
2–3 mm, with conspicuous marginal wing. Distinct within Sect. Littaea (as Group Filiferae) by its relatively large simple stem with innumerable, very narrow, pliable, smooth leaves and large ‘spikes’ with relatively remote long flowers. It is most closely related to A. ornithobroma which differs esp. in its asymmetrical, few-leaved, surculose rosettes (Gentry 1982: 112). The species was of unknown wild origin until Gentry (1968) reported it from a small area N of Ocotillo, Nayarit. Recently, 2 further populations in Nayarit (Sierra de Álica, La Galinda) and a first occurrence in Jalisco at Xiloasco were found (Cházaro Basáñez & al. 2004b, Hernández-Vera & al. 2007a, Vázquez-García & al. 2007a). The report of a specimen of this taxon from S Sinaloa by McVaugh (1989: 135) is erroneous, since the collection cited (Gentry 18358) is the type collection of A. ornithobroma. The plant came in 1797 from the Botanical Garden Bologna to Milano (Italy), where it first flowered in the garden of Duke Litta of Lainate in 1815, after whom it was named in a new genus as Littaea geminiflora (Berger 1915) (see also the historical account by Richter 2011: 24–32). The epithet of A. geminiflora (dating from 1816) is antedated by the name Yucca boscii (1813),

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which would therefore be the correct epithet for the species (Govaerts 2014+). A proposal to conserve the long-established name A. geminiflora was consequently published by Figueiredo & Smith (2013b) and was recommended for acceptance (Applequist 2014: 1366). A. gentryi B. Ullrich (Succulenta 69(10): 210–214, ills., 1990). Type: Mexico, Nuevo León (Gentry 20159 [ARIZ, US]). — Lit: Gentry (1982: 598–601, figs. 21.8, 21.9, as A. macroculmis); Villarreal (1996); González-Elizondo & al. (2009: 80–84, with ills.); Starr (2012: 98–105, 325–328, with ills.); Greulich (2012c: with ills.). Distr: Mexico (Coahuila, Nuevo León, Tamaulipas, Durango, Zacatecas, San Luis Potosí, Guanajuato (?), Querétaro, Hidalgo, Puebla); mountain regions in open pine-oak forests, 1800–3400 m; flowers in April. I: Heller (2006: 89); Spracklin (2007: 148); Richter (2011: 72); Pilbeam (2013: 91); Nájera Quezada & al. (2015: 5); Hochstätter (2015: III: 3); Greulich (2016b: 38). [2m] Ros 0.9–1.5 (–1.8)
(0.6–) 0.9–1.8 (–2.2) m, solitary or clustering with rhizomatous offsets; L 30–45, either broad from the base to above the middle before quickly tapering to the sharp tip, or long triangular and widest at the base before gradually narrowing to the tip, concave, 60–120
15–26 cm, light yellowish-green to deep rich green, with strong imprints from the central bud created by the marginal teeth; marginal teeth nearly straight, small to large, 8–12 mm, 20–40 mm apart; terminal Sp stout, 45–75 mm; Inf 2.5–5.5 m, paniculate, peduncle with overlapping, thick and succulent, entire Bra, fertile part elliptic, covered in a bulb-like manner by fleshy bracts before anthesis, Fl densely clustered in 10–18 (–21) short-stalked part-Inf; Fl 70–110 mm; Tep yellow, reddish in bud, tube funnel-shaped, 11–16 mm, lobes 22–28 mm; Ov 35–50 mm; Fr 60–75
20–25 mm; Se 7–8
6 mm. The usage of the name A. macroculmis by Gentry (1982: 598) must be rejected, since this name is a synonym of A. atrovirens (Ullrich 1990i). This left Gentry’s distinctive plant without a name, which led Ullrich (1990i) to describe it as a new species, A. gentryi, using Gentry’s

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J. Thiede

Fig. 24 Agave ghiesbreghtii. (Copyright: J. Etter & M. Kristen)

description of A. macroculmis. However, this description also includes features of the subsequently separated A. montana. The description given above for A. gentryi is based on a combination of data from Villarreal (1996), Starr (2012), and Greulich (2012c). See under A. montana for differences and the classification of both species. A. gentryi includes 3 distinct and geographically separated forms (Starr 2012). It moreover appears to form hybrids with A. americana ssp. protoamericana, A. asperrima, A. salmiana ssp. crassispina, A. mapisaga, and A. montana (Starr 2012, Greulich 2012c). González-Elizondo & al. (2009) report the species from Durango. The report of “A. macroculmis” from Guanajuato (Zamudio & Galván Villanueva 2011) may refer to A. gentryi. — ‘Jaws’ is a cultivar with exceptionally large teeth (Starr 2012: 105). Greulich (2012c) depicts flowering specimens of it and considers ‘Jaws’ to be a long-leaved transitional form to A. salmiana with longer stalked and compound part-inflorescences. ‘La Pena Colorada’ is a variegated cultivar (Starr 2014: 222, with ill.). The flowers are visited at day by birds and insects and at night, when nectar production and sugar content increases, by bats and insects (Castillo-Hernández & Treviño-Carreón 2014). The maroon-fronted parrot (Rhynchopsitta terrisi) feeds on flowers and fruits of the species during the breeding season (Ortiz-Maciel & al. 2010).

A. ghiesbreghtii Verschaffelt (Cat., 71: 3, 1862). Type [neo]: Cult. BG La Mortola (Berger s.n. [US [neo/syn?]]). — Lit: Berger (1915: 116–117, with ill.); Standley & Steyermark (1952: as A. huehueteca); Gentry (1982: 141–142, with ills.); Lott & García-Mendoza (1994: onlineversion with ills.); García-Mendoza (2011a: 26–27, 30). Distr: Mexico (México, Guerrero, Puebla, Oaxaca, Veracruz, Chiapas), W Guatemala; limestone rock slopes and outcrops in desert scrub, tropical deciduous forests, and pine-oak forests, 975–2500 m; flowers mainly June to September, also October, November and January. I: Heller (2006: 40, 90, 91, 156); Richter (2011: 48, as A. purpusiorum, 66, 113); Pilbeam (2013: 92). – Fig. 24. Incl. Agave rohanii Jacobi (1864); incl. Agave leguayana Verschaffelt (1868); incl. Agave gilbeyi Hort. Haage & Schmidt (1873)  Agave horrida var. gilbeyi (Hort. Haage & Schmidt) Baker (1877)  Agave roezliana var. gilbeyi (Hort. Haage & Schmidt) Trelease (1914); incl. Agave roezliana Baker (1877); incl. Agave roezliana var. inghamii hort. ex Baker (1877); incl. Agave purpusorum A. Berger (1915); incl. Agave roezlii hort. ex A. Berger (1915) (nom. inval., ICN Art. 61.1); incl. Agave huehueteca Standley & Steyermark (1943). [1g] Acaulescent; Ros compact, 0.4–0.7
0.6–0.8 m, solitary or copiously suckering; L 70–90, broadly lanceolate, ovate or deltoid,

Agave AGAVACEAE

erect, straight or upcurving, thick, rigid, narrowed above the thick base, broadest at the base or in the middle, apex acuminate, convex below, flat to slightly concave or guttered above, smooth, (20–) 30–40 (–65)
(5–) 7–12 cm, or more rarely broadly linear and 35–38
5.5–6 cm, dark or light green to yellow-green, sometimes with a lighter mid-stripe, margins horny, relatively straight, narrow, continuous, brown or greyish; marginal teeth triangular, strong, frequently straight, sometimes curved upwards or downwards, 5–8 (–15)
3–6 (–7) mm, brown to greyish, (10–) 25–40 (–60) mm apart, with some intermittent small teeth, reduced at the leaf tip for 2.5–4.5 (–12) cm; terminal Sp conical or subulate, shortly shallowly grooved above, rounded below, (10–) 25–35 (–40)
4–6 (–10) mm, brown to grey, decurrent on the lower face for 2–4 mm; Inf (3–) 3.5–5 (–6) m, ‘spicate’, peduncle green with purple tinge, with linear papery brown Bra 13–17 cm long, fertile part in the upper , densely flowered; Ped geminate, 1–4 (fruiting 5–8) mm, thick; Fl campanulate, 30–40 (–50) mm; Tep purple to greenish-red, inside yellowish, tube broadly funnel-shaped, grooved, 2–5 (–10)
5–7 (–10) mm, lobes broadly linear to oblong, apex rounded, thickly apiculate, at first outcurving, later involute and clasping the filaments, subequal, 15–21
3–5 (–8) mm, inner narrower and with broad low keel; Fil (35–) 40–45 mm, inserted on the rim of the tube at the lobe bases, reddish; Anth centric, 10–15 (–22) mm, brownish to brownish-red; Ov cylindrical, 15–20
3–5 (–8) mm, green, neck 2–4 mm, constricted; Sty 45 to >70 mm, thickened below the stigma, brown or reddish; Sti capitate; Fr oblong, globose-ovoid or ellipsoid to obovoid, 17–23
8–15 mm, brownish; Se semicircular, finely punctiform, wing inconspicuous, (3–) 4–5
2–3.5 mm, shiny black. — Cytology: 2n = 120 (Simpson & al. 2011: p.p., as A. gilbeyi). The neotype cited above was designated by Gentry (1982: 142). A species of uncertain circumscription, closely related to A. kerchovei, but different by its shorter and broader leaves with fewer and smaller teeth on narrower and darker horny margins (Gentry 1982: 142, García-Mendoza 2011a). Characterized by its small rosettes, deltoid or broadly

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lanceolate leaves, narrow margins with few short and widely spaced teeth, short tepals, long filaments, and punctiform seeds (García-Mendoza 2011a). The species was introduced by Ghiesbreght and brought into the trade by Verschaffelt in 1862 (Berger 1915). The species (vernacular names: “pichomel”, “jabalí”) is used for living fences, and strips of dried leaves are used to make ties (GarcíaMendoza 2011a). Rivera-Hernández & al. (2015) provide a new record from W-C Veracruz. See also notes under A. angustiarum, A. funkiana, A. horrida ssp. horrida, and A. kerchovei. A. gigantensis Gentry (Occas. Pap. Calif. Acad. Sci. 130: 63–67, ills. (pp. 65–66 [bottom], 68, t. 2 [top]), 1978). Type [syn]: Mexico, Baja California Sur (Gentry & McGill 23320 [US, ARIZ, ASU, CAS, DES, MEXU, MICH, SD]). — Lit: Gentry (1982: 386–388, ills. pp. 387, 388 [bottom]); Webb & Starr (2014a: 94); Webb & Starr (2014b: 7–8); Webb & Starr (2015: 73); all with ills. Distr: Mexico (C Baja California Sur: N Sierra de la Giganta [Cerro El Potrero & Sierra de las Palmas]); mountain slopes, mesas and cliffs, 900–1520 m; flowers April to May and November. [2j] Acaulescent; Ros rather open, 0.5–1
0.8–1.2 m, solitary; L few, broadly lanceolate, flat, rigid, thick, fleshy, markedly narrowed at the base, widest at or above the middle, acuminate, smooth, 40–75
11–16 cm, dark green to occasionally glaucous grey-green, with distinct imprints from the central bud, turning red to purplish when plants are flowering, margins repand to prominently mammillate; marginal teeth variously curved, basis thick, sometimes grotesquely so, frequently 2–3 teeth on the same base, confluent along the upper leaf margins, 10–20 mm and more, brown to light greyish, generally remote and to 60–80 mm apart; terminal Sp strongly subulate, straight or sinuous, deeply sulcate above, 30–60 mm, grey, long decurrent as pronounced horny margin, sometimes to mid-leaf; Inf 4–5 m, paniculate, peduncle with narrowly lanceolate, thickly chartaceous Bra, fertile part rather stout, compact and narrow, part-Inf nearly horizontal, 18–25 in the upper of the inflorescence; Fl

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slender, 48–60 mm; Tep bright pale yellow, buds whitish or pale green, tube shortly discoid, spreading, thickened at the lobe bases, 4–5
11–13 mm, lobes linear, spreading, rounded and shortly hooded at the apex, 18–25
5–6 mm, inner with prominent keel, grooved within and strongly overlapped at the base by the outer; Fil slender, 30–45 mm, variable in length, inserted on the rim of the tube at the lobe bases; Anth somewhat excentric, 18–25 mm; Ov slender, fusiform, neck short, slightly grooved, constricted; Sty slightly longer than the filaments; Sti capitate; Fr oblong, 35–40
12–15 mm, neither beaked nor stipitate; Se unknown. — Cytology: 2n = 60 (Simpson & al. 2011). Gentry’s holotype indicated in the protologue for US consists of 3 sheets which are not crosslabelled, thus representing syntypes. Recent field and herbarium studies (Webb & Starr 2014a, b) showed that Gentry (1978) and Gentry (1982) included material with glaucous-blue to bluegreen leaves in fact belonging to A. sobria ssp. sobria in his original concept of A. gigantensis. Webb & Starr (2014b) restricted the concept of A. gigantensis to plants with dark-green leaves and occurring only at higher elevations on the Cerro El Potrero and Sierra de las Palmas. In the above description based on Gentry (1982), the leaf colour is corrected to dark green and some further data were added from Webb & Starr (2014b) and Webb & Starr (2015), but it may still include incorrect data from A. sobria. Some of Gentry’s photographs labelled “A. gigantensis” (1978: p. 66 top; 1982: p. 388 top) belong to A. sobria (Webb & Starr 2014b), and “A. gigantensis” plants with bluish leaves (Richter 2011: 79, 105, Pilbeam 2013: 93, Moore 2016: 277) apparently belong to A. sobria. According to Gentry (1982), the closest relatives appear to be A. avellanidens (with leaves not or scarcely narrowed towards the base) and A. moranii (with much longer leaves), both disjunct to the north. These three taxa as well as A. turneri are now placed in a new Sect. Conicae (see under A. avellanidens). A. gilbertii A. Berger (Monatsschr. Kakt.kunde 14: 126, 1904). Type [lecto]: Ex cult.

J. Thiede

(Anonymous s.n. [K [sub A. bakeri]]). — Lit: Watson (1902: protologue A. bakeri); Hooker (1903: as A. bakeri); Berger (1915: 41–42); Gentry (1982: 71–72); Roosbroeck (2004); all with ills. Distr: Known from cultivation only. I: Heller (2006: 93); Richter (2011: 60, as A. bakeri); Hochstätter (2015: VIII: 21); Meer (2016c: 144). Incl. Agave bakeri W. Watson (1902) (nom. illeg., ICN Art. 53.1). [1b] Stems short; Ros solitary, not suckering; L many, lanceolate, recurving, coriaceous, narrowed and thickened near the base, broadest in the middle, tapering to the base to 6.25 cm, tapering to the apex to a sharp point, convex below, concave to flat above, 90–100
10–12 cm, glaucous-green, with paler median stripe, margins thin, with continuous narrow brown border; marginal teeth none; terminal Sp slender, 5–20 mm, brown; Inf to 3 m, ‘spicate’, peduncle short, with linear-lanceolate Bra longer than the lowest flowers, fertile part cylindrical, densely flowered from near the base; Fl 50–60 mm, geminate; Tep pale greenish-yellow outside, whitish within, tube 6-furrowed, 11–12 mm, lobes reflexed or revolute, obtuse, 20 mm, greenish outside, whitish within, inner broader, with dark green keel; Fil rigid, 40–50 mm, whitish, inserted at the orifice of the tube 1 mm above the tube base; Anth 14 mm, golden-yellow; Ov fusiform, slender, 20 mm, neck slender, 6-sulcate; Sty rigid, shorter than the filaments but longer at the female phase (Watson 1902); Sti 3-lobed; Fr and Se unknown (although Berger (1904) reported that the plant which flowered at Kew in 1902 yielded many seeds, which were widely distributed). The illegitimate name and replaced synonym A. bakeri is usually ascribed to Hooker (1903), but was published earlier by W. Watson (Watson 1902), based on a flowering plant at Kew purchased at the sale of the Agave collection of the late Mr. Peacock in 1889. The notion of Gentry (1982) that the species “disappeared in cultivation” is erroneous; it is cultivated in Europe (see references) and the USA, but has never been located in habitat (Gentry 1982). The recently published A. chazaroi is somewhat similar, but stemless and with broader leaves with a longer terminal spine.

Agave AGAVACEAE

A.
glomeruliflora (Engelmann) A. Berger pro sp. (Hort. Mortol. 12, 1912). Type: [lecto — icono]: Gard. Chron. ser. nov., 1883: 19, fig. 6. — Lit: Berger (1915: 94–95, as A. glomeruliflora); Muller (1939: with ill., as A. chisoensis); Gentry (1982: 142–144, with ills., as A. glomeruliflora); Reveal & Hodgson (2002: as A. glomeruliflora); Greulich (2016a: with many ills.). Distr: USA (W Texas), Mexico (Coahuila); gravelly calcareous slopes in grasslands and oak-juniper woodlands, 600–1600 m; flowers mid-spring to early fall. I: Kümmel & Klügling (2006: as A. lechuguilla); Richter (2011: 86); Pilbeam (2013: 94–95); Hochstätter (2015: IX: 47).  Agave heteracantha var. glomeruliflora Engelmann (1883)  Agave lechuguilla fa. glomeruliflora (Engelmann) Trelease (1920); incl. Agave chisosensis C. H. Muller (1939). [1g
2l] Obviously of intersubgeneric hybrid origin with morphological gradations between A. lechuguilla and A.
gracilipes, A. (parryi ssp.) neomexicana and/or A. havardiana (Gentry 1982: 143) and thus only representing an aggregate of different habitually similar natural hybrids. According to Reveal & Hodgson (2002), the name A.
glomeruliflora “should probably be more appropriately applied only to crosses and back-crosses between A. lechuguilla and A. havardiana”. Gentry’s selection of a neotype is superseded by his simultaneous selection of a lectotype (Gentry 1982: 143). Baker & al. (2009: 283) report a chromosome count of n = 60 which is congruent with a spontaneous origin between A. lechuguilla
A. parryi ssp. neomexicana, both with 2n = 120; counts for the remaining two possible parental taxa are unknown. Few if any substantial threats were identified, and substantial populations in Big Bend and Guadalupe Mountains National Parks are under protection (Reichenbacher 1985: 103–105). Magallán Hernández & Hernández Sandoval (2001: 108) report “A. aff. glomeruliflora” from Municipio Cadereyta in Coahuila. Plants have survived 30 years outdoors in a mild climate in Germany with rain protection in winter (Kümmel & Klügling 2006: as A. lechuguilla, Greulich 2016a).

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A. gomezpompae Cházaro & Jimeno-Sevilla (Cact.-Avent. Int. 88: 2–11, ills., 2010). Type: Mexico, Veracruz (Jimeno-Sevilla & al. 247 [XAL, CHAP, CHAPA, ENCB, IBUG, IEB, MEXU]). — Lit: Jimeno-Sevilla (2010: 97–99, with ills.). Distr: Mexico (C Veracruz: near Córdoba); karstic limestone rocks and outcrops, 850 & 1400 m, in tropical “subperennial” (semideciduous) or cloud forests; flowers May to July; only known from 4 populations. I: Richter (2011: 56); Meer (2012); Pilbeam (2013: 96); Hochstätter (2015: VIII: 21). [1f] Caulescent, stems creeping, to >150
8–15 cm, bifurcating 3
or more, each stem with a repeatedly flowering Ros that turns upright, not surculose; L few, lanceolate to oblanceolate, flexible, fleshy, concave above, convex below, 80–85
10–13 cm, dark green, margins denticulate throughout; marginal teeth none; terminal Sp 20 mm or longer, dark; Inf lateral according to the protologue but actually likely terminal on rudimentary lateral rosettes, erect, to 2.7 m, ‘spicate’, peduncle with triangular Bra 12–16
3–5 mm, fertile part only 50 cm; Ped 3–4 mm, single or geminate; Fl 32–49 mm (74 mm with stamens); Tep spatulate, 20–28
2–4 (at the base) mm, green or creamish-yellow inside, tube  as long as the obtuse lobes; Fil to 43 mm; Anth 7–9 mm, yellowish; Ov ovoid or barrel-shaped, 15–21
3–5 mm, greenish, not constricted; Sty slender, to 43 mm; Sti slightly thickened; Fr trigonous, 20 mm with a 5 mm long stipe, blackish; Se 3–4
2 mm, black, shiny. Placed in Sect. Micracanthae (as Group Polycephalae) according to the protologue, and closest to the allopatric A. pendula with which it shares long and bifurcating stems and seemingly lateral inflorescences, but differs by thicker stems, broader and pure green leaves without yellow mid-stripe and denticulate all along the margins, and longer inflorescences. A.
gonzaloi D. Guillot & P. Van der Meer pro sp. (Flora Montiber. 27: 55–56, 2004). Type: Spain, Valencia (Guillot & Van der Meer s.n. [VAL 151000]). — Distr: Cultivated only.

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Mangave gonzaloi (D. Guillot & P. Van der Meer) hort. (s.a.) (nom. inval., ICN Art. 29.1). [1
3a] A commercially offered garden hybrid with spotted leaves, most probably between a species of Subg. Manfreda (A. maculosa or A. variegata?) and an unknown species of Subg. Littaea (and thus belonging to the synonymized nothogenus
Mangave), found at the Botanical Garden of Valencia (Spain). A. gracielae Galván & Zamudio (Acta Bot. Mex. 105: 2–4, ills., (pp. 3, 6–7), 2013). Type: Mexico, Querétaro (González 569 [IEB, ENCB, MEXU, XAL]). — Distr: Mexico (SE San Luis Potosí, NE Querétaro); on karstic limestone rocks, in pine-oak or cloud forests, 1200–2400 m; flowers April to July. I: Magallán Hernández & al. (2012: 26, as Agave sp. 1); Hochstätter (2015: VIII: 21); Perez Badillo (2016: 120); De-Nova & al. (2018: 24). [1a] Stems short; Ros hemispherical, 16–60 (–70)
35–75 cm, caespitose, forming dense groups/clusters; L (65–) 100–230, linear, at the base with a deltoid sheath 2.5–5
1.8–2.5 (–3) cm at the base, whitish, fleshy, lamina straight to slightly incurved at maturity, fleshy-coriaceous, flexible, keeled below, flat or slightly concave above, esp. towards the apex, smooth on both faces, 13.5–55 (–60)
(1–) 1.2 (–1.5) cm, light green, striated when dried, margins corneous, 3–5 mm wide, with hyaline or yellowish border, minutely serrulate, denticles sometimes bicuspid, sometimes reddish or only at the tip; marginal teeth none; terminal Sp conical-subulate, weak, keeled below, somewhat flat above, 4–6.5 mm, reddish-brown, slightly decurrent; Inf 1.5–2.45 m, ‘spicate’, straight, peduncle 95–185 cm, green, often tinged reddish or reddish-brown, with linear, reddish-brown, often decurrent Bra 2–5 cm long, fertile part  in the upper , 35–75 cm; Fl geminate, 22–27 mm, sessile; Tep green or yellowish-green, often tinged purple, tube slightly flaring, 8–13
4–5 mm, grooved, lobes slightly spreading, ovate-oblong, unequal, 7–11
3–4.5 mm, cucullate, apex with a tuft of hairs; OTep acute; ITep somewhat shorter, rounded, with a broad keel; Fil 30–40 mm, green with reddish tinge or purple, inserted at 2 levels in the

J. Thiede

of the tube; Anth slightly eccentric, upper 10 mm, yellow or purple; Ov cylindrical, grooved, 6–8
4–5 mm, neckless, intruding the tepal tube; Sty 48–55 mm, shorter than the filaments, green with reddish tinge, purple at the apex; Sti slightly thickened; Fr ovoid-ellipsoid, 15–17
10–12 mm, shortly beaked, dark brown; Se semicircular, 3.8–5
2.2–2.5 mm, black, shiny, with marginal wing, hilar notch shallow. According to the protologue closest to A. dasylirioides and A. petrophila, differing from both esp. in its smooth leaves without striae in fresh condition, larger marginal denticles, and flowers tubular below and campanulate above. A. dasylirioides differs further in its usually solitary rosettes, its arching inflorescences, and its occurrence on igneous rocks, and A. petrophila in its campanulate flowers with a shorter tube, globular fruits and its occurrence in tropical deciduous forests. A. gracielae is also similar to A. rzedowskiana, which differs in its smaller inflorescences and its inclined to decumbent peduncle, shorter tepal lobes and the occurrence on igneous rocks. A. kavandivi is a further close relative (see there for differences). The floral buds are eaten as vegetable, and the young inflorescences are cooked and eaten as sweets according to the protologue. The specimen Rzedowski 7128a from San Luis Potosí, placed in A. dasylirioides by Gentry (1982: 249) may belong here according to the protologue. A. gracilipes Trelease (Annual Rep. Missouri Bot. Gard. 22: 95, 1912). Type [syn]: USA, Texas (Mulford 293 + 293a [MO, ILL, NY]). — Lit: Berger (1915: 180–181); Breitung (1963: 120–121, with ills.); Gentry (1982: 526–530, 536, with ills.); Reveal & Hodgson (2002); GonzálezElizondo & al. (2009: 85–87, with ills.). Distr: USA (SE New Mexico, W Texas); Mexico (Chihuahua); gravelly to rocky, often calcareous places in grasslands, desert scrub, and pinyonjuniper woodlands, 1200–1900 m; flowers summer to early fall. I: Pilbeam (2013: 97); Hochstätter (2015: VII: 100); Greulich (2016a: 35). [2l
1g?] Acaulescent; Ros compact, small, 0.6–0.8 m ∅, sparingly and closely suckering; L 60–100, lanceolate, thick at the base, rigid,

Agave AGAVACEAE

gradually acuminate, convex below, openly concave above, smooth, 18–30
5–7 cm, mostly glaucous-green, margins nearly straight to wavy because of the teeth bases, slightly narrowed above the base; marginal teeth narrowly triangular, mostly rather straight or curved, regular, abruptly lenticular and often confluent at the base, larger along mid-leaf and 2–10 mm, light grey, 15–25 mm apart, regularly spaced, 8–12 per side, intervening margin usually slightly concave; terminal Sp usually straight, narrowly to openly grooved above, 20–50 (–100)
4–5 mm, dark reddish-brown to light grey, long-decurrent; Inf 1.8–5 m, paniculate, peduncle slender, with scarious lanceolate Bra, fertile part frequently narrow and deep, with 30–40 ascending part-Inf in the upper of the inflorescence, or fertile part short, with only 10–12 ascending part-Inf in the upper of the inflorescence Ped < 5 mm, very slender; Fl 35–55 mm; Tep yellow at anthesis, red in bud, tube shallow, broadly campanulate, 4–7
9–15 mm, knobby by the base of the filaments, grooved below the sinuses, lobes spreading to ascending at anthesis, overlapping at the base with thin margins becoming involute above, soon narrowing, withering and sinuses spreading, slightly unequal, 14–18 mm; Fil 30–45 mm, slender, inserted just below the rim of the tube, yellow, sometimes red; Anth 16–20 mm, yellow; Ov slender to broadly cylindrical to fusiform, 20–32 mm, neck constricted; Sty intermediate in length between lobes and filaments, yellow, sometimes red; Sti capitate; Fr oblong to ovoid, 35–40
20 mm, sessile or shortly stipitate, straw-coloured; Se 5–6.5
4 mm. — Cytology: 2n = 60 (Reveal & Hodgson 2002). A very variable complex, hypothetically of ancient hybrid (allopolyploid?) origin between A. (parryi ssp.) neomexicana
A. lechuguilla, but presently back-crossing with A. lechuguilla, which explains the dimorphic variation (Gentry 1982). Pinkava & Baker (1985) list the taxon as putative intersubgeneric hybrid A. neomexicana
A. lechuguilla. Reveal & Hodgson (2002) are reluctant to designate the taxon as nothospecies until populations are studied cytologically in detail, and this view is followed here. The fact that A. parryi ssp. neomexicana is absent from

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major parts of the area of A.
gracilipes excludes a spontaneous hybrid origin with A. lechuguilla and would favour the hypothesis of an allopolyploid origin. The diploid chromosome number, however, dismisses an allopolyploid origin, since both putative parental species are tetraploid. For the time being, A. gracilipes must be upheld as valid species, but its origin and status needs further study. A. gracilipes is very similar in habit to A. applanata (Berger 1915) and was formerly included in the concept of the latter as “Texas form”, e.g. by Mulford (1896: 83), but differs from that species in its much smaller flowers with much shorter tubes (Berger 1915). In the protologue, Mulford, 293, 293a, 1895 are designated as “the types” (plus several paratypes), so that all four specimens of both gatherings (ILL 00009818, ILL 0009819, MO 202075, NY 00170405) represent syntypes. Similar plants from N Durango are provisionally placed here by González-Elizondo & al. (2009) as “A. aff. gracilipes”. A. gracillima A. Berger (Agaven, 33, 288 [erratum], 1915). Type: Mexico, Durango (Rose 2341b [US]). — Lit: Rose (1903c: 21); VerhoekWilliams (1975: 283–286); Piña Luján (1986: 13–15); McVaugh (1989: 229–230); CastillejosCruz (2009: [127]–[132], with ills.); all as Manfreda elongata. Distr: Mexico (S Durango, S Nayarit, S Zacatecas (?), C Jalisco); valleys and grassy plains in the mountain region, in oak and tropical deciduous forests and grasslands, 1000–1380 m; flowers mid-August to September. I: Hochstätter (2016: I: 18); Castro-Castro & al. (2017: 62); both as Manfreda elongata.  Manfreda elongata Rose (1903)  Polianthes elongata (Rose) Shinners (1966) (nom. illeg., ICN Art. 53.1); incl. Agave gracilis A. Berger (1915) (nom. illeg., ICN Art. 53.1); incl. Polianthes rosei Shinners (1967) (nom. illeg., ICN Art. 52.1). [3a2] Herbaceous; corm 5–10
2.5–4 cm, cylindrical, bulb 4–6.5
2.5–3 cm, ovoid to cylindrical, covered with fibrous dry leaf bases 6–12 cm long; R fleshy, with filiform ramifications, contractile; L drought-deciduous, 2–5, recurvate to arching, extended, linear-lanceolate,

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slightly canaliculate, glabrous on both faces, apex acute, with a short soft tip, 40–60 (–95)
2–4 (–5) cm, light green, shiny, margins entire, with a hyaline band, slightly revolute; Inf 120–140 (–160) cm, erect, glaucous-green, ‘spicate’, Bra 8–10, similar to the leaves but smaller, fertile Inf part 20–40 (–70) cm, lax, with 20–30 flowers; floral Bra (0.6–) 1.4–2
0.3–0.7 cm, narrowly deltoid; Fl 28–32 mm, solitary, sessile, ascending; Tep yellowish-green to reddish-green, forming a funnel-shaped tube (9–) 13–18
4–5 mm, slightly recurved, lobes oblong, extended to revolute, 11–16
3–6 mm, tip cucullate, with a soft tip and a bundle of short white trichomes; Fil 25–35 mm, erect at anthesis, exceeding the tube for 24–28 mm, inserted in the upper part of the tube on the same level, greenish-yellow with purple dots; Anth 10–12 mm, greenish-yellow; Ov cylindrical, 12–16
4–5 mm, greenish, not extending into the tube, not constricted above; Sty 30–42 mm, yellowish-green with purple dots; Sti capitately 3-lobed; Fr cylindrical, 20–26
12–14 mm, slightly apiculate, tepals persisting; Se deltoid, plane-concave, 3–4
3–5 mm, dull black. Distinguished by its long, recurving, channelled leaves, the long-acuminate floral bracts, and the styles, which are usually much longer than the stamens. The flowers of A. gracillima are similar to those of A. scabra but with a proportionately longer style and a bluntly angled perianth tube (Verhoek-Williams 1975: 285, as Manfreda elongata and M. scabra). The protologue wrongly cites the number of the type collection as “2341” (Verhoek-Williams 1975: 283). The map in Castillejos-Cruz (2009) shows a locality in S Zacatecas, but does not cite a specimen from that state. A. graminifolia (Rose) Govaerts & Thiede (Willdenowia 43(2): 332, 2013). Type: Mexico, Jalisco (Rose 2571 [US, GH, K, MO, NY]). — Lit: Rose (1903a: 11–12, as Polianthes); McVaugh (1989: 251–252, as Polianthes geminiflora var.); Solano Camacho & García-Mendoza (1998: 476–477, as Polianthes). Distr: Mexico (S Zacatecas, Aguascalientes, Jalisco); grasslands, rocky slopes and grassy openings in oak forests, (1400?–) 2000–2250 m; flowers July to September. I:

J. Thiede

Castro-Castro & al. (2015: 145); Castro-Castro & al. (2016: 721); Hochstätter (2016: II: 14); all as Polianthes.  Polianthes graminifolia Rose (1903)  Bravoa graminifolia (Rose) Conzatti (1903)  Polianthes geminiflora var. graminifolia (Rose) McVaugh (1989)  Agave duplicata ssp. graminifolia (Rose) Thiede & Eggli (1999) (incorrect name, ICN Art. 11.4). [3b2] Herbaceous; corm short, 0.8
0.8 cm, bulb oblong, 3
1.6–1.7 cm, covered with dry leaf bases; R contractile, thickened; L 6, linear, ascending or drooping, involute,  densely pubescent on the lower face on the nerves and margins and on exposed portions of the leaf sheath, glabrous on the upper face, 15–20 (–30 (–60))
0.1–0.2 (–0.8) cm; Inf (20–) 50–85 cm, ‘spicate’, peduncle with narrow subulate-tipped Bra 2–6 cm long, pubescent in the lower part, fertile Inf part 15–30 cm, with 8–15 widely spaced flowering nodes, lower ones often on 1 cm long peduncles, lower internode 5–7 cm, internode length decreasing towards the distal portion; Ped 3 mm, elongating in fruit to 4–4.5 (–10?) mm; Fl geminate, 25 mm; Tep rose to dark red, forming a tube curved downwards near the base, widening between the curvature and the base of the lobes, 22–25
3–4 (at the tip) mm, lobes unequal, outer 2–3
(1.5–) 3–4 mm, inner 1.5–3
1.5–2.5 mm, apex obtuse to rounded, rose; St included; Fil filiform, inserted low in the tube 3.5–8 mm above the tip of the ovary; Anth not described; Ov cylindrical, 5
1.5 mm; Sty not exserted at anthesis; Sti 3-lobate, with 3 flat suborbicular lobes,  reaching the anther tips; Fr and Se not described. For some time considered to represent merely an infraspecific taxon of A. coetocapnia (McVaugh 1989: as Polianthes geminiflora var., Thiede & Eggli 1999: as A. duplicata ssp.), it is here accepted at species rank based on Solano Camacho (2002) (where it represents an independent clade in the morphological analysis), García-Mendoza & Solano (2007), and Solano & García-Mendoza (2013) (all as Polianthes). Its most characteristic feature is the pubescence of distinct, upright, stiff hairs to 0.2 mm that cover the veins and margins of the leaf sheath and lamina and the lower part of

Agave AGAVACEAE

the peduncle (McVaugh 1989: as Polianthes geminiflora var. graminifolia). Albeit well collected (29 records from 12 localities; Solano & Feria 2007: 1888), no detailed morphological description is yet available. A. grisea Trelease (Mem. Nation. Acad. Sci. 11: 34–35, tt. 54–56, 1913). Type: Cuba, Santa Clara (Grey 1 [MO [4 sheets, syn?]]). — Lit: Berger (1915: 208–209); Álvarez de Zayas (1996: 118–119, 123, with ills.); Guillot Ortiz & Meer (2011). Distr: S-C Cuba (Bahía de Cienfuegos); on calcareous coastal cliffs; flowers January to March. I: Hochstätter (2015: VII: 70). Incl. Agave grisea var. grisea; incl. Agave grisea var. cienfuegosana Trelease (1913); incl. Agave grisea var. obesispina Trelease (1913). [2p] Acaulescent, not surculose; Ros semiglobular, solitary; L lanceolate, somewhat concave, or oblanceolate, (100–) 110–130 (–160)
(10–) 12–16 (–20) cm, yellowish grey-green, passing into glaucous, or grey, rather dull, margins between the teeth from nearly straight to conspicuously concave; marginal teeth gently curved, broadly triangular, or acuminately tapered from the hardened tops of asymmetrical green prominences, sometimes wider or sublenticular at the leaf base, 2–5 (–10)
1–4 mm, 10–25 mm apart; terminal Sp triquetrously conical or somewhat subulate, slightly curved, flattened or shallowly concave to or beyond the middle or becoming subinvolute, smooth, (10–) 12–22 (–25)
3–6 mm, reddish chestnut-brown or brown,  dull, decurrent for its length or more; Inf 4–6 (–8) m, paniculate, fertile part oblong, part-Inf in the upper of the inflorescence, 45–50 cm, not bulbilliferous; Ped 5–10 mm; Fl 42–47 (–50) mm; Tep golden-yellow, tube 4–8
7–9 mm, lobes 14–16 (–19)
4–5 mm; Fil 30–36 (–40) mm, inserted nearly in the tube throat; Anth 14–18 mm; Ov fusiform, trigonous, 25–30 mm; Sty longer than the filaments; Sti capitate; Fr cylindrical, sometimes subpyriform, 35–40
14–17 (–20) mm, beaked, shortly stipitate; Se subtriangular, 6–7
5–7 mm. The 4 type specimens at MO are cross-labelled, but the cross-labels do not appear to be original (Smith & Figueiredo 2014e: 235). A polymorphic

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species. Since the 2 published varieties merely differ gradually and in addition appear to be connected by transitional forms (Berger 1915: 209), they were included in the synonymy by Álvarez de Zayas (1996) and in the first edition of this handbook. A. grisea differs from A. brittoniana and A. acicularis esp. in its longer leaves, and from A. offoyana (as A. legrelliana) in its greyish leaves and golden-yellow flowers (Álvarez de Zayas 1996). The omnivorous bat Phyllonycteris poeyi feeds predominantly on the nectar of the species (Mancina 2010). According to the IUCN Red List categories, the species is considered as “Endangered” (Berazaín & al. 2005, González-Torres & al. 2016). The plant depicted by Guillot Ortiz & Meer (2011: 12) has teeth on distinct prominences and is most probably misidentified. A. guadalajarana Trelease (Contr. US Nation. Herb. 23: 123, 1920). Type: Mexico, Jalisco (Pringle 4473 [MO, BR, COLO, E, F, JE, K, MEXU, NY, NDG, RM, S, US, VT]). — Lit: Gentry (1982: 531–532, with ills.); McVaugh (1989: 135, 137); Cházaro Basáñez & Mostul (1999: with ills.); Kemble (2006: with ills.); Hernández-Vera & al. (2007b); Starr (2012: 312–313, with ill.). Distr: Mexico (Jalisco, known only from the region of Guadalajara; S Nayarit); grassy slopes in oak woodlands and ecotone with tropical deciduous forests, in volcanic rocky soils, 1400–2000 m; flowers April to November. I: Kemble (2004: 208); Heller (2006: 16, 93); Etter & Kristen (2012: 83); Pilbeam (2013: 99–101); Hochstätter (2015: IV: 22); Moore (2016: 279); Castro-Castro (2017: 136). [2l] Ros compact, 25–50 (–70) cm ∅, broader than tall, solitary, rarely suckering; L numerous, obovate to oblong, rigid, closely imbricate, obtuse, flat to incurved, 20–30
8–12 cm, inner leaves shiny but glaucous, outer leaves dull grey, margins nearly straight, upper part with prominences; upper marginal teeth (5–) 8–10 mm, remote, on strong prominences, those below mid-leaf much smaller, 3–4 mm, reddish-brown to dusty grey, 5–10 mm apart; terminal Sp subulate, straight to sinuous, flat to shallowly hollowed above, roundly keeled below, 25 mm, greyish; Inf

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3–5 m, paniculate, peduncle slender, pruinose, fertile part very lax, part-Inf nearly horizontal, small, remote, 15–25 in the upper of the inflorescence, pale reddish; Fl 60 mm; Tep lobes slender, much longer than the short tube, 20 mm, tips cucullate, greenish or yellow-green to pale yellow, tips in bud sometimes purplish, in flower tips of OTep rose to purple, tips of ITep less so; Fil slightly exserted, rose to purplish; Ov fusiform, 25 mm, green, neck short, slightly constricted and slightly sulcate; Sty longer than the tepals, rose to purplish; Sti capitate; Fr narrowly oblong, thick-walled, 38–45
15–18 mm, shortly beaked, stipitate, dark brown; Se crescent-shaped, 6
4 mm. The protologue is rather short, and the description in Gentry (1982) is incomplete, lacking data on filaments and ovary, which were added here from Cházaro Basáñez & Mostul (1999), Kemble (2006) and Etter & Kristen (1997+: accessed Dec. 2015). The specimen Norris & Taranto 14380 (MICH) from Nayarit doubtfully assigned here by Gentry (1982: 546), is listed under A. guadalajarana by McVaugh (1989). Portal Datos Abiertos UNAM (2018+: accessed Jan. 2019) lists three further specimens from S Nayarit (MEXU 606850, 615834, 878517). Resembling the other members of Sect. Parryanae in the compact rosettes with closely imbricate, short, broad, rigid leaves with the largest teeth confined to the upper third of the margin, and the proportions of flower tube and lobes. The short obtuse leaves resemble those of A. parryi var. truncata, but the conspicuous prominences along the leaf margins and the lax panicles distinguish A. guadalajarana from that species and all others of Sect. Parryanae (as Group) (Gentry 1982). — Hummingbirds are (occasional?) flower visitors (photo in Cházaro Basáñez & Mostul 1999: 12). A.
guemensis D. Guillot & P. Van der Meer (Stud. Bot. (Salamanca) 24: 87–89, 2006). Type: Spain (Anonymous s.n. in VAL 169523 [VAL]). — Distr: Cultivated only. [1f
1f] This is the garden hybrid A. horrida ssp. perotensis (as “A. polyacantha”)
A. warelliana.

J. Thiede

A. guerrerensis (Matuda) G. D. Rowley (Repert. Pl. Succ. 26: 4, 1977). Type: Mexico, Guerrero (González Medrano & al. s.n. [MEXU 231698, MEXU 526057]). — Lit: Matuda (1975: protologue, with ill.); García-Mendoza (2003); Castillejos-Cruz (2009: [145]–[149], with ills.); all as Manfreda. Distr: Mexico (Guerrero); oak forests, 1250 m; flowers in October.  Manfreda guerrerensis Matuda (1975). [3a3] Plants herbaceous; corm 4–8.2
2.2–3.5 cm, succulent, cylindrical, bulb 3–4.2
1.5–3.5 cm, conical to ovoid, covered with dried fibrous leaf bases 1.5–6.5 cm long; R fleshy, with filiform ramifications, contractile; L droughtdeciduous, 2–3, 40–45
6–8 cm, prostrate to semi-erect, linear-lanceolate, narrowed towards the base, semi-amplexicaul, slightly pubescent at the leaf base, apex acute to obtuse, dark green to glaucous-green, with cherry to purple dots, margins entire; Inf (120–) 150–260 cm, erect, glabrous with purple dots at the base, ‘spicate’, peduncular Bra 5–12, lower similar to the leaves, fertile Inf part 35–40 cm, lax, with 6–12 flowers; floral Bra 0.7–3.5
0.4–0.8 cm; Fl 40–50 mm, solitary, ascending, sessile or the basal ones shortly stipitate; Tep reddish-green, semisucculent, forming a tube 27–33
5–7 mm, lobes narrowly elliptic to oblong, erect to extended, 13–17
5–7 mm, tip cucullate, with a bundle of short white trichomes; Fil 30–47 mm, erect at anthesis, exceeding the tube for 23–40 mm, inserted at of the tube length on the same level, reddishgreen; Anth 8–11 mm, yellow with reddish dots; Ov cylindrical, (6–) 8–12
3–5 mm, not prolonging into the tube, not constricted above; Sty 30–38 mm, yellowish-green; Sti capitately 3-lobed; Fr oblong-ellipsoid, 15–21
9–18 mm, without scar near the apex, tepals persisting; Se deltoid, plane-concave, 4–5
3–4 mm, shiny black. According to the protologue, the species keys out near A. debilis (as Manfreda pringlei) in the key to Manfreda of Rose (1903c). A full description complementing the partly sparse data from the protologue was provided by Castillejos-Cruz (2009). The species is remarkable for its pubescent leaves (although the pubescence is less pronounced than in A. maculata and A. pubescens),

Agave AGAVACEAE

127

Fig. 25 Agave guiengola (Etter & Kristen 83: Mexico; Oaxaca, Presa Lic. Benito Juárez, 190 m). (Copyright: J. Etter & M. Kristen)

and is characterized by its broad, ovate-lanceolate leaves gradually narrowing towards the base. Morphologically it is closest to A. scabra and A. pubescens. At present known from 4 collections only (Castillejos-Cruz 2009). — For details of the typification see González Medrano (1991). — The species is erroneously omitted from the Manfreda accounts of Piña Luján (1985), Piña Luján (1986), and Hochstätter (2016). It has to be considered as “Rare” in the IUCN Red List Categories A. guiengola Gentry (Brittonia 12: 98–100, ills., 1960). Type: Mexico, Oaxaca (Gentry 16436 [US, ARIZ, DES, MEXU, MICH]). — Lit: Gentry (1982: 97–99, with ills.); Ullrich (1991k: with ills.); García-Mendoza (2003: with ill.); (García-Mendoza & al. 2019: 8). Distr: Mexico (Oaxaca: Cerro Guiengola); limestone rocks in short-tree forests, 100–1000 m, flowers in March. I: MacDougall (1956: as ‘blue Agave’); MacDougall (1966: as A. guiengolensis); MacDougall (1970: as A. guiengolensis); Irish & Irish (2000: t. 19); Heller (2006: 95, 160); Richter (2011: 62); Pilbeam (2013: 102–103); Greulich (2013); Gonzales & Mera (2015: 12); Moore (2016: 226, 227). – Fig. 25. [1c] Acaulescent; Ros open, mostly solitary; L few, at maturity 9–16 (to 25–30 in cultivation), broadly lanceolate, sometimes ovate, openly ascending, fleshy-leathery, short-acuminate, nearly

flat above but briefly and narrowly channelled apically, 30–50
14–25 cm (larger in cultivation), light grey or white-glaucous, with transversal bands, epidermis finely and densely papillate, margins repand, discontinuous, variously serrate; marginal teeth flattened, blunt, fine or coarse, unequal, 1- to 3-cuspidate, 1–10
2–9 mm, (5–) 15–20 mm apart, often basally confluent, dark brown, on prominences or on a lenticular base; terminal Sp acicular, rounded above and below, pungent, 6–8 (–15) mm, dark brown, not decurrent; Inf erect, (1.6–) 2–4 (–5) m, ‘spicate’, peduncle with recurved, long attenuate upper Bra and bracteoles, Bra scarious, with a dark midvein, subulatelanceolate, 10–20
3 mm at the deltoid base, fertile Inf part from near the base, part-Inf with 2–3 flowers; Ped 5–8 mm; Fl inconspicuous, 30–45 mm; Tep greenish-yellow, pale yellow or yellowish-white, tips reddish or rose on the outer face, 17–18
4–5 mm, erect or slightly incurved, tube funnel-shaped, 2–4 mm (GarcíaMendoza 2003; “virtually tubeless” according to the protologue), lobes elliptic to lanceolate, openly ascending, straight; Fil 15–21 mm, yellowish, inserted at the lobe bases; Anth excentric, versatile, 13 mm, yellow; Ov cylindrical to obconical, sulcate, 10–14 mm, neck narrowly constricted, green; Sty longer than the filaments; Sti thickened; Fr oblong-ovate, thin-walled, 11–18
5–6 mm (from García-Mendoza 6062,

128

ASU0000570!; protologue: 22–24 mm), shortly beaked; Se “small”, without further data. A strikingly ornamental and very distinct species esp. due to its broad, thick, white-glaucous leaves. In the protologue, where the flowers are described as “nearly tubeless”, placed in Sect. Choritepalae (as Group), but differing from the other species (A. bracteosa, A. ellemeetiana, A. gypsicola) geographically as well as in the shape and colouration of the toothed leaves, its strictly monocarpic rather than polycarpic habit (protologue), and its seed morphology (Ullrich 1991k). Ullrich (1991k) therefore regards Sect. Choritepalae (as Group) as artificial (see also the introduction to the genus). The description by García-Mendoza (2003) differs from the protologue, esp. by giving a tube length of 2–4 mm. The fruits, as described above, are among the smallest in the genus. — For the closely related A. gypsicola see there. The type locality Cerro Guiengola is a karstic limestone formation 20 km NW Tehuantepec, where the species grows “on sheer cliff-faces in prodigious numbers, resembling dense colonies of sea-stars” (Trager 2011) and even on the ancient Zapotec pyramids (MacDougall 1956, Ullrich 1991k). The species was also collected at the microwave tower at Santa María Jalapa del Marqués slightly further northwest (MEXU AGA807760). Well-nourished plants in cultivation may develop much larger rosettes and inflorescences (Greulich 2013). A. guiengola is propagated via tissue culture at the Huntington Botanical Garden (Gomez Rhine 2009). ‘Moto Sierra’ K. Griffin (= ISI 2011–14) is a cultivar with confluent teeth giving the leaf-margin a chain-saw-like appearance (Trager 2011), and ‘Creme Brulee’ is a variegated cultivar with cream-coloured margins (Starr 2014: 222, for illustrations see Moore 2016: 227). A.
guignardii Trelease (in L. H. Bailey, Stand. Cycl. Hort. 1: 233, 235, 1914). Nom. inval., ICN Art. 38.1a. Type: not typified. — Distr: Cultivated only. [1b
2o] This is the artificial intersubgeneric hybrid A. attenuata
A. potatorum (as A. verschaffeltii). Apparently lost from cultivation.

J. Thiede

A. guttata Jacobi & C. D. Bouché (Hamburg. Gart.- & Blumenzeit. 21: 561, 1865). Type: not typified. — Lit: Rose (1903c: 21–22); VerhoekWilliams (1975: 233–244); Piña Luján (1985: 35, 85–86, with ill.); McVaugh (1989: 230–231); Castillejos-Cruz (2009: [150]–[156], with ills.); all as Manfreda. Distr: Mexico (Chihuahua, Durango, Zacatecas, San Luis Potosí, Aguascalientes, Nayarit, Jalisco, Guanajuato, Querétaro, Hidalgo, Michoacán); open grassy roadsides, rocky fields, summit of hills, roadcuts, in oak, pine-oak and tropical deciduous forests, desert scrub and grasslands, 1220–2440 m; flowers mid-July to September. I: Curtis’s Bot. Mag. 86: t. 8429, 1912, as A. protuberans; Anonymous (1940); Rodríguez & Castro-Castro (2006: 624); Rodríguez & Castro-Castro (2007: 9); Richter (2007: 42); Hochstätter (2016: I: 21); CastroCastro & al. (2017: 62); all as Manfreda.  Manfreda guttata (Jacobi & C. D. Bouché) Rose (1903)  Polianthes guttata (Jacobi & C. D. Bouché) Shinners (1966); incl. Agave protuberans Engelmann ex Baker (1888)  Leichtlinia protuberans (Engelmann ex Baker) H. Ross (1894); incl. Leichtlinia commutata H. Ross (1896). [3a3] Plants herbaceous; corm 2–5
2–4.5 cm, oblong to subcylindrical, succulent, stoloniferous; bulb 1.8–3
1.8–3.5 cm, cylindrical, nearly completely covered with dried leaf bases 4.5–6 cm long, these slightly membranous at the base and fibrous above; R fleshy, with filiform ramifications, contractile; L drought-deciduous, 2–7 (–13), extended, lanceolate to lanceolate-elliptic, narrowed towards the base, semisucculent, canaliculate, slightly wavy, lower face slightly papillose above the veins, upper face glabrous, apex obtuse, apiculate, (8–) 17–34
(0.9–) 1.2–3.5 cm, light green to glaucous, sometimes with small to large confluent dark-green to purple dots, margins entire, with a cartilaginous, finely denticulate to erose hyaline band; Inf 60–160 cm, erect, light green to yellowish, ‘spicate’; Bra 8–12, lower ones similar to the leaves; fertile Inf part (2.3–) 4–14 cm, dense, with (2–) 4–25 (–33) flowers; floral Bra 0.8–3.1
0.3–1 cm; Fl 27–42 mm, solitary, sessile, ascending; Tep yellowish-green to reddish with purple tinge, semisucculent, forming a tube 7–11
4–5 mm, lobes oblong to

Agave AGAVACEAE

narrowly elliptic, recurved to revolute, 10–15 (–17)
4–5 mm, tip cucullate, with a bundle of short white trichomes; Fil 26–38 (–46) mm, arching upwards at anthesis, exceeding the perianth tube for 19–32 (–42) mm, inserted at the upper end of the tube on the same level, reddishgreen with small purple dots; Anth (9–) 12–15 mm, yellow; Ov cylindrical to slightly ellipsoid, 7–13
4–7 mm, greenish, prolonging 1–5 mm into the perianth tube, slightly constricted above; Sty 36–52 mm, yellowish-green; Sti capitately 3-lobed; Fr ellipsoid, 13–22 (–25)
10–18 mm, with a scar near the apex, base of tepals and style persisting; Se deltoid, planeconcave, with narrow margin, 3–4
2–3 mm, shiny black. The cylindrical storage rhizome and the presence of spreading rhizomes, the denticulate-erose leaf margin, dense inflorescence, and the stubby flowers with revolute lobes and exserted stamens and styles are diagnostic for this widespread species (Verhoek-Williams 1975: 238, CastillejosCruz (2009). Morphologically closest to A. planifolia (as Manfreda) esp. in its flowers, and to an as yet undescribed species “Manfreda valsequillana” (Castillejos-Cruz 2009); the latter appears to be the same as the undescribed M. riosramirii from Mpio. Valsequillo in Puebla, dealt with in the same publication. A. gypsicola García-Mendoza & D. Sandoval (Acta Bot. Mex. 126: e1461: 6, ills. (pp. 7, 15), 2019). Type: Mexico, Oaxaca (García-Mendoza & Franco Martínez 11218 [MEXU [2 parts], IEB, MEXU, MO]). — Distr: Mexico (W-C Oaxaca: Tlaxiaco Distr., upper Río Verde basin); on gypsisols in xerophytic scrub, 1350–1600 m; flowers January to March. [1c] Acaulescent; Ros open, solitary, 0.7–1
1–1.4 m; L 20–40, broadly ovate to semiorbicular, erect, soft, thickened at the base, with acute apex or caudate, flattened, 45–60
20–26 cm, 1.8–2.5
longer than broad, glaucous or glaucous-yellowish, epidermis densely papillate, margins straight, horny, continuous, reddishbrown or whitish-yellow when young, dentate; marginal teeth deltoid, straight or antrorse, 1–3 (–4)
2–4 mm, brownish-red, 10 cm, not bulbilliferous; Fl erect, 68–90 mm;

Tep yellow to yellow-green, tube deeply funnelshaped, 14–22
15–22 mm, lobes erect, slightly unequal, 18–24
4–5 mm, inner shorter, narrower, involute-conduplicate, wilting early and inwards; St long-exserted; Fil erect, longtapered to the apex, 50–65 mm, yellow, inserted rather irregularly above mid-tube; Anth excentric, 25–30 mm, yellow; Ov fusiform, 30–40 mm, green, neck short, thick, constricted; Sty stout, longer than the tepals; Sti capitate, 3-lobate; Fr oblong to obovoid, 40–55
14–20 mm, apex beaked, broadly short-stipitate, on strong pedicels; Se 6–7
4–5 mm, smoothly black, marginal wing low, hilar notch small. Distinguished within Sect. Parryanae (as Group) by the very broad-based acuminate leaves with reflexed teeth, and tepals forming a deep tube and with relatively short lobes (Gentry 1982: 535). A. havardiana hybridizes with A. lechuguilla (= A.
glomeruliflora), and backcrosses with that hybrid (see there). The 4 type sheets at MO are not cross-labelled and represent syntypes. The species has a multiple pollinator syndrome and is pollinated by birds during the day and by bats during the night; nectar production and stigma receptivity is both nocturnal and diurnal (Slauson 2002a). The feeding ecology of flower-visiting hummingbirds was studied by Kuban & Neill (1980) and Kuban & al. (1983), and the nectar sugar variability by Reid & al. (1985).

Agave AGAVACEAE

A. hiemiflora Gentry (Agaves Cont. North Amer., 480–482, ills., 1982). Type: Guatemala (Gentry 23640 [US, ARIZ, DES, MEXU, MICH]). — Lit: Lott & García-Mendoza (1994: online version with ills.); Véliz Pérez (2013: 237–239, 479, 489, with ill.). Distr: Mexico (Chiapas: E mountains), Guatemala (Huehuetenango); montane slopes, often on limestone, in pine-oak or also in montane rain forests, rarely epiphytic, 800–2290 m; flowers October to December. I: Richter (2011: 114, 123); Pilbeam (2013: 106); Hochstätter (2015: VI: 7). [2o] Acaulescent; Ros compact, 1–1.5 m ∅, solitary, not surculose; L 50–90, lanceolate to ovate, openly spreading, rather softly fleshy, gradually narrowed and thickened towards the base, acuminate, flat to slightly concave above, mostly 30–60
8–15 cm, light grey-glaucous to pale green, margins repand to deeply crenate; larger marginal teeth curved or retrorse, rarely straight, (1–) 5–8 mm at mid-leaf, (10–) 20–40 mm apart, light to dark brown or greyish, or teeth smaller and more distal on the repand margins, all on distinct prominences, the slender tips variously curved up or down; terminal Sp slender or thick, sinuous or contorted to straight, openly grooved to flat above, generally 20–40 mm, brown, decurrent; Inf 2–5 m, paniculate, peduncle with remote, triangular, quickly drying Bra, fertile part slender, narrow, part-Inf small, dense, densely tufted with scarious, frequently dark-coloured bractlets, 20–30 in the upper of the inflorescence, on very short (4–5 cm) or longer (10–20 cm) stalks; Ped slender, to 5 mm; Fl slender, 45–55 (–70) mm; Tep yellow, sometimes red in bud, tube broadly tubular to funnel-shaped, angled to rather knobby, lightly grooved, 5–13 mm, lobes oblong, unequal, 15–23 mm, involute; OTep linearlanceolate, erect-ascending, obtuse, frequently reddish towards the tip, overlapping the inner tepals at the base; ITep smaller, narrower but widened at the base, keel narrow; Fil slender, flattened adaxially, (30–) 45–60 mm, inserted in the middle or near the mouth of the tube, reddish; Anth straight, regular, centric, 15–27 mm, yellow; Ov angled-cylindrical, 20–30 (–40) mm, neck short, smooth, little-constricted; Sty finally longer than the stamens; Sti capitate, triquetrous;

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Fr ovoid to oblong, 30–40
14–20 mm, scarcely beaked; Se 5–6
3.5–4 mm. A highland relative of A. seemanniana and A. congesta, sharing with both moderate leaf variability, and an openly grooved to flat terminal spine. It differs from A. seemanniana in a more regular leaf form and size with a less narrowed base, and the densely tufted bracteolate apex of the part-inflorescences, and from A. congesta in its small rosette, less robust and more richly branched open inflorescence with fewer partinflorescences, and filaments more evenly inserted at a lower level in the tube (Gentry 1982: 480). The species is used for fibre. The IUCN Red List Category is “Vulnerable” according to Véliz Pérez (2013). A. hookeri Jacobi (Hamburg. Gart.- & Blumenzeit. 22(4): 168, 1866). Type [neo]: Ex cult. Kew (Brown s.n. [K]). — Lit: Berger (1915: 162–163); Gentry (1982: 338–340, with ills.); McVaugh (1989: 137–138); Hernández-Vera & al. (2007b). Distr: Mexico (Jalisco, Michoacán); apparently cultivated only or as spontaneous escape (?), in mountain regions in pine-oak forests, 1920–2285 m; flowers October to January. I: Richter (2011: 111); Pilbeam (2013: 107); Greulich (2014a: 57–59, 61); [2d] Stems short, thick; Ros to 2 m, solitary; L lanceolate, arching in age, thickly fleshy, gradually narrowed towards base and tip, generally concave above, 120–175
20–25 cm, glaucous to green or yellow-green, margins repand to crenate, esp. in the middle of the lamina, nearly straight below; marginal teeth straight or curved, 8–12 mm (middle of the lamina), dark brown to greyish-brown, 20–50 mm apart, with few smaller intermittent teeth, much reduced and closely spaced towards the base, broadly based on fleshy prominences; terminal Sp subulate, shortly openly grooved above, roundly keeled below, and with linguiform protrusion into the leaf, 35–60 mm, edges decurrent as toothless smooth horny border for 15–20 cm; Inf 7–8 m, paniculate, peduncle with narrow, triangular, reflexed Bra, part-Inf compact, 20–40 in the upper of the inflorescence; floral Bra caducous; Ped long; Fl slender, 63–80 mm; Tep yellow, red to pink in

134

bud, tube grooved, 5–8
13–14 mm, lobes red to pink, linear, erect, elongate, conduplicate, unequal, outer 28–32 mm, rounded on the back, cucullate, inner 2–3 mm shorter, with prominent keel; Fil slender, flattened, 50–60 mm, inserted above mid-tube; Anth excentric, 26–34 mm, yellow; Ov cylindrical, angled, smooth, 34–41 mm, neck long, grooved, constricted; Sty almost as long as the filaments; Sti capitate; Fr oblong, rounded at the tip, thick-walled, 50–55
25 mm, stipitate; Se broadly crescent-shaped, rugose, 8–9
6–7 mm, lucid black, hilar notch broad, marginal wing broad but little raised. Recognizable among the species of Sect. Crenatae (as Group) by its large size, the glaucous leaves with long-decurrent spine-bases, the short flower tube and the very long tepal lobes (Gentry 1982: 340). Gentry’s “type” represents in fact a neotype (McVaugh 1989: 138). The collection Diguet s.n. (P) cited by McVaugh for Jalisco is likely a misidentification with A. inaequidens, and the collection Gentry s.n. (DES, now at ARIZ 266166) from Jalisco cited by Gentry (1982: 352) appears to represent a cultivar introduced from Michoacán according to Hernández-Vera & al. (2007a); a further collection from Jalisco (García-Mendoza & al. 6409, ASU 0000674!, MEXU AG817141) is designated as “semicultivada”. Widely cultivated for pulque in Michoacán. Morphometric studies showed that A. hookeri is most similar to cultivated A. inaequidens, supporting the hypothesis that it could merely represent the extreme of a domestication gradient A. inaequidens wild ! A. inaequidens cultivated ! A. hookeri cultivated (Figueredo-Urbina & Casas 2012, Figueredo & al. 2014). A cluster analysis of morphological data showed A. hookeri to be nested within A. inaequidens (Figueredo-Urbina & al. 2017). For differences from A. inaequidens see there. The name A. hookeri was independently published three times for different taxa: The oldest, A. hookeri K. Koch (1865), is a synonym of A. spicata Cavanilles (1802), the younger A. hookeri Jacobi (1866) was based on cultivated plants and applied by Gentry to plants he collected in Michoacán, and are described above, and the illegitimate A. hookeri Baker (1881) is a synonym

J. Thiede

of A. inaequidens K. Koch (1860). A proposal to conserve the established name A. hookeri Jacobi against A. hookeri K. Koch was published by Figueiredo & Smith (2014) and was recommended by the relevant committee (Applequist 2014: 1368). A. horrida Lemaire ex Jacobi (Hamburg. Gart.- & Blumenzeit. 20: 546–547, 1864). Type [neo]: Mexico, Morelos (Pringle 8206 [US, CM, E, G, MEXU, MO, NDG, NY, P, S, VT]). — Distr: S Mexico. The neotype cited above was designated by Gentry (1982: 146). A. horrida ssp. horrida — Lit: Berger (1915: 100–101); Gentry (1982: 144–146, with ills.); Ullrich (1993b: with ills.); Starr (2012: 110–114, with ills.). Distr: Mexico (México, Morelos, C Puebla, C Guerrero, NW Oaxaca); volcanic rocks and rocky slopes in mountain regions, pineoak forests and rosette scrub, 1600–2440 m; flowers January to March and July. I: Curtis’s Bot. Mag. 106: t. 6511, 1880; García-Mendoza (2002: 186); Heller (2006: 97, 156); Richter (2011: 68, 122); Pilbeam (2013: 108); Hochstätter (2015: IX: 33); Moore (2016: 254–255). Incl. Agave grandidentata Hort. Belg. ex Jacobi (1866); incl. Agave maigretiana Jacobi (1866)  Agave horrida var. maigretiana (Jacobi) Trelease (1914); incl. Agave horrida var. laevior Jacobi (1869); incl. Agave horrida var. micracantha Baker (1877); incl. Agave desmetiana hort. ex Baker (1877) (nom. illeg., ICN Art. 53.1); incl. Agave regeliana hort. ex Baker (1877) (nom. illeg., ICN Art. 53.1); incl. Agave horrida var. macrodonta Van Geert (1878); incl. Agave artichaut Hort. C. Besserer ex A. Berger (1915) (nom. inval., ICN Art. 36.1c); incl. Agave horrida var. latifrons Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave horrida var. monstruosa Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/ 38.1a); incl. Agave horrida var. recurvispina Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave horrida var. viridis Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a); incl. Agave killischkii hort. ex A. Berger (1915) (nom. inval.,

Agave AGAVACEAE

135

Fig. 28 Agave horrida ssp. perotensis (Etter & Kristen 994: Mexico; San Luis Potosí, Sierra Alvarez, 2350 m). (Copyright: J. Etter & M. Kristen)

ICN Art. 36.1c); incl. Agave regelii Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a). [1g] Acaulescent; Ros compact, formidably armed, 0.3–0.6
0.45–0.9 m, solitary; L 80–100 in mature rosettes, ovate to elliptic-lanceolate, patulous, rigidly thick-fleshy, slightly narrowed above the base, short-acuminate, convex below, flat to guttered above, generally 18–35
4–7.5 cm, yellowish-green, green or dark olivegreen, margins thickly horny, straight to sinuous between the teeth; marginal teeth straight to variously curved, sometimes hooked, broadly flattened at the base, massive, generally 10–15 mm, rarely much smaller, light grey, 5–10 mm apart, continuing to near the base of the terminal spine; terminal Sp semicircular to subdeltoid in crosssection, very pungent, flattened rather than grooved above, 25–40 mm; Inf 2–2.5 m, ‘spicate’, slender, peduncle 1–1.5 m, with deltoidattenuate, narrow, appressed Bra, fertile part slender, part-Inf with 1–2 flowers; Ped slender, 4–8 mm; Fl 35–40 mm; Tep dark purple, red or yellow, tube shortly funnel-shaped, 3–5
8–9 mm, lobes linear, obtuse with small hood, equal, 15–16
4–6 mm, the inner broader, with a prominent keel; Fil slender, regular, flattened, 36–42 mm, mostly reddish, inserted near the rim of the tube; Anth centric, regular, 14–16 mm, bronze to red or purple; Ov fusiform, 17–20 mm, neck constricted, smooth or slightly grooved; Fr ovoid, 20–25
12–14 mm, beaked, on strongly

jointed pedicels 10–12 mm long; Se crescentshaped, 3–4
2–3 mm. Exhibited for the first time by Verschaffelt in 1862, and introduced by the collector C. Besserer as A. artichaut (Berger 1915). Hardly separable from some small-leaved forms of A. ghiesbreghtii, from which it otherwise differs by larger, more numerous leaf margin teeth continuing nearly to the base of the terminal spine (Gentry 1982). Plants with yellow or reddish flowers may occur in the same population (e.g. Gentry 23368 (DES 00009541!, ASU 131463!); García Mendoza 6143 (ASU 0000670!)). The collection Steinmann & RamírezAmezcua 5771 (MEXU; det. A. García-Mendoza) is a new record for Guerrero (Portal Datos Abiertos UNAM 2018+: accessed Dec. 2018). The species is self-compatible and mainly pollinated by a nocturnal bat, but two species of diurnal hummingbirds are also regular visitors and successful, though less efficient pollinators (Flores-Torres & Galindo-Escamilla 2017). A. horrida ssp. perotensis B. Ullrich (Cact. Suc. Mex. 35(4): 80, ill. (p. 96), 1990). Type: Mexico, Veracruz (Gentry & al. 20417 [US, ARIZ, MEXU]). — Lit: Gentry (1982: 161–164, as A. obscura); Ullrich (1992f); Jimeno-Sevilla (2010: 377–378); Starr (2012: 111, 113–114); all with ills. Distr: Mexico (N Puebla, C Veracruz). I: Alsemgeest & al. (2006b: 214); Heller (2006: 97); Richter (2011: 115); Pilbeam (2013: 109); Hochstätter (2015: IX: 34). – Fig. 28.

136

[1g] Differs from ssp. horrida: Ros with somewhat more leaves, small to medium-sized; L narrowly linear to ovate, 25–40
5–8 cm, pale green to green; marginal teeth variable, small (3–5 mm) or large (10–15 mm), straight to curved or flexuous, frequently slanted downwards and curved, 10 cm long in the upper of the inflorescence, bulbilliferous after flowering; Fl erect, 40–65 mm, unpleasantly scented; Tep greenish-yellow, tube broadly urceolate, 15–18
10–12 mm, lobes linear-lanceolate, erect, equal, 17–18
5–6 mm, St long-exserted; Fil erect, 50–60 mm, inserted  at mid-tube, yellow; Anth 20–25 mm, yellow; Ov shortly fusiform, 15–25
8–9 mm, neck slightly constricted, 2–4 mm; Fr and Se unknown. — Cytology: 2n = 138, 140–149, 147, 149, 150 (5
), 180 (Satô 1935, Doughty 1936, Granick 1944, Castorena Sanchez & al. 1991, Robert & al. 2008); in cultivars 54–65 (2
), 77–99 (3
), 118–128 (4
), 137–151 (5
) (Lv & al. 2009). Easily recognizable by its green, unarmed mature leaves with a short, dark brown, conical to subulate, non-decurrent terminal spine. The taxon appears to represent a sexually sterile clone that is widely cultivated in fibre plantations and could be of hybrid origin, possibly between A. angustifolia of Sect. Rigidae (as Group) and A. kewensis of Sect. Sisalanae (as Group) (Gentry 1982: 630). Ullrich (1990d) consequently removed it from Gentry’s Group Sisalanae and placed it in

Agave AGAVACEAE

Sect. Rigidae (as Group), but this is not followed here. Molecular AFLP data (Gil-Vega & al. 2007) place A. sisalana closest to A. kewensis and both close to, but separate from species of Sect. Rigidae (as Group). The species is not known from the wild, and was probably widely distributed by pre-Columbian people (Reveal & Hodgson 2002). It is naturalized in (sub-)tropical regions, e.g., in the Mediterranean (e.g., Spain: Guillot Ortiz & Meer (2006c) and Guillot Ortiz & Meer (2008b: as A. amaniensis), both with ills.) and in RSA (Smith & Mössmer 1996, the plants described appear to be esp. well-nourished individuals and reach much larger size than otherwise reported). Cultivars cultivated in Spain are presented by Guillot Ortiz & Meer (2009a: with ills.). For the synonymous A. franceschiana, see also under A. weberi. A. sobolifera Houttuyn (Nat. Hist. 2(8): 374, 1777). Type: not typified. — Lit: Baker (1887: 543, 549, Fig. 165, as A. morrisii); Trelease (1913: 32–33, tt. 44–48); Berger (1915: 207); Hummelinck (1927: with ills.); Adams (1972: 81); Hummelinck (1984: 208–212, 226–228, with ills.); Acevedo-Rodríguez & Strong (2012: 85). Distr: Jamaica; dry rocky well-drained calcareous hillsides, 30–850 m; flowers February to April. I: Richter (2010: 41); Richter (2011: 90, 91, 138); Pilbeam (2013: 200); Hochstätter (2015: VII: 75). Incl. Agave americana Lamarck (1783) (nom. illeg., ICN Art. 53.1); incl. Agave laetevirens K. Koch (1865); incl. Agave morrisii Baker (1887); incl. Agave sobolifera [?] marginata hort. ex Trelease (1908); incl. Agave morrisii Kent (1908) (nom. illeg., ICN Art. 53.1); incl. Agave sobolifera fa. spinidentata Hummelinck (1984) (nom. inval., ICN Art. 38.1a). [2p] Acaulescent; Ros to 2
2.5 m, solitary; L 20 or more, variably lanceolate, massive, curved, gradually acute or somewhat subacuminate, often deeply and conduplicately or undulately concave, 125–200
15–24 cm, 9 cm thick near the base, rather light green, somewhat glossy; marginal teeth curved or reflexed-triangular (rarely straight), 1–4 mm, glossy dark brown, 5–15 mm apart, often hardened on the tops of green prominences

241

of the margin, intervening margin  concave; terminal Sp conical, nearly straight, slightly flattened, grooved or slightly involutely channelled below the middle when mature, smooth, somewhat glossy, 15–25
3–4 mm, reddish-brown, not decurrent; Inf 5–9 m, paniculate, oblong, partInf on rather short spreading branches, above the middle of the inflorescence, freely bulbilliferous; Ped stout, 8–10 mm; Fl 50 mm; Tep goldenyellow to light orange, tube open, 5–7 mm, lobes bluntly triangular-oblong, 20
7 mm; Fil 45 mm, inserted towards mid-tube; Ov narrowly fusiform, 15–20 (–25) mm, from slightly shorter to longer than the perianth; Fr narrowly oblong, turbinately narrowed at the base, 45–50
13–25 mm, shortly beaked at the tip, stipitate; Se 7
4–5 mm. Previously ascribed to Salm-Dyck (Hort. Dyck., 307, 1834), but published earlier, as given above (Govaerts 2014+: accessed Oct. 2018). See Trelease (1913: 33) on the difficult interpretation of this name. Plants from the Cayman Islands previously included in A. sobolifera were published as a separate species, A. caymanensis (see there for differences). A. sobria Brandegee (Proc. Calif. Acad. Sci., ser. 2, 2: 207, 1889). Type [syn]: Mexico, Baja California Sur (Brandegee 2 [UC, CAS 0000145]). — Lit: Gentry (1982: 394–404, with ills.); Turner & al. (1995: 65–67, with ill.). Distr: Mexico (Baja California Sur). The protologue gives “Comondu on the mesas” as type indication, which refers to Brandegee 2 (Trelease 1912a: 56); the 2 available sheets represent syntypes. A. sobria ssp. frailensis Gentry (Occas. Pap. Calif. Acad. Sci. 130: 54–56, ills., 1978). Type: Mexico, Baja California Sur (Gentry & Cech 11264 [US 2562946, ARIZ, MEXU, US]). — Lit: Gentry (1978: 54–58); Gentry (1982: 401–404); Turner & al. (1995: 66); Webb & Starr (2015); all with ills. Distr: Mexico (Baja California Sur: Cape region); on granite hillslopes; flowers May and October. I: Richter (2011: 80, 105); Pilbeam (2013: 202); Hochstätter (2015: V: 19).

242

[2i] Differs from ssp. sobria: Ros compact, smaller, sparingly caespitose; L more numerous, broadly lanceolate, mostly 20–35
6–8 cm, glaucous-green to bluish-glaucous, margins with pronounced prominences; marginal teeth numerous, smaller, mostly 6–10 mm, chestnut-brown to greying, closely spaced; terminal Sp frequently sinuous or contorted, 30–40 mm; Inf with 10–15 part-Inf; Fl slender, 45–63 mm; Ov cylindrical, 25–40 mm, lobes 4–6 mm wide. Ssp. frailensis occupies the S part of the total range of A. sobria, and is the S-most taxon of Sect. Deserticolae. The protologue indicates “Gentry & Cech 11264” at US as “Type”. From the 2 sheets at US, US 2562946 is annotated as “Type” by Gentry (= holotype); US 2540165, annotated as “Type coll.”, is an isotype. A. sobria ssp. roseana (Trelease) Gentry (Occas. Pap. Calif. Acad. Sci. 130: 54, 1978). Type: Mexico, Baja California Sur (Rose 16854 [US, MO, NY]). — Lit: Trelease (1912b: 59, tt. 58–60, as A. roseana); Berger (1915: 267, as A. roseana); Breitung (1963: 173, with ills., as var.); Gentry (1978: 54–56, with ills.); Gentry (1982: 400–403, with ills.); Ullrich (1994b: with ills.); Turner & al. (1995: 66); Webb & Starr (2015: 77, 83, 87, with ill.). Distr: Mexico (Baja California Sur: Espírito Santo Island and adjacent mainland); flowers February to March, May, June and October. I: Pilbeam (2013: 203); Hochstätter (2015: V: 20).  Agave roseana Trelease (1912)  Agave sobria var. roseana (Trelease) I. M. Johnston (1924). [2i] Differs from ssp. sobria: Ros lax, openly spreading, sparsely surculose; L fewer, broadly lanceolate, frequently twisted, acuminate, smaller, 35–50
7–10 cm, yellow-green, margins with prominent prominences, tubercles 1–1.5 mm; marginal teeth flexuous, few, larger, larger teeth 10–25 mm, remote; terminal Sp sinuous to contorted, 50–70 mm; Inf with 8–12 part-Inf; Fl 45–65 mm, lobes 4–5 mm wide. Ssp. roseana occupies the C part of the total range of A. sobria. A. sobria ssp. sobria — Lit: Trelease (1912b: 56, tt. 50–51, as A. sobria); Berger (1915: 265, as

J. Thiede

A. sobria); Breitung (1963: 149, 151, with ills.); Gentry (1978: 48–54, with ills.); Gentry (1982: 396–399, with ills.); Turner & al. (1995: 65–67); Webb & Starr (2014b: 8–11, with ills.). Distr: Mexico (Baja California Sur); widely scattered but common on mountain slopes on both sides of the Sierra de la Giganta, 0–1070 m; flowers February to April, and September. I: Heller (2006: 134); Richter (2011: 80); Pilbeam (2013: 201); Hochstätter (2015: V: 18). Incl. Agave affinis Trelease (1912); incl. Agave carminis Trelease (1912); incl. Agave slevinii I. M. Johnston (1924). [2i] Stems short, or appearing stemless; Ros open, 0.5–1.5
0.5–1.5 m, usually caespitose; L few, linear to lanceolate, straight to curved, sometimes twisted, long-acuminate, thick and convex below towards the base, flat to somewhat concave above, variable, mostly 45–80
5–10 cm, bright glaucous-grey, frequently cross-zoned, margins repand to tuberculate; marginal teeth variously curved or straight, flattened, base broad, mostly 5–10 mm, grey, reddish towards the tips, mostly 30–40 mm apart; terminal Sp acicular, narrowly grooved above, mostly 30–60 mm; Inf 2.5–4 m, paniculate, peduncle with small triangular chartaceous Bra, fertile part slender, sometimes arching, part-Inf compact, nearly globose, 12–20 per inflorescence, on short lateral branches; Ped 2–7 mm; Fl slender, 45–55 mm; Tep pale yellow, tube broadly funnel-shaped, 3–4
9–11 mm, lobes very narrow, linear, rather thin, involute at anthesis, rounded at the tip, cucullate,  equal, 17–22
3–4 mm, inner keeled; Fil slender, 35–47 mm, inserted at the lobe base on a nectary disc; Anth centric, 18–23 mm; Ov tapering at the base, 25–35 mm, neck short, scarcely constricted; Sty slightly longer than the tepals; Sti capitate; Fr oblong, thick-walled, 50–65
15–18 mm, shortly stipitate, apiculate; Se crescent-shaped, 7–8
5–6 mm, narrowly winged. — Cytology: 2n = 60 (Pinkava & Baker 1985). A. sobria is the S-most species of Sect. Deserticolae, occupying the E parts of Baja California Sur, and possibly the most variable species in Baja California (Webb & Starr 2015). Ssp. sobria differs from ssp. roseana and ssp. frailensis by its longer, narrower, very light-glaucous and

Agave AGAVACEAE

long-lanceolate leaves with a straight to repand margin and with remote marginal teeth, and its slender flowers with long, narrower lobes (Gentry 1978, Gentry 1982). Some forms of ssp. sobria were misidentified by Gentry (1982) as A. gigantensis (Webb & Starr 2014b, Webb & Starr 2015; see under that species). A. spicata Cavanilles (Anales Ci. Nat. 5(15): 261, 1802). Type: Ex cult. Madrid (Cavanilles s.n. [MA 215275]). — Lit: Berger (1915: 43–44, as A. yuccaefolia), Breitung (1960: 181–182, with ill., as A. yuccaefolia); Gentry (1982: 85–86, with ill., as A. yuccaefolia); Ullrich (1995b: with ills.); Ullrich (1996b: with ills.); Guillot Ortiz & Meer (2002b); Alsemgeest & al. (2006a: with ills.). Distr: Not known from the wild; flowers September to March (in cultivation). I: Redouté (1802–1816: tt. 328–329, as A. yuccaefolia, 1811); Curtis’s Bot. Mag. 86: t. 5213, 1860, as A. yuccaefolia; Heller (2006: 135); Richter (2011: 61); Hochstätter (2015: VIII: 28).  Agave yuccifolia var. spicata (Cavanilles) A. Terracciano (1885) (incorrect name, ICN Art. 11.4, 25.1); incl. Agave yuccifolia F. Delaroche (1811); incl. Agave spicata Gussone (1825) (nom. illeg., ICN Art. 53.1); incl. Agave guccifolia Mannetti in F. A. Gera (1834) (nom. inval., ICN Art. 61.1); incl. Agave hookeri K. Koch (1865) (nomen rejiciendum, ICN Art. 56.1); incl. Agave cohniana Jacobi (1866); incl. Agave yuccifolia var. caespitosa A. Terracciano (1885); incl. Agave yuccifolia var. viridis Hort. Besaucèle ex A. Berger (1915) (nom. inval., ICN Art. 36.1c/38.1a). [1b] Stems short or none; Ros open, small to medium-sized, suckering, forming dense groups with age; L 12–40, linear, recurving at maturity, soft, pliable, scarcely succulent, weakly and finely fibrous, not or hardly narrowed at the base, long and finely acuminate, convex below, concave above, 50–65
3–3.5 cm, mostly green with pale midstripe, sometimes reddish- or purple-spotted, margins with red cartilaginous band, finely serrulate with unequal denticles, between the larger mostly a minute one; terminal Sp conical to subulate, fine, pungent, 3–8 mm, brown; Inf 2–3 m, ‘spicate’, peduncle with triangular-lanceolate long-acuminate erect

243

Bra, fertile part slender, arching, 27–30 cm, part-Inf mostly with geminate flowers; Ped short; Fl 40 mm, unpleasantly scented; Tep greenish-yellow, tube narrowly cylindrical, 8 mm, lobes linear, obtuse, tip revolute, inner ones somewhat narrower, spreading to reflexed, 15–16 mm, inner narrower; Fil 40–45 mm, inserted at the orifice of the tube, reddish to reddishbrown; Anth 15 mm, yellow; Ov roundly 3-angled, 16–18 mm, neck short; Sty filiform, finally exceeding the filaments; Sti 3-lobed; Fr obovate, broadly 3-angled, thinly woody, 20
17 mm, shortly beaked, light grey-glaucous; Se nearly semicircular, 5 mm, shiny black. The name A. yuccifolia (usually as A. yuccaefolia), ascribed to Redouté (1811) and published with two colour plates (reproduced by Ullrich (1995b: 115) and by Smith & Figueiredo (2016: 11)), was widely applied to this species. The herbarium specimen B-W 06789-01 0 labelled “Malmaison Febr. 1811” is probably taken from Redouté’s plant. Hooker (1860) published a colour plate of an “A. yuccaefolia” plant flowering at Kew that purportedly was received from Real del Monte (Hidalgo, Mexico). Koch (1865: 104) considered Hooker’s plant to be different from Redouté’s plant and published the new name A. hookeri K. Koch 1865 for the former, which antedates the widely applied name A. hookeri Jacobi (1866), thus rendering the latter illegitimate. A. hookeri K. Koch differs from Redouté’s plant mainly in its distinct stem and much longer inflorescence which may be due to its long cultivation under lush conditions, and the name can be referred to the synonymy of A. yuccaefolia (see also under A. hookeri). Smith & Figueiredo (2016) established the correct spelling “yuccifolia” and author (F. Delaroche). Ullrich (1995b) and Ullrich (1996b) replaced A. yuccifolia by the older name A. spicata, which was based on a plant cultivated at Madrid purportedly received from the BG La Habana, Cuba, and established the specimen cited above with the original handwriting of Cavanilles, preserved in 1802, as holotype (Ullrich 1996b: 25). A detailed historical account of both species is provided by Guillot Ortiz & Meer (2002b).

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A. spicata is a very distinct species without close relatives within Sect. Inermes (as Group Amolae) (Gentry 1982). This lead Ullrich (1996b: 29) to place it in a section of its own (Sect. Yuccifoliae (A. Terracciano) Ullrich). Molecular AFLP data place A. spicata closest to A. nizandensis and close to other members of Sect. Inermes (as Group Amolae). The spotted leaves, a feature otherwise only known from Sect. Herbaceae, might indicate that the species is of hybrid origin involving a species of Sect. Herbaceae and one of Subgen. Littaea (Ullrich 1996b: 28: the former as Manfreda). A. stictata Thiede & Eggli (Kakt. and. Sukk. 50(5): 111, 1999). Type: not typified. — Lit: Rose (1903c: 18–19); Verhoek-Williams (1975: 301–306); Piña Luján (1986: 17, with ill.); Castillejos-Cruz (2009: [204]–[209]); all as Manfreda maculata. Distr: Mexico (SW México, N Guerrero, C Oaxaca); disturbed places (rocky slopes and moist shady areas) derived from oak and pine-oak forests, 1180–2000 m; flowers mid-July to mid-September. I: Rodríguez & Castro-Castro (2007); Hochstätter (2016: I: 29); both as Manfreda maculata. Incl. Polianthes maculata Martius (1831)  Manfreda maculata (Martius) Rose (1903). [3a2] Plants herbaceous; corm 2–3.5 (–5)
(0.7–) 1.5–2.5 cm, subcylindrical, bulb 2.5–4 (–5.5) cm, ovoid, covered with dried fibrous leaf bases 2–4 cm long; R fleshy and thickened, with filiform ramifications, contractile; L drought-deciduous, 2–6 (–8), prostrate, lanceolate to narrowly elliptic, narrowed towards the base, canaliculate, densely pubescent on both faces with straight simple hairs 0.6–0.8 mm long, apex acute, 7–22 (–32)
0.6–2.6 (–4.5) cm, dark green with purple dots, margins entire, slightly wavy and with a hyaline band; Inf 30–95 (–120) cm, erect, ‘spicate’, reddish to greenish, pubescent at its base, peduncular Bra 3–5, base truncate, lower similar to the leaves, decreasing in size upwards, fertile part (7–) 12–28 cm, lax, with 6–22 flowers; floral Bra 3–5
3–5 mm; Fl 25–32 mm, solitary, sessile, ascending; Tep greenish-yellowish to reddish with purplish tinge, semisucculent, forming a tube 12–20
4–5 mm, not constricted at the apex

J. Thiede

of the ovary, lobes narrowly elliptic to oblong, revolute, 8–12
3–4 mm, tip cucullate, with a bundle of short white trichomes; Fil 17–22 mm, erect at anthesis, exceeding the tube for 7–14 mm, inserted at mid-tube all on the same level, yellowish-green with cherry-red dots; Anth 6–10 mm, yellow; Ov cylindrical, (6–) 8–12
2–4 mm, greenish, not protruding into the tube, not constricted above; Sty 21–37 mm, yellowishgreen; Sti capitate, 3-lobed, yellowish-green; Fr cylindrical to subellipsoid, 11–17
9–15 mm, without scar near the apex, tepals persisting; Se deltoid, plane-concave, with narrow margin, 3–4
2–3 mm, shiny. Differentiated from most other species of Sect. Herbaceae by its ovoid bulbs, and by the pubescent leaves narrowed at the base into a petiolar portion; pubescent leaves are otherwise only found in A. pubescens (Verhoek-Williams 1975: 303–304, Castillejos-Cruz 2009: [205]), while A. guerrerensis has leaves that are slightly pubescent at the base. When transferring Manfreda maculata to the genus Agave, a new epithet was necessary because of the earlier name A. maculata Regel 1856. The lectotypification attempt (as “Type”) by Verhoek-Williams (1975: 301) is ineffective because the work was not effectively published. A. striata Zuccarini (Flora 15: 2(Beiblatt 2): 98, 1832). Type [neo]: Cult. RBG Kew (Anonymous s.n. [K]). — Lit: Gentry (1982: 242–247, with ills.). Distr: Mexico (Coahuila, Nuevo León, Tamaulipas, Durango, Zacatecas, San Luis Potosí, Querétaro, Hidalgo); limited to drier valleys and plains with annual rainfall 100, long-lanceolate, linear, upcurved to straight, rigid, widest near the base, somewhat keeled above and below, rhombic in cross-section, 25–35 (–50)
0.5–1 cm, glaucous-green, greenish-yellow or purple to

Fig. 54 Agave stricta. (Copyright: U. Eggli)

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reddish-purple in the upper part, striate, margins thin, whitish or pale yellow to reddish, cartilaginous, scabrous-serrulate; terminal Sp acicular, 15–20 (–25) mm, angularly keeled below, flat above, grey, decurrent along the margin for 0.5–1.5 cm, at the base with a grey to blackish projection penetrating into the leaf tissue; Inf (1.5–) 2–3 m, ‘spicate’, erect or ascending, sometimes twisted, peduncle with linear or filiform deciduous whitish Bra 3–6.5 cm long, fertile part in the upper or a little less, part-Inf with geminate flowers; Ped 2–3 mm; Fl ascending to outcurved, campanulate, 25–30 mm; Tep reddish to purplish, tube funnelshaped, angled, grooved, 6–11
7–10 (at the mouth) mm, lobes erect to spreading with incurved tips, deltoid or elliptic, apiculate, equal, 7–10
3–6 mm, inner ones broadly keeled; Fil 30–37 mm, inserted at 2 levels at mid-tube, reddish; Anth centric, 10–13 mm, bronze to brownish-red or purple, opening yellow; Ov cylindrical, sharply 6-angled, 8–14
2–4 mm, slightly intruding the base of the tepal tube, neckless, wine-red; Sty 35 mm, reddish; Fr subglobose to pear-shaped, trigonous, 10–14
8–10 mm, truncate at the base, apiculate, reddish or purplish; Se semicircular, thickly discoid, wing inconspicuous, 3–4
2–3 mm, black. — Cytology: 2n = 50, 60 (Brandham 1969, Simpson & al. 2011). Distinguished from the vegetatively very similar A. striata by the short flower tube, equalled or exceeded in length by the tepals, and from A. dasylirioides by the short, stiff, narrow leaves on elongate stems (Gentry 1982; see also Starr 2012: 314). From A. petrophila, A. stricta differs esp. in its much smaller rosettes, shorter and narrower leaves, larger and reddish or purple flowers, and its subglobose to pear-shaped, slightly larger fruits (García-Mendoza 2011a). Plants with reddish-purple leaves co-occur with typical green-leaved plants without intermediates (Pilbeam 2013: 206). Ullrich (1990c) emphasizes vegetative similarities and suggests subspecific rank under A. striata, but he was not followed by later authors. Gentry (1982: 248) designated the neotype cited above. Ullrich (1991d) provided a first record for N Oaxaca based on Leuenberger 2747 (B, MEXU), but further collections from Oaxaca are

248

meanwhile known (García-Mendoza 2011a). The species is endemic to the Tehuacán-Cuicatlán valley (García-Mendoza 2011a). Photographs from San Luis Potosí labelled A. stricta (Pilbeam 2013: 206) most probably belong to A. striata ssp. striata. A. stringens Trelease (Contr. US Nation. Herb. 23: 114, 1920). Type: Mexico, Jalisco (Trelease s. n. (1904) [MO]). — Lit: Gentry (1982: 582); McVaugh (1989: 143); Hernández-Vera & al. (2007a: 500); Vázquez-García & al. (2007b: 73, t. GG1, GG2). Distr: Mexico (Jalisco: Río Blanco barranca near Guadalajara); on rocks and cliffs. I: Pilbeam (2013: 207). [2g] Ros to 1.5–2 m ∅, apparently solitary; L 18–25, narrowly linear, lowermost nearly on the ground, uppermost upright-recurved, those in between  recurved, concave, thin,  60
1–2 cm, very glaucous, with bud imprints created by the marginal teeth, margins cartilaginous, narrow, yellowish-white, nearly straight; marginal teeth numerous, slightly upcurved, very sharp and slender, 1–2 mm, red or brown, 10 mm apart (Trelease: scarcely 5 mm apart), with small intermittent denticles; terminal Sp conical, 8
2 mm, dark; Inf 2.5–3.5 m, paniculate, peduncle very long, making up  of the inflorescence, part-Inf 10, ascending-spreading, lowermost long-stalked; Tep yellow but flowers not further described; Fr and Se not described. A hardly known and apparently rare but distinctive species, known for long only from the sterile type specimen collected near Guadalajara (Jalisco) and a specimen from a sterile plant grown at Missouri BG, both appearing immature (Gentry 1982). McVaugh (1989) did not know the plant and repeats data from Trelease and Gentry. Hernández-Vera & al. (2007a) could not find the plant at its type locality and assume that it could be a form of A. angustifolia. The species was not known from habitat until its rediscovery by Kristen & Etter (Pilbeam 2013: 207; first published photograph). Etter & Kristen (1997+: accessed 2015) provide photographs of a plant with an infructescence and few additional data confirming the placement in Sect. Rigidae, following McVaugh (1989). The photograph in Hochstätter (2015: VII: 48) labelled “Agave

J. Thiede

stringens” apparently shows a different plant. Further studies are needed. A. subsimplex Trelease (Annual Rep. Missouri Bot. Gard. 22: 60, tt. 63–64, 1912). Type: Mexico, Sonora (Rose 16811 [US, MO, NY]). — Lit: Berger (1915: 268); Breitung (1963: 149, 151, with ill., as A. sobria); Gentry (1972: 131–134, with ills.); Gentry (1978: 27, 31, 34–35, with ills.); Gentry (1982: 404–407, with ills.); Turner & al. (1995: 67–68); Wilder & al. (2008: 136–137, 140, with ills.). Distr: Mexico (Sonora); thinly scattered in small colonies on outwash slopes of granitic and volcanic mountains and the adjacent islands (Tiburón, Dátil, Cholludo), in desert scrub, strictly coastal at low elevations; flowers April to July and October. I: Richter (2011: 81); Pilbeam (2013: 208); Hochstätter (2015: V: 13). [2i] Ros low-spreading, 20–35
50–70 cm, solitary or caespitose; L variable, lanceolate to ovate, thick, rigid, long- to short-acuminate, only a little narrowed towards the base, rounded below, hollowed above, 12–35
3–5 cm, grey-glaucous or light yellow-green, or sometimes purple-tinged, margins nearly straight or with strong prominences; marginal teeth variable, friable, straight or variously curved, rarely 2-tipped, larger teeth 3–15 mm, brown or more often yellowish-grey; terminal Sp subulate, frequently sinuous, shallowly grooved above, 20–40 mm, glaucous-grey, not or only a little decurrent; Inf 2–3.5 m, paniculate, slender, narrow, part-Inf short and small, 5–8 per inflorescence; Fl 40–45 mm; Tep yellow to pink, tube shallow, spreading, 3–4
10 mm, lobes elliptic, ascending, flat, widest at the middle, apiculate and scarcely hooded, equal, 12–15
6–7 mm; Fil 25–28 mm, round in cross-section, inserted below the base of the lobes 3 mm above the tube bottom; Anth centric, 13–15 mm, yellow; Ov 25 mm, with unconstricted neck 5 mm long; Sty slightly longer than the tepals; Sti capitate; Fr variable, (narrowly) oblong, 35–40
10–15 mm, bluntly apiculate, narrowly or broadly stipitate, light glaucous; Se roughly crescent-shaped, mostly 4.5–5
3–4 mm, sooty black, hilum notch narrow, the opposite corner frequently apiculate, margin with a sharp winglike flange.

Agave AGAVACEAE

Closely related to A. deserti and A. cerulata with which it shares small variable leaves, nearly tubeless flowers, and narrow small panicles, all characteristic for Sect. Deserticolae (as Group) (Gentry 1982). A. subsimplex may be closest to the polymorphic A. cerulata of Baja California, as indicated by the small, variable leaves and the narrow, oblong, waxy capsules, but A. subsimplex differs in its more spreading rosettes, wider, less acuminate grey or yellow-green leaves, and filaments inserted below the base of the tepal lobes. The pink to red colour frequently occurring in the style, filaments, and corolla is distinctive for A. subsimplex, but not known in A. deserti and A. cerulata (Gentry 1982). In the population genetic studies of Navarro-Quezada & al. (2003), the three samples of the species form a clade nested within the A. deserti/A. cerulatacomplex, but none of the latter two species is monophyletic. Diurnal (insects, birds) and nocturnal (insects, bats) pollinators are equally effective (Molina-Freaner & Eguiarte 2003). See also notes under A. deserti. A.
taylorii Hort. B. S. Williams (Gard. Chron., ser. nov. 1: 363, 1874). Type: not typified. — Lit: Baker (1877a: 620, 621: fig. 125); Baker (1888a: 186); Coulson (1897: with ill.); Trelease (1914: 236, 238); Berger (1915: 71, as A. taylorii). Distr: Cultivated only. Incl. Agave
taylorea Hort. B. S. Williams ex Rafarin (1877) (nom. inval., ICN Art. 61.1). [1d
1d] This is the garden hybrid A. geminiflora
A. filifera (as A. filamentosa), commercially offered by the nursery of B. S. Williams. Baker (1877a) provides a plate of the plant. The second parent was given as A. densiflora (= A. mitis) in the protologue, but corrected to A. filamentosa (= A. filifera) by Baker (1888a) in agreement with the very narrow leaves. Coulson (1897) provides a detailed description and photograph of a plant in bloom. J. N. Rose prepared 3 specimens labelled A. taylorii from a plant at US: Rose 1132, 1897 (US 302815; leaves), and Rose 4128, 1898 (US 399839 & 399840; leaves, flowers, photographs of a flowering plant; later identified as A. colimana by H. S. Gentry in 1977). Trelease (1914) erroneously lists both parentages.

249

According to Berger (1915), the plant could be identical with A. wrightii (= A. schidigera) and would antedate that name. Apparently no longer extant in cultivation. A. tecta Trelease (Trans. Acad. Sci. St. Louis 23: 145–146, tt. 26–27, 1915). Type: Guatemala (Trelease 17 [ILL [2 sheets, syn]]). — Lit: Standley & Steyermark (1952: 118–119); Breitung (1964: 36, with ill.); Gentry (1982: 612–614, with ills.); Lott & García-Mendoza (1994: & online version); Véliz Pérez (2013: 240–242, with ill.). Distr: NW Guatemala (region of Quetzaltenango); cultivated only, 1500–2600 m; flowers in April. I: Lyons (2002: 162); Hochstätter (2015: II: 9). [2b] Stems very thick and broad; Ros semiglobose, open, 2
3–4 m, broad, freely suckering; L lanceolate or broadly lanceolate, straightly ascending and often outcurved, very thick, base deeply convex and thick, becoming thinner upwards, acuminate, sigmoidally upcurved at the tip, concave to guttered, smooth, 100–200
30–50 cm, greyish or rather dark green, margins almost straigt; marginal teeth recurved, often lustrous, on triangular often curved prominences often dilated at the base to 15 mm or more, 8–10 (–15) mm (at mid-leaf), dull brown or chestnut-brown becoming grey, 20–70 mm apart, tips flattened, triangular, straight or curved from low bases, dull brown, teeth sometimes on a discontinuous horny band formed by their bases; terminal Sp conical or subacicular, subulate, often slightly flexuous, narrowly shortly grooved above to the middle, 40–70
5–7 mm, dull brown or chestnutbrown, becoming grey, long decurrent for 10–15 cm, somewhat intruding dorsally into the green leaf tissue; Inf 5–7 m, paniculate, thick, with fleshy broadly triangular amplexicaul Bra, basal Bra sometimes densely imbricate below, sometimes completely covering the peduncle, fertile part ellipsoid, lax, with 25 slightly ascending part-Inf, apparently not bulbilliferous; Ped thick, to 10 mm; Fl 70–95 mm; Tep yellow to greenish-yellow, buds tinged red, tube funnelshaped, deeply grooved, thick-walled, 10–18
20 mm, lobes linear-lanceolate, unequal, outer 20–33
8–9 mm, thickly apiculate, deeply

250

cucullate, inner shorter, narrower, thickly keeled; Fil broadly flattened, 40–65 mm, inserted at 2 levels 10–13 mm above the tube base; Ov 30–43 mm, neck grooved, not constricted; immature Fr oblong, 60
30 mm; mature Fr and Se unknown. Geographically isolated from other members of Sect. Salmianae (as Group); possibly a remnant of former use and now cultivated as a fence plant (Gentry 1982: 614). Cultivated in many miles of hedges around Quetzaltenango (Guatemala), and easily distinguished from other Guatemalan Agaves by its massive leaves of trough-like shape (Standley & Steyermark 1952). The report from oak forests at Tixtla de Guerrero (Guerrero, Mexico; Velázquez & al. 2003: 9) appears doubtful, or might represent the native habitat? The two type specimens at ILL are not cross-labelled and thus represent syntypes (Smith & Figueiredo 2014e: 244). A. telegraphica H. Jekyll (Garden (London, 1871–1927) 1: 51, ill., 1871). Incorrect name, ICN Art. Preamble 8. Type: not typified. — Lit: Anonymous (1872: 172); Cutak (1943: 151); Mitich (1975: 106–107); Nelson (2001: 9–10); all with ill. Distr: Printed matter only, and on the verge of extinction.  Polea telegraphica (H. Jekyll) E. C. Nelson (2001) (incorrect name, ICN Art. Preamble 8). [2b?] Ros open, solitary; L 7, broadly lanceolate, older arching over, to 200
40 cm, margins with 4–5 teeth, concave between the teeth; Inf paniculate, with 12 regularly horizontal partInf in the upper of the inflorescence, part-Inf always 2 at the same level, and all in the same plane. The species, a “rather entertaining bit of English humor” (Cutak 1943), is based on a drawing of a huge Agave with an inflorescence representing a telegraph pole and its 12 “partinflorescences” being the carrying structure for the telegraph wires. In habit, it resembles species of Sect. Salmianae. A. temacapulinensis A. Vázquez & Cházaro (Novon 22(2): 236–237, ills., 2012). Type: Mexico, Jalisco (Vázquez-García & Cházaro 9092

J. Thiede

[IBUG, MEXU, MICH, MO]). — Lit: Etter & Kristen (2012: 94–95, with ills.); Cházaro Basáñez & al. (2013: with ills.); Vázquez-Garcia & al. (2016: with ill.). Distr: Mexico (Jalisco: Río Verde drainage); calcareous outcrops, 1600–1700 m, ecotones between juniper, thorn, and Taxodium gallery forests, 1600–1700 m; flowers May to June. I: Pilbeam (2013: 209–210); Hochstätter (2015: IV: 15). [2h] Ros open, closed or compact, 0.8–1.15
1.72 m, solitary or occasionally surculose; L 20–25, ovate to lanceolate, firm and smooth, 55–90
(13–) 15–20 (–24) cm, glaucous to light grey, margins crenate; marginal teeth variously curved, usually upwards, with broad base, 8–10
7 mm, 15–21 mm apart at mid-leaf, 12–29 mm apart towards the tip, 6–10 mm apart towards the base; terminal Sp flexuous, channelled, 35–45
9 mm, black to bluish-grey, long-decurrent; Inf 5–6.5 m, ‘paniculate’, peduncle with reflexed triangular Bra, these 20
3 cm, fertile part open, part-Inf 16–18 in the upper , 20–22 cm, with 50–70 flowers; Fl 60–71 mm; Tep yellow, outer face greenish, tube 9–15
8–9 mm, equal to or longer than the lobes, lobes dimorphic, outer 10–15
5–6 mm, apex red, horny, inner 12–13 mm, white-puberulent, apex galeate, all dorsally with a prominent longitudinal central keel except at the apex, drying leathery, persisting erect; Fil inserted unequally at mid-tube, outer inserted 1 mm higher, yellow; Ov 17
5 mm, neck 7
3 mm; Sty slender, nearly as long as the filaments, yellow; Sti 3-lobed, clavate, glandular; Fr oblong to obovoid, 40–51
14–18 mm, stipe 2–8 mm; Se subcircular to semicircular or triangular, 5–6
3–4 mm, black, shiny, wingless. The light glaucous rosettes, dimorphic tepal lobes, and deep floral tube clearly place the species in Sect. Ditepalae where it is most similar morphologically to A. wocomahi (with dark green to glaucous leaves with straight to repand margins, longer teeth, shorter inflorescences with a lower number of branches, and larger seeds), and also to A. durangensis (with rough-textured leaves, longer teeth, and taller inflorescences with more numerous branches). A. temacapulinensis is a locally abundant local endemic, and is threatened

Agave AGAVACEAE

by land degradation and the El Zapotillo dam project, which may flood nearly its entire habitat (Vázquez-Garcia & al. 2016). — According to the protologue, the flowers are visited by hummingbirds during the day and by bats in the evening. A. tenuifolia Zamudio & E. Sánchez (Acta Bot. Mex. 37: 47–52, ills., 1995). Type: Mexico, Querétaro (Carranza 1905 [IEB, MEXU, TEX]). — Distr: Mexico (C Tamaulipas, S Guanajuato, NE Querétaro, Hidalgo, Michoacán); limestone slopes with thin soil, in pine-oak or tropical (sub-)deciduous forests, 450–1500 m; flowers mid-April to July. I: Magallán Hernández & al. (2012: 30, as Agave sp. 2). Pilbeam (2013: 211); Hochstätter (2015: VIII: 12); Greulich (2018b: 90). [1a] Ros lax, caespitose, forming dense groups by axillary or rhizomatous branching; L 30–50 (–90), linear, subcoriaceous, very thin and flexible, young straight, mature recurved, keeled on both faces, (29–) 50–100 (–130)
0.25–0.5 (middle, to 1.3 at the base) cm, green, striate, margins horny, 100 cm with 3–5 Br in old plants; racemes cylindrical, 30–35  4–5 cm, very dense, Fl sessile, arranged in 13 spirally twisted rows; Bra ovate-obtuse, 8–10  6–8 mm, fleshy, reddish; Fl yellow with orange tips, 15 mm, slightly campanulate, base rounded, 7 mm ∅ across the ovary, enlarging to 9 mm at the mouth; OTep free to the base; St exserted 3–4 mm; Sty exserted 5 mm. Flowering starts on the sunny side of the raceme, where the flowers are more orange in colour. A. bicomitum L. C. Leach (Kirkia 10: 385–386, 1977). Type: Tanzania, Western Prov. (Reynolds 8948 [PRE, EA, K]). — Lit: Reynolds (1959: protologue A. venusta). Distr: SE Tanzania, N Zambia: shores of Lake Tanganyika; rock outcrops in Brachystegia woodland, 780–1400 m. I: Reynolds (1966: 174–175, as A. venusta); Carter & al. (2011: 346).  Aloe venusta Reynolds (1959) (nom. illeg., ICN Art. 53.1). [5] Acaulescent, simple or in small groups; L 20, densely rosulate, triangular, 50  9 cm, dull grey-green with many elliptic pale green spots, more crowded towards the base, smaller and more crowded on the lower face, with continuous pinkish margin, marginal teeth 3 mm, uncinate, pinkish, 8–10 mm apart; Inf 1–1.2 m, with 10 Br; racemes cylindrical-conical, 15–20 cm, dense; Bra ovate-cuspidate, 11  10 mm, somewhat fleshy in the middle, imbricate in bud stage; Ped 13 mm; Fl pale scarlet, minutely pubescent, 32 mm, base very shortly attenuate, 7 mm ∅ across the ovary, slightly narrowed above, then enlarging towards the mouth; OTep free for 12 mm; St and Sty exserted 3–5 mm. A. boiteaui Guillaumin (Bull. Mus. Nation. Hist. Nat., Sér. 2, 14: 349, 1942). Type: Madagascar, Toliara (Boiteau s.n. [P]). — Distr: S Madagascar (Toliara: near Tolanaro); known only from the type collection, extinct in the wild. I: Reynolds (1966: 420); Castillon & Castillon (2010: 234–235); Carter & al. (2011: 231).  Lemeea boiteaui (Guillaumin) P. V. Heath (1994).

Aloe ASPHODELACEAE

[1] Acaulescent or with short stem, suckering; L 10, rosulate, triangular, 15–20  1.4 cm, olive-green, with narrow pinkish cartilaginous margin, marginal teeth 0.5–1 mm, pinkish, 2 mm apart, absent towards the apex; Inf 10–15 cm, simple; raceme 5 cm, lax, with 10 flowers; Bra deltoid-acute, 5  3 mm; Ped 10 mm; Fl bright scarlet, 25 mm, base shortly attenuate, slightly narrowed above the ovary, then enlarging towards the mouth; OTep free for 18 mm; St and Sty slightly exserted. A. boscawenii Christian (J. South Afr. Bot. 8 (2): 165–167, ills., 1942). Type: Tanzania, Tanga Distr. (Boscawen 7 in Christian 902 [SRGH, EA, K, PRE]). — Distr: NE Tanzania (Tanga Distr.); scrub on sandy soil along coast, to 60 m. I: Reynolds (1966: 365); Carter & al. (2011: 596). [14] Caulescent, branching near the base; stems 1–2 m  5–7 cm, erect for 20–30 cm, when longer sprawling or supported by surrounding vegetation; L scattered along the stem for 20–30 cm, ovate-lanceolate, 44–50  8 cm, light green, with narrow cartilaginous margin, surface smooth, exudate yellow, marginal teeth 2–3 mm, brown-tipped, pungent, 7–18 mm apart; Inf 90 cm, with 3–9 Br, lower ones sometimes rebranched; racemes cylindrical, 10–12  7 cm, lax below, more dense towards the apex; Bra long-acuminate, 7  3 mm; Ped 18 mm; Fl yellow, becoming brownish towards the apex, 30 mm, base attenuate, 9 mm ∅ across the ovary, very slightly narrowed above; OTep free for 18 mm; St and Sty scarcely exserted. A. bosseri J.-B. Castillon (Adansonia, sér. 3, 22(1): 135–137, ills., 2000). Type: Madagascar, Toliara (Castillon 1 [P]). — Distr: W Madagascar (Toliara, Mahajanga); steep limestone cliffs along the Manambolo River, and adjacent “tsingy” areas. I: Castillon & Castillon (2010: 296–297). [6, 7] Acaulescent or caulescent to 40 cm, suckering at the base; L 10–15, rosulate, tip attenuate, 30–70  4 cm, blue-green to yellowishgreen, upper face finely lineate, margins a 1 mm wide whitish or rose border, marginal teeth absent or 0.1–0.5 mm, 0.5–1 mm apart, leaf sheath

515

1 cm; Inf 70 cm, erect or ascending, simple or with 2–6 Br; racemes cylindrical, 10 cm, lax, with 15–20 flowers; Bra ovate-acute, 2.5–3.5 mm; Ped 12–15 mm; Fl red, with green tips, inner lobes whitish, 20–25 mm, 8 mm ∅ across the ovary; OTep free for 16–21 mm; St and Sty scarcely exserted. A. bowiea Schultes & Schultes fil. (Syst. Veg. 7(1): 704, 1829). Type: [neo — icono]: K [unpubl. drawing]. — Lit: Smith (1990); Smith (1991a). Distr: RSA (Eastern Cape: around Port Elizabeth and Kariega); Valley Bushveld vegetation, dense thickets, 0–100 m. I: Carter & al. (2011: 106); Wyk & Smith (2014: 358–359). Incl. Bowiea africana Haworth (1824)  Chamaealoe africana (Haworth) A. Berger (1905). [1] Acaulescent, suckering to form dense groups; R fusiform; L 18–25, rosulatemultifarious, linear, 10–15  1.25 cm, pale glaucous green with scattered white spots, more numerous on the lower face, marginal teeth minute, soft, white; Inf 45 cm, simple; racemes cylindrical, 15 cm, lax; Bra deltoid; Ped 1–2 mm; Fl greenish-white, 8–15 mm, clavate, base shortly attenuate, enlarging above the ovary to the mouth; OTep free to the base; St and Sty exserted 6–8 mm. A. boylei Baker (Bull. Misc. Inform. Kew 1892: 84, 1892). Type: RSA, KwaZulu-Natal (Allison s.n. in Boyle s.n. [K]). — Distr: RSA, Swaziland. I: Carter & al. (2011: 147); Wyk & Smith (2014: 298–299). Included in A. ecklonis by Glen & Hardy (2000) but treated as distinct by Carter & al. (2011) and Wyk & Smith (2014). Flowers are visited by nectar-seeking birds, and also by the bee Amegilla natalensis, probing the flowers for nectar, but the bees are less efficient pollinators than the birds (Hargreaves & al. 2012). A. boylei ssp. boylei — Distr: RSA (Eastern Cape, KwaZulu-Natal, Mpumalanga, Limpopo); grasslands, often in the mist belt. I: Craib (2006: 33–35); Wyk & Smith (2014: 298–299).

516

Incl. Aloe micracantha Pole-Evans (1923) (nom. illeg., ICN Art. 53.1); incl. Aloe agrophila Reynolds (1936). [4] Caulescent, simple or branching; stem short or up to 20  6 cm with age, simple or branching at the base to form groups of 6–12 shoots; L 10–14, rosulate, lanceolate to ensiform, 50–60  1.5–3 cm, dark green, upper face sometimes lineate and with few scattered elliptic spots near the base, lower face with many white spots from the base to the middle, with soft white margin 2 mm, marginal teeth 1–3 mm, soft, white, 2–5 mm apart, smaller and more crowded towards the leaf tip; Inf 40–60 cm, simple; raceme capitate, 10–12  10–12 cm, with 40 flowers; Bra ovate-acuminate, 20–23 mm; Ped 40–45 mm; Fl salmon-pink, greenish-tipped, 30–35 mm, base attenuate, 11–12 mm ∅ across the ovary, narrrowing to 8–9 mm at the mouth; OTep free almost to the base; St and Sty exserted 0–1 mm. — Cytology: 2n = 14 (Müller 1945). A. boylei ssp. major Hilliard & B. L. Burtt (Notes Roy. Bot. Gard. Edinburgh 42(2): 252, 1985). Type: RSA, KwaZulu-Natal (Hilliard & Burtt 8438 [E, NU]). — Distr: RSA (Eastern Cape, KwaZulu-Natal, Mpumalanga, Limpopo); dry stony grassland, 680–1800 m. I: Reynolds (1950: 154–155, as A. boylei); Carter & al. (2011: 147). [4] Differs from ssp. boylei: Plants more robust; L 7 cm broad; Fl 40 mm. Plants regarded by Reynolds (1950) as a “weak form” of this species (also described as A. agrophila) match Baker’s description of A. boylei, and plants regarded by Reynolds as typical are now ssp. major. Craib (2006) and Wyk & Smith (2014) do not mention this subspecies and presumably do not accept it. A. braamvanwykii Gideon F. Smith & Figueiredo (Bradleya 30: 162–165, ills., 2012). Type: RSA, North-West Prov. (Smith & Figueiredo 1 [PRE]). — Lit: Smith & al. (2017: portrait). Distr: RSA (C-S North-West Prov.); wooded grassland on red sandy loam or occasionally clay. I: Wyk & Smith (2014: 216–217).

L. E. Newton

[7] Acaulescent, solitary or with up to 70 Ros; L 12–15, rosulate, linear-attenuate, 17–26  3.5–5.5 cm, dull mid-green with elongated white spots in irregular transverse bands, smooth, exudate pale yellowish drying purple, marginal teeth 4 mm, with green base and light brown tips, 10–13 mm apart; Inf to 75 cm, erect, with 5–7 Br; racemes cylindrical, 14–17 cm, semidense; Bra narrowly triangular, long attenuate, 5–9  4–5 mm, straw-coloured; Ped 10–12 mm, pinkish-brown; Fl orange-red to bright red, 20–25 mm, base truncate, 6 mm ∅ across the ovary, narrowed above to 3 mm, enlarging to 6–7 mm at the mouth; OTep free for 7 mm; St exserted 2–5 mm; Sty exserted 1–2 mm. Similar to A. transvaalensis, which may also be its closest relative (Smith & al. 2017: 18). A. brachystachys Baker (Curtis’s Bot. Mag. 121: t. 7399 + text, 1895). Type: “Zanzibar” (Kirk s.n. [K]). — Distr: SE Tanzania; rock outcrops in montane bushland and at the edge of mist forest, 1450–2100 m. I: Lavranos & Newton (1976: 279–280, as A. schliebenii). Incl. Aloe lastii Baker (1901); incl. Aloe schliebenii Lavranos (1970). [4] Acaulescent, or with short procumbent stem with age, simple or suckering to form small groups; L 20–30, rosulate, lanceolateattenuate, 30–60  4–8 cm, yellowish-green, red-brown in sun, longitudinally lineate, marginal teeth 1–3 mm, brown-tipped, curved, 10 mm apart; Inf 40–100 cm, simple or occasionally with 1 short Br; raceme cylindrical, 15–20 cm, dense; Bra ovate, 12–14  5–10 mm, fleshy, red, becoming scarious with age, imbricate in bud stage; Ped 16–22 mm; Fl pale orange-red, yellow-green at the mouth, 25–32 mm, clavate, base attenuate, 4–5 mm ∅ across the ovary, not narrowed above, enlarging to 7–8 mm at the mouth; OTep free for 8–10 mm; St and Sty exserted 2–4 mm. One of many plants sent to Kew from Zanzibar by the governor, Sir John Kirk, without locality information. Most of these plants had been collected on the African mainland, in parts of Kenya and Tanzania.

Aloe ASPHODELACEAE

A. branddraaiensis Groenewald (Flow. Pl. South Afr. 20: t. 761 + text, 1940). Type: RSA, Mpumalanga (van der Merwe s.n. [PRE 24208]). — Distr: RSA (Mpumalanga, Limpopo); Bushveld vegetation in frost-free areas, 850–1000 m. I: Reynolds (1950: 219–220); Carter & al. (2011: 183); Wyk & Smith (2014: 218–219). [5] Acaulescent, usually simple; L 20–25, rosulate, sometimes subdistichous or somewhat spirally twisted, lanceolate-attenuate, 35  8–10 cm, green with many longitudinal dull whitish striations and many irregular, somewhat H-shaped, whitish spots, marginal teeth 2–3 mm, pale brown, deflexed, 10–15 mm apart; Inf 1–1.5 m, with many Br, lower ones rebranched; racemes capitate, 3–6  7 cm, with 15 flowers; Bra deltoid-acuminate, 8 mm; Ped 20 mm, shorter on lateral racemes; Fl dull scarlet at the base, paler at the mouth, 27 mm, base rounded, 5.5 mm ∅ across the ovary, narrowed to 3.5 mm above, then enlarging to 6 mm at the mouth; OTep free for 7 mm; St and Sty exserted 1 mm. — Cytology: 2n = 14 (Brandham 1971). A natural hybrid with A. burgersfortensis has been reported (Reynolds 1950). A. brandhamii S. Carter (Fl. Trop. East Afr., Aloaceae, 32–33, ills., 1994). Type: Tanzania, Iringa Distr. (Carter & al. 2600 [K, DAR, EA, NHT]). — Distr: C Tanzania; riverine deciduous forest, light shade on rocky slopes, 750–1200 m. I: Carter & al. (2011: 382). [7] Acaulescent or shortly caulescent, suckering to form small groups; stem to 1 m on old plants, ascending; L densely rosulate, lanceolate, 50–80  10–20 cm, dull dark green, often bronzed, surface smooth, exudate drying yellow, marginal teeth 2–3 mm, pungent, browntipped, 10–20 mm apart; Inf 1.5–2 m, with up to 25 Br, lower ones sometimes rebranched; racemes with flowers secund, 15–30 cm, lax; Bra ovate, 12–15  5–7 mm; Ped 5–9 mm; Fl coral-pink, paler at the mouth, minutely whiteflecked, 30–40 mm, base rounded, 7–8 mm ∅ across the ovary, scarcely narrowed above; OTep free for 10–13 mm; St and Sty exserted 3–4 mm.

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A. brevifolia Miller (Gard. Dict. Abr. ed. 6, [no. 8], 1771). Type: [proposed lecto — icono] Commelin, Praeludia Bot., 73, t. 22, 1703. — Distr: RSA. I: Carter & al. (2011: 398–399); Wyk & Smith (2014: 288–289). Incl. Aloe perfoliata var. δ Linné (1753). A. brevifolia var. brevifolia — Distr: RSA (S Western Cape); open bushland, clay soil in rocky places, to 150 m; neophyte in Spain (Valencia). I: Reynolds (1950: 184–186); Carter & al. (2011: 398); Wyk & Smith (2014: 288–289). Incl. Aloe prolifera Haworth (1804); incl. Aloe prolifera var. major Salm-Dyck (1817); incl. Aloe postgenita Schultes & Schultes fil. (1830)  Aloe brevifolia var. postgenita (Schultes & Schultes fil.) Baker (1880). [6] Acaulescent, suckering to form dense clumps; L 30–40, densely rosulate, lanceolatedeltoid, 6  2 cm, glaucous, lower face with a few soft prickles in a median line or irregular in the upper , tip a firm spine, marginal teeth 2–3 mm, whitish, 10 mm apart; Inf 40 cm, simple; raceme conical, 15  7 cm, subdense; Bra ovatelanceolate, 15 mm; Ped 15 mm; Fl pale scarlet, 38 mm, base truncate, very slightly narrowed above the ovary, slightly enlarging to the mouth; OTep free to the base; St and Sty exserted 5 mm. — Cytology: 2n = 14 (Resende 1937). A. postgenita and A. prolifera var. major are said to be intermediate between this and var. depressa (Wyk & Smith 2014). The taxon was recently recorded as neophyte from Spain (Guillot 2013). A natural hybrid with A. mitriformis has been reported (Reynolds 1950), referred to as “A. nobilis” without authorship by Zonneveld (2003), though with reference to Jacobsen’s Lexicon, which includes A.  nobilis Haworth (see entry under this name). A. brevifolia var. depressa (Haworth) Baker (J. Linn. Soc., Bot. 18(108): 160, 1880). Type: not typified. — Distr: RSA (S Western Cape: near Caledon only); shale cliffs, to 100 m. I: Reynolds (1950: 189); Carter & al. (2011: 399).  Aloe depressa Haworth (1804); incl. Aloe perfoliata var. ζ Linné (1753); incl. Aloe serra De

518

Candolle (1799)  Aloe brevifolia var. serra (De Candolle) A. Berger (1908). [6] Differs from var. brevifolia: L 60, 12–15  6 cm, surface smooth or with white subtuberculate spots in the upper , marginal teeth 2–4 mm; Inf 60 cm; Bra 15 mm; Ped to 20 mm; Fl 40 mm, flame-scarlet. — Cytology: 2n = 14 (Resende 1937). Larger than the typical variety. A. breviscapa Reynolds & P. R. O. Bally (J. South Afr. Bot. 24(4): 176–177, t. 19, 1958). Type: Somalia, Northern Region (Reynolds 8542 [PRE, EA, K]). — Distr: N Somalia; arid gypsum plains, 1400 m. I: Reynolds (1966: 267–268); Carter & al. (2011: 443). [7] Caulescent, suckering to form small to large dense clumps, with almost horizontal Ros; stem short or to 50 cm, decumbent, to 1 m with age; L 24, densely rosulate, lanceolateattenuate, 30–35  8–10 cm, bluish-grey tinged reddish, exudate drying yellow, marginal teeth absent or few in the basal , 1–2 mm, blunt, 10 mm apart; Inf 50 cm, arcuate-ascending, with 4–8 Br; racemes cylindrical, 20–25  6 cm, lax; Bra ovate-acute, 6  3 mm; Ped 10–14 mm; Fl scarlet with a bloom, greenish at the mouth, 26–30 mm, base shortly attenuate, 8 mm ∅ across the ovary, slightly narrowed above; OTep free for 10 mm; St and Sty exserted 2–4 mm. A. broomii Schönland (Rec. Albany Mus. 2: 137, 1907). Type: RSA, Northern Cape (Broom s.n. [not located]). — Distr: RSA, Lesotho. A. broomii var. broomii — Distr: W Lesotho, RSA (Northern Cape, Western Cape, Eastern Cape, S Free State); rocky slopes, amongst grass and bushes, (1000–) 1400–1900 m, flowers in September. I: Reynolds (1950: 163–165); Carter & al. (2011: 263); Wyk & Smith (2014: 144–145). [4] Caulescent, usually simple; stem short or up to 1 m, procumbent, usually simple, sometimes dividing into 2 or 3, covered with dried leaves; L densely rosulate, ovate-lanceolate, acuminate with pungent terminal spine, 30  10 cm, green, obscurely lineate, with reddish-brown horny margin, marginal teeth 1–2 mm, reddish-brown with

L. E. Newton

paler tips, 10–15 mm apart; Inf 1–1.5 m, mostly simple, rarely with 1 Br; racemes cylindrical, slightly acuminate, to 100  6–8 cm, very dense; Bra lanceolate-acute, 30  15 mm, rather fleshy, white to pale lemon with brownish tips; Ped 1–2 mm; Fl pale lemon, 20–25 mm, base rounded, enlarging above the ovary and narrowing to the mouth, completely hidden by the bracts; OTep free to the base; St and Sty exserted 12–15 mm. Natural hybrids with other species have been reported (Reynolds 1950). A. broomii var. tarkaensis Reynolds (J. South Afr. Bot. 2(2): 72–73, ills., 1936). Type: RSA, Eastern Cape (Reynolds 1777 [PRE]). — Distr: RSA (SE Eastern Cape, adjacent Northern Cape); rocky slopes, 1200–1500 m, flowers February– March. I: Reynolds (1950: 166); Carter & al. (2011: 263). [4] Differs from var. broomii: Plants more luxuriant, L 2–3 broader at the base; Bra dry and much shorter; Ped 4 mm; Fl 30 mm, not hidden by the bracts. A. brunneodentata Lavranos & Collenette (Cact. Succ. J. (US) 72(2): 86, ill. (p. 84), 2000). Type: Saudi Arabia, Asir Prov. (Collenette 5826 [K]). — Distr: Saudi Arabia (Asir Prov.); on granite, 1800 m; known only from the type locality. I: Collenette (1999: 20); Carter & al. (2011: 258). [2] Acaulescent, solitary; L densely rosulate, 28–35  7 cm, pale bluish-green, marginal teeth short, brown; Inf 60 cm, simple or with 1 Br; racemes cylindrical, lax; Bra 12–15  4–6 mm; Fl reddish, downy, 24–26 (–35) mm, base rounded, not constricted above the ovary; OTep free for 11 mm; St and Sty not exserted. A. brunneostriata Lavranos & S. Carter (Cact. Succ. J. (US) 64(4): 206–208, ills., 1992). Type: Somalia, Bari Region (Migurtein) (Lavranos & Horwood 10187 [K]). — Distr: NE Somalia (Puntland: Bari Region); sandy plains on limestone, 640 m. I: Carter & al. (2011: 312). [7] Shortly caulescent, suckering from the base to form small groups; Br erect or ascending, 40

Aloe ASPHODELACEAE

cm; L up to 10, laxly rosulate, lanceolate, acute, 30  7 cm, creamy-yellow with many longitudinal reddish-brown lines, surface smooth, marginal teeth absent or of the Per length. — 8 species from the Lower Karoo regions and the Eastern Cape. [2] Ser. Namaquana Van Jaarsveld 1992: Per 25–45 ( 50) mm, gasteriform part  of the Per length, globose but often  indistinct from the upper part of the Per and of  the same ∅. — Only G. pillansii from the Namaqualand. • Sect. Longiflorae Haworth 1827: L tapering, smooth or tuberculate (rarely asperulous), rosulate; Per 18–45 ( 50) mm, gasteriform portion narrowly ellipsoid. [3] Ser. Longifoliae (Haworth) Van Jaarsveld 1992: Per 35–50 mm, hardly gasteriform. — 3 species from the Eastern Cape, Mpumalanga, as well as Swaziland. [4] Ser. Multifariae (Haworth) Van Jaarsveld 1992: Per 18–33 mm, gasteriform part  well defined. — 14 species from the Western Cape and Eastern Cape.

E. Van Jaarsveld

Conservation: Jaarsveld and Raimondo have included 18 Gasteria species in the Red List of South African Plants (Raimondo & al. 2009). Their status ranges from “near threatened” (G. batesiana), “rare” (G. baylissiana, G. bicolor var. liliputana, G. retusa, G. thunbergii, G. ellaphieae, G. rawlinsonii, G. tukhelensis, G. vlokii), “vulnerable” (G. croucheri, G. pillansii var. hallii) to “critically rare” (G. batesiana var. dolomitica, G. disticha, G. doreeniae, G. glauca, G. glomerata, G. armstrongii, G. polita). Most, although rare, are secure due to habitat inaccessibility. However, the biggest threat, apart from unsustainable harvesting for the traditional medicinal market (Crouch & al. 2000), remain urban and agricultural expansion. Horticulture: Gasteria are popular indoor horticultural subjects. Most require some form of shading. They are easily propagated by off-shoots, leaf cuttings or seed which should be sown when fresh. They hybridize readily, and some cultivars have been developed (see, e.g., Jaarsveld 2009). The following names are of unresolved application but are referred to this genus: Aloe angulata var. striata Willdenow (1811); Aloe guttata Salm-Dyck (1834)  Gasteria nigricans var. guttata (Salm-Dyck) Baker (1880); Aloe linguaefolia Willdenow (1809); Aloe linguaefolia var. angustifolia Willdenow (1809); Aloe linguaefolia var. latifolia Willdenow (1809); Aloe linguiformis De Candolle (1800); Aloe macchiata Da Pare (1835); Aloe nigricans var. crassifolia Salm-Dyck (1817); Aloe nigricans var. fasciata Salm-Dyck (1821)  Gasteria nigricans var. fasciata (Salm-Dyck) Haworth (1821)  Gasteria fasciata (Salm-Dyck) Haworth (1827); Aloe nigricans var. latifolia Salm-Dyck (1821) (nom. illeg., Art. 52.1?); Aloe nitida var. brevifolia Salm-Dyck (1816) (nom. inval., Art. 43.1); Aloe pseudonigricans Salm-Dyck (1817)  Gasteria pseudonigricans (Salm-Dyck) Haworth (1821); Aloe quadrangularis Da Pare (1835); Aloe subcarinata var. striata Salm-Dyck (1821) (nom. illeg., Art. 52.1); Aloe subnigricans var. canaliculata Salm-Dyck (1834)  Gasteria subnigricans var. canaliculata (Salm-Dyck) A. Berger (1908)

Gasteria ASPHODELACEAE

(incorrect name, Art. 11.4)  Gasteria pseudonigricans var. canaliculata (Salm-Dyck) H. Jacobsen (1955); Aloe trigona Salm-Dyck (1821)  Gasteria trigona (Salm-Dyck) Haworth (1827); Aloe trigona var. elongata Salm-Dyck (1821) (nom. illeg., Art. 52.1); Gasteria acinacifolia var. spathulata hort. (s.a.) (nom. inval., ICN Art. 29.1?); Gasteria angulata var. truncata (Willdenow) A. Berger (1908); Gasteria brevifolia Haworth (1812); Gasteria brevifolia var. laetevirens Haworth (1827); Gasteria brevifolia var. perviridis Haworth (1827); Gasteria crassifolia Haworth (1827); Gasteria dicta N. E. Brown (1876); Gasteria elongata Baker (1896); Gasteria fasciata var. laxa Haworth (1827); Gasteria formosa Haworth (1827)  Gasteria picta var. formosa (Haworth) Baker (1880); Gasteria gracilis Hort. Saunders ex Baker (1880); Gasteria laevis Haworth (1827); Gasteria linita Haworth (1827); Gasteria nigricans var. polyspila Baker (1880)  Gasteria fasciata var. polyspila (Baker) A. Berger (1908); Gasteria prolifera Lemaire (1869); Gasteria subnigricans Haworth (1827) (nom. illeg., Art. 52.1)  Aloe subnigricans (Haworth) Sprengel (1828)  Gasteria nigricans var. subnigricans (Haworth) Baker (1880); Gasteria subnigricans var. glabrior Haworth (1827) (incorrect name, Art. 11.4)  Gasteria pseudonigricans var. glabrior (Haworth) H. Jacobsen (1955); Gasteria transvaalensis Hort. De Smet ex Baker (1889). G. acinacifolia (Jacquin) Haworth (Suppl. Pl. Succ., 49, 1819). Type: [lecto — icono]: Jacquin, Eclog. Pl. Rar. 49, t. 31, 1811–1816. — Distr: RSA (Eastern Cape); Alagoa and Albany Dune Strandveld vegetations, flowers spring to mid-summer. I; Jaarsveld & Ward-Hilhorst (1994: 40–41); Retief (2015).  Aloe acinacifolia Jacquin (1811); incl. Gasteria acinacifolia var. acinacifolia; incl. Aloe acinacifolia var. angustifolia Salm-Dyck (1817); incl. Aloe acinacifolia var. laetevirens Salm-Dyck (1817); incl. Gasteria nitens Haworth (1819)  Aloe nitens (Haworth) Roemer & Schultes (1829)  Gasteria acinacifolia var. nitens (Haworth) Baker (1880); incl. Gasteria candicans Haworth (1821)  Aloe candicans (Haworth) Roemer &

753

Schultes (1829); incl. Gasteria ensifolia Haworth (1825)  Aloe ensifolia (Haworth) Roemer & Schultes (1829)  Gasteria acinacifolia var. ensifolia (Haworth) Baker (1880); incl. Gasteria pluripuncta Haworth (1827)  Aloe pluripuncta (Haworth) Roemer & Schultes (1829)  Gasteria acinacifolia var. pluripuncta (Haworth) Baker (1896); incl. Gasteria venusta Haworth (1827)  Aloe venusta (Haworth) Roemer & Schultes (1829)  Gasteria acinacifolia var. venusta (Haworth) Baker (1896); incl. Gasteria inexpectata Von Poellnitz (1938). [4] Acaulescent, decumbent to erect, 25–75  65 cm, solitary or proliferating to form small groups; L rosulate, 22–60  4.5–10 cm, linearlanceolate to lorate, erectly spreading and sometimes falcate, keeled, both surfaces dark green, with dense white spots arranged in transverse bands, epidermis smooth, rarely slightly tuberculate, margin cartilaginous, serrulate, rarely entire, tip acute, rarely obtuse, mucronate; juvenile L distichous, lorate, patent to erectly spreading, tuberculate, rarely smooth; Inf variable racemose panicles, usually flat-topped, to 1 m, branched, branches horizontal to erectly spreading; Per 35–45 ( 50) mm, gasteriform part narrowly ellipsoid, of the perianth length, 5–9 mm ∅, pink, often not constricted and indistinct, upper white with green striations; Sty included or exserted for up to 5 mm; Fr 35–43 mm, truncate or obtuse; Se 6–8  5–6 mm. One of the largest species of the genus, confined to the SE coast of RSA. G. armstrongii Schönland (Rec. Albany Mus. 2(4): 258, 1912). Type: RSA, Eastern Cape (Anonymus s.n. [K]). — Distr: RSA (Eastern Cape): Humansdorp Shale Renosterveld vegetation, well camouflaged on the pebble background, flowers in mid-summer. I: Jaarsveld & WardHilhorst (1994: 49, as G. nitida var.).  Gasteria nitida var. armstrongii (Schönland) Van Jaarsveld (1992). [4] Plants solitary or proliferating to form small clusters; L remaining distichous, to 5  3 cm, patent, lorate, surface tuberculate, rarely smooth, tip somewhat retuse, obtuse or truncate, mucronate; Inf racemose, 40–50 cm; Per 20 mm, bright reddish-pink, stipitate for 2–3 mm, gasteriform for

754

E. Van Jaarsveld

slightly > of the length, 5–8 mm ∅, narrowly ellipsoid, constricted above into a tube 4–5 mm ∅, lobes erectly spreading, obtuse, yellowish; Sty included; Fr oblong, 24–30  8 mm; Se 3–4  2 mm. Treated at species rank on the base of the DNA analyses of Zonneveld & Jaarsveld (2005). This is a very rare species, and its coastal habitat is threatened by agriculture and urban expansion. G. barbae Van Jaarsveld (CactusWorld 32(4): 257, ills. (pp. 257–260), 2015). Type: RSA, Western Cape (Van Jaarsveld & al. 25246 [NBG]). — Distr: RSA (Western Cape); sheer coastal cliffs, South Outeniqua Sandstone Fynbos vegetation, flowers late spring. [3] Acaulescent, decumbent, 8–12  8–24 cm, solitary or proliferating from the base forming clusters of 3–5 heads; L 8–12 per rosette, rosulate (but distichous in juvenile plants), linear-lanceolate, linear or deltoid-lanceolate to (4-) 12–17  2.2– 3.8 cm, (slightly) falcate and ascending-spreading, adaxial face flat to becoming canaliculate, flat towards the apex, matt dark green to greyish-green (becoming reddish during the dry season), densely white-tuberculate in obscure transverse bands, abaxial face somewhat convex with a distinct eccentric keel, densely tuberculate, tubercles arranged in obscure transverse bands, margins denticulate to almost entire, apex acute or subacute, sometimes acuminate, mucronate; Inf racemose, to 40–55 cm, to 11-flowered; Fl secundly arranged, pendent, all open at the same time; Ped 8–12 mm, pink; Per 40–43 mm, stipitate for 5 mm, subcylindrical and curved, hardly gasteriform, orange-pink, upper white with green striations, tips erect becoming erectly spreading, obtuse, margins of the ITep free and channelled at the base for 10–12 mm, diverging gradually towards the apex; St 34–37 mm; Anth 3  1.5 mm, included or shortly exserted; Ov 8  3 mm ∅, green; Sty 31 mm; Sti included or shortly exserted, curved upwards, minute; Fr 18–30  7–8 mm ∅, narrowly obovoid, triangular in cross-section, with obtuse apex; Se 3.5–4  1.5–2 mm, black. G. batesiana G. D. Rowley (Nation. Cact. Succ. J. 10(2): 32, 1955). Type [neo]: RSA,

Fig. 1 Gasteria batesiana var. batesiana. (Copyright: U. Eggli)

Mpumalanga (Rowley s.n. [RNG]). — Distr: RSA (Mpumalanga, KwaZulu-Natal). G. batesiana var. batesiana — Distr: RSA (Mpumalanga, KwaZulu-Natal); S-facing cliff faces in river valleys, Valley Bushveld vegetation; flowers spring to mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 37). – Fig. 1. [4] Acaulescent, decumbent to erect, 3–10  8–30 cm, proliferating from the base to form small to large groups, rarely solitary; L distichous at first, becoming rosulate, 5–18  1.5–4 cm, triangular-lanceolate to linear, erectly spreading, becoming recurved, dark green with white spots densely arranged in transverse bands, densely rugulose-tuberculate, margin cartilaginous, serrulate (rarely denticulate), tip acute, rarely obtuse, mucronate; juvenile L lorate, densely tuberculate, tip obtuse, mucronate; Inf racemose, 30–45 cm; Bra 6–12  2–5 mm; Ped 9 mm; Per 35–40 m, stipitate for 3–5 mm, gasterifom part narrowly ellipsoid,

Gasteria ASPHODELACEAE

755

of the perianth length, 6–9 mm ∅, light pink, upper of the perianth white with green striations, inflated to the same ∅ as the lower portion, with a slight constriction in the middle, tips obtuse, white with green median stripes; Sti included or exserted for up to 5 mm; Fr 16–20 mm; Se 4–6  2–3 mm. G. batesiana var. dolomitica Van Jaarsveld & E. A. van Wyk (Aloe 36(4): 74, 2000). Type: RSA, Mpumalanga (Van Jaarsveld & Hankey 15081 [NBG]). — Distr: RSA (Mpumalanga); sheer dolomite cliffs in Origstad Mountain Bushveld vegetation. I: Jaarsveld (2008: 85). [4] Differs from var. batesiana: L linear, 10  1–2 cm, becoming biconvex when turgid, tip obtuse. The distribution (disjunct from var. batesiana) represents the N-most within the genus. G. baylissiana Rauh (J. South Afr. Bot. 43(3): 187–191, ills., 1977). Type: RSA, Eastern Cape (Bayliss s.n. [HEID 30517, PRE]). — Distr: RSA (Eastern Cape); Kowie Thicket vegetation on quartzitic sandstones along the Witterivier (Suurberg), spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 75). [1] Acaulescent, decumbent to erect, 0.5–4 cm tall, proliferating from the base to form small dense groups to 8 cm ∅, rarely solitary; L distichous, 2.5–5.5  2–2.3 cm, lorate, erectly spreading, often becoming patent or recurved, epidermis with dense white cartilaginous tubercles, these very dense, domed to globose and confluent forming a dense reticulation, margin crenulate, becoming continuous towards the tip, tip obtuse, truncate or retuse, mucronate; juvenile L lorate, tuberculate and slightly asperulous; Inf racemose, 8–35 cm, erectly spreading, occasionally with a pair of branches; Per 14–16 mm, stipitate for 1 mm, gasteriform part of the perianth length, red-pink, 6–7.5 mm ∅, then abruptly constricted into a tube 3–4 mm ∅, tube white with green striations; Sty included; Fr 14–20 mm; Se oblong, 4  3 mm. G. bicolor Haworth (Philos. Mag. J. 1826: 275, 1826). Type: [neo — icono]: Salm-Dyck,

Fig. 2 Gasteria bicolor var. bicolor. (Copyright: E. Van Jaarsveld)

Monogr. Gen. Aloes & Mesembr., fasc. 7, Aloe sect. 29, fig. 5, 1863, as Aloe bicolor. — Distr: RSA (Eastern Cape).  Aloe bicolor (Haworth) Schultes (1829). G. bicolor var. bicolor — Distr: RSA (Eastern Cape); Albany Thicket vegetation, flowers in spring and summer. I: Jaarsveld & Ward-Hilhorst (1994: 69). – Figs. 2 and 3. Incl. Aloe maculata Thunberg (1785) (nom. illeg., Art. 53.1)  Gasteria maculata (Thunberg) Haworth (1827)  Gasteria maculata var. maculata; incl. Aloe maculata var. obliqua Aiton (1789) (nom. inval., Art. 43.1); incl. Aloe obliqua Haworth (1802)  Gasteria obliqua (Haworth) Duval (1809); incl. Aloe lingua Ker (1807) (nom. illeg., Art. 53.1); incl. Gasteria picta Haworth (1827); incl. Gasteria retata Haworth (1827); incl. Aloe dictyodes Roemer & Schultes (1829); incl. Aloe zeyheri Salm-Dyck (1836)  Gasteria zeyheri (Salm-Dyck) Baker (1880); incl. Aloe planifolia Baker (1870) (nom. illeg., Art. 53.1)  Gasteria planifolia (Baker) Baker (1880); incl. Gasteria variolosa Baker (1871); incl. Gasteria colubrina N. E. Brown (1877); incl. Gasteria marmorata Hort. Peacock ex Baker (1880); incl. Gasteria spiralis Baker (1880); incl. Gasteria spiralis var. tortulata Baker (1880); incl. Gasteria maculata var. dregeana A. Berger (1908); incl. Gasteria caespitosa Von Poellnitz (1937); incl. Gasteria chamaegigas Von Poellnitz (1937); incl. Gasteria herreana Von Poellnitz (1938); incl. Gasteria longiana Von

756

Fig. 3 Gasteria bicolor var. bicolor. (Copyright: E. Van Jaarsveld)

Poellnitz (1938); incl. Gasteria longibracteata Von Poellnitz (1938); incl. Gasteria salmdyckiana Von Poellnitz (1938); incl. Gasteria biformis Von Poellnitz (1940); incl. Gasteria kirsteana Von Poellnitz (1940); incl. Gasteria loeriensis Von Poellnitz (1940); incl. Gasteria multiplex Von Poellnitz (1940). [1] Decumbent to erect, 8–50 cm tall, with a short leafy stem to 20 cm, proliferating from the base to form small groups; L distichous or rosulate, 8–40  1.5–6 cm, lorate to linear, erectly spreading, slightly falcate and twisted sideways when distichous, with an asymmetrical keel when spirally arranged, dark green and with dense white spots arranged in obscure transverse bands, epidermis smooth, rarely slightly asperulous, margin entire, cartilaginous, serrulate, tubercles becoming confluent towards the tip, tip obtuse, rarely acuminate, with an asymmetrical mucro; juvenile L patent or erectly spreading, lorate, asperulous, obtuse, mucronate; Inf 16–150 cm, rarely simple or mostly branched from the middle, with up to 8 erectly spreading branches; Per 12–20 mm, stipitate for 1–2 mm, gasteriform part > of the perianth length, light pink (rarely white), globose to globose-ellipsoid, 6–9 mm ∅, then abruptly constricted into a tube 3–4 mm ∅, tube white with green striations; Fr 10–25  6–10 mm; Se oblong to rectangular, 2–4 mm. A very variable taxon with many local forms. Three varieties are recognized that often grade into each other.

E. Van Jaarsveld

G. bicolor var. fallax (Haworth) Van Jaarsveld (Aloe 44(4): 98, ills. (pp. 98–99), 2008). Type [neo]: RSA, Eastern Cape (Van Jaarsveld 18794 [NBG]). — Distr: RSA (Eastern Cape); Kowie Thicket vegetation, spring-flowering. I: Jaarsveld (2008: 99).  Gasteria maculata var. fallax Haworth (1827)  Aloe obliqua var. fallax (Haworth) Roemer & Schultes (1829). [1] Differs from var. bicolor: L linear, 13–15  1–1.2 cm. The type specimen cited above is here designated as neotype. G. bicolor var. liliputana (Von Poellnitz) Van Jaarsveld (Aloe 29(1): 21, 1992). Type: RSA, Eastern Cape (Dyer & Britten 508 [PRE]). — Distr: RSA (Eastern Cape); Great Fish Noorsveld vegetation, spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 71).  Gasteria liliputana Von Poellnitz (1938)  Gasteria maculata var. liliputana (Von Poellnitz) Hort. Flevohof (s.a.) (nom. inval., Art. 29.1). [1] Differs from var. bicolor: L distichous or rosulate, 1.5–10  0.8–1.4 cm, epidermis smooth, rarely slightly tuberculate or somewhat asperulous, tip obtuse or acute; Inf simple racemes, 16–40 cm, occasionally with a pair of branches; Per 12–15 mm, diameter of the gasteriform part variable. G. brachyphylla (Salm-Dyck) Van Jaarsveld (Aloe 29(1): 19, 1992). Type: [lecto — icono]: Salm-Dyck, Monogr. Gen. Aloes & Mesembr. 29, t. 8. — Distr: RSA (Western Cape).  Aloe brachyphylla Salm-Dyck (1840). G. brachyphylla var. bayeri Van Jaarsveld (Aloe 29(1): 20, Fig. 21 (p. 19), 1992). Type: RSA, Western Cape (Bayer 1751 [NBG]). — Distr: RSA (Western Cape: Little Karoo); Succulent Karoo and Renosterveld vegetations, flowers late spring to mid-summer. I: Jaarsveld & WardHilhorst (1994: 67). [1] Differs from var. brachyphylla: L shorter, 1.5–5  2.2–2.8 cm, at first erect, becoming patent and recurved, tips often incurved, truncate, epidermis smooth or slightly asperulous; Inf racemose, 25–28 cm; Per 18 mm.

Gasteria ASPHODELACEAE

G. brachyphylla var. brachyphylla — Distr: RSA (Western Cape); Succulent Karoo and Renosterveld vegetations, spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 65). Incl. Aloe nigricans var. marmorata Salm-Dyck (1821); incl. Gasteria nigricans var. marmorata Haworth (1821); incl. Gasteria nigricans var. platyphylla Baker (1880); incl. Gasteria angustiarum Von Poellnitz (1937); incl. Gasteria triebneriana Von Poellnitz (1938); incl. Gasteria joubertii Von Poellnitz (1940); incl. Gasteria vlaaktensis Von Poellnitz (1940). [1] Acaulescent, decumbent to erect, 9–23  7.5–23 cm, proliferating from the base to form small groups; L distichous, 8.5–23  2.2–8 cm, lorate, rarely triangular-lanceolate, epidermis smooth, dark green, both faces with densely arranged white spots in obscure transverse bands, margin crenulate, becoming continuous towards the tip, tip acute, obtuse or truncate; juvenile L lorate, asperulous, densely spotted, obtuse; Inf racemose, simple or with a pair of branches, 0.2–1.1 m; Per 12–22 mm, stipitate for 2–3 mm, gasteriform part > of the perianth length, pink, 5–7 mm ∅, globose or globose-ellipsoid, then constricted into a tube 3–4 mm ∅, tube white with green striations; Sty 7–10 mm, included; Fr oblong, 15–23  7 mm; Se 3–4  2–3 mm. G. carinata (Miller) Duval (Pl. Succ. Horto Alencon., 6, 1809). Type: [lecto — icono]: Commelin, Hort. Med. Amstel., t. 9, 1701. — Lit: Jaarsveld (1998). Distr: RSA (Western Cape). Incl. Aloe disticha var. δ Linné (1753);  Aloe carinata Miller (1768). [4] Acaulescent, decumbent to erect, 3–18 cm tall, proliferating from the base to form small dense groups 15–80 cm ∅; L distichous at first, remaining distichous or becoming rosulate, 3–28  1–10 cm, triangular to triangularlanceolate, erectly spreading, lower leaves spreading, both faces tuberculate or smooth, rarely asperulous, spotted with raised or immersed white domed tubercles in obscure transverse bands, margin cartilaginous, tuberculate-crenulate, rarely denticulate, tip acute, rarely obtuse, truncate or retuse, mucronate; juvenile L distichous, erectly spreading, tuberculate or smooth, lorate; Inf racemose,

757

12–90 cm, occasionally with a pair of side branches; Per variable, 16–27 mm, gasteriform part > of the perianth length, pink, narrowly ellipsoid to rarely globose-ellipsoid, above constricted into a tube 3–5 mm ∅, lobes light pink to white with central green stripes; St included; Ov 6–7  2.5 mm; Sty 14 mm; Fr 19–23  7 mm; Se oblong, 3–4  2 mm. G. carinata var. carinata — Distr: RSA (Western Cape); Renosterveld vegetation and thickets on shale or quartz, springflowering. I: Jaarsveld & Ward-Hilhorst (1994: 55). Incl. Aloe tristicha Medikus (1786); incl. Aloe carinata var. subglabra Haworth (1804); incl. Aloe lingua var. angulata Haworth (1804); incl. Aloe lingua var. multifaria Haworth (1804); incl. Aloe angulata Willdenow (1811)  Gasteria angulata (Willdenow) Haworth (1819)  Gasteria disticha var. angulata (Willdenow) Baker (1880); incl. Aloe excavata Willdenow (1811)  Gasteria excavata (Willdenow) Haworth (1827); incl. Aloe laevis Salm-Dyck (1817)  Aloe angustifolia var. laevis (Salm-Dyck) Salm-Dyck (1821) (nom. inval., Art. 43.1); incl. Aloe obscura var. truncata Salm-Dyck (1817); incl. Aloe pseudoangulata Salm-Dyck (1817); incl. Aloe subcarinata SalmDyck (1817)  Gasteria subcarinata (SalmDyck) Haworth (1819); incl. Gasteria latifolia var. multifaria Haworth (1819); incl. Aloe sulcata Salm-Dyck (1821)  Gasteria sulcata (SalmDyck) Haworth (1827); incl. Gasteria laetepuncta Haworth (1827)  Aloe laetepuncta (Haworth) Roemer & Schultes (1829); incl. Gasteria parva Haworth (1827)  Gasteria carinata var. parva (Haworth) Baker (1896); incl. Gasteria strigata Haworth (1827)  Gasteria carinata var. strigata (Haworth) Baker (1896); incl. Aloe carinata var. laevior Salm-Dyck (1836); incl. Gasteria pallescens Baker (1880); incl. Gasteria parvifolia Baker (1880); incl. Gasteria porphyrophylla Baker (1880); incl. Gasteria carinata var. falcata A. Berger (1908); incl. Gasteria carinata var. latifolia A. Berger (1908); incl. Gasteria trigona var. kewensis A. Berger (1908); incl. Gasteria humilis Von Poellnitz (1929); incl. Gasteria bijliae Von Poellnitz (1937); incl. Gasteria schweickerdtiana

758

E. Van Jaarsveld

Von Poellnitz (1938); incl. Gasteria patentissima Von Poellnitz (1940). [4] L first distichous, becoming rosulate or spirally arranged, 3–18  2.5–10 cm, triangular to triangular-lanceolate with a distinct keel, epidermis with raised white tubercles, tip acute or obtuse. Differs from the other varieties by the triangular to triangular-lanceolate tuberculate and keeled leaves. G. carinata var. glabra (Salm-Dyck) Van Jaarsveld (Cact. Succ. J. (US) 70(2): 70, ill. (p. 68), 1998). Type: [neo — icono]: SalmDyck, Monogr. Gen. Aloes & Mesembr., fasc. 5, Aloe sect. 29, fig. 19, 1849. — Distr: RSA (Western Cape: E of the Gouritz River, around Mossel Bay etc.); Mossel Bay Shale Renosterveld vegetation. I: Jaarsveld (2008: 88). – Fig. 4.  Aloe glabra Salm-Dyck (1817)  Gasteria glabra (Salm-Dyck) Haworth (1819); incl. Aloe glabra var. brevifolia Salm-Dyck (1817); incl. Aloe glabra var. major Salm-Dyck (1821) (nom. illeg., Art. 52.1). [4] Differs from var. carinata: L 3–18  2.5–10 cm, without tubercles. G. carinata var. verrucosa (Miller) Van Jaarsveld (Aloe 29(1): 15, 1992). Type: [lecto — icono]: Boerhaave, Ind. Alter Hort. Lugd.Bat., t. 2, p. 131 (no. 36), 1720. — Distr: RSA (Western Cape); Fynbos vegetation on limestone or shale, mainly spring-flowering.  Aloe verrucosa Miller (1768)  Aloe linguiformis var. verrucosa (Miller) De Candolle (1799)  Gasteria verrucosa (Miller) Duval (1809); incl. Aloe racemosa Lamarck (1783); incl. Aloe carinata De Candolle (1799) (nom. illeg., Art. 53.1); incl. Aloe intermedia Haworth (1804)  Gasteria intermedia (Haworth) Haworth (1812); incl. Aloe lingua Ker (1810) (nom. illeg., Art. 52.1); incl. Aloe subverrucosa Salm-Dyck (1817)  Gasteria subverrucosa (Salm-Dyck) Haworth (1827); incl. Aloe verrucosa var. striata SalmDyck (1817)  Gasteria verrucosa var. striata (Salm-Dyck) Von Poellnitz (1938); incl. Aloe intermedia var. asperrima Salm-Dyck (1821)  Gasteria intermedia var. asperrima (Salm-Dyck)

Fig. 4 Gasteria carinata var. glabra. (Copyright: E. Van Jaarsveld)

Haworth (1821) (nom. inval., Art. 43.1)  Gasteria verrucosa var. asperrima (Salm-Dyck) Von Poellnitz (1938) (nom. inval., Art. 43.1); incl. Aloe subverrucosa var. parvipunctata Salm-Dyck (1821)  Gasteria subverrucosa var. parvipunctata (SalmDyck) Haworth (1827); incl. Aloe verrucosa var. latifolia Salm-Dyck (1821)  Gasteria verrucosa var. latifolia (Salm-Dyck) Haworth (1821); incl. Gasteria repens Haworth (1821); incl. Aloe subverrucosa var. grandipunctata Salm-Dyck (1821) (nom. inval., Art. 26.1)  Gasteria subverrucosa var. grandipunctata (Salm-Dyck) Haworth (1827) (nom. inval., Art. 26.1); incl. Gasteria intermedia var. laevior Haworth (1827); incl. Gasteria intermedia var. longior Haworth (1827); incl. Aloe scaberrima Salm-Dyck (1834)  Gasteria verrucosa var. scaberrima (Salm-Dyck) Baker (1880); incl. Gasteria subverrucosa var. marginata Baker (1880); incl. Gasteria radulosa Baker (1889). [4] Differs from var. carinata: L distichous, rarely becoming rosulate, 3–28  1.5–3.5 cm, linear-lanceolate to lorate, distinctly tuberculate due to raised semitranslucent white or green domed tubercles, green to glaucous-green; Inf 12–30 cm; Per 20–25 mm. Aymerich & Gustamante (2015) report that the taxon is likely to become naturalized in Spain, based on a small number of recent sightings at ruderal places. G. croucheri (Hooker fil.) Baker (J. Linn. Soc., Bot. 18(109): 196, 1880). Type: RSA

Gasteria ASPHODELACEAE

(Cooper s.n. [K]). — Distr: RSA (KwaZuluNatal, N Eastern Cape).  Aloe croucheri Hooker fil. (1869). G. croucheri ssp. croucheri — Distr: RSA (KwaZulu-Natal, N Eastern Cape); quartzitic sandstone cliff faces in Eastern Valley Bushveld vegetation, flowers mid-summer. Incl. Gasteria disticha var. natalensis Baker (1880). [4] Acaulescent, decumbent to erect, 25–40  60 cm ∅, solitary or dividing to form dense groups; L rosulate, 15–36  3–10 cm, triangular to linearlanceolate, erectly spreading, rarely patent or recurved or pendent, both surfaces dark green, often glaucous, with dense white spots arranged in transverse bands, epidermis smooth, rarely slightly asperulous, margin tuberculate, serrulate, rarely denticulate, tip obtuse or acute, mucronate; juvenile L distichous, lorate, patent to erectly spreading, smooth or asperulous, tip acute, rarely obtuse, mucronate; Inf variable flat-topped panicles to 50 cm, or racemose, with or without a pair of branches; Bra 5 mm, piliferous; Ped 6–7 mm, pink; Per 28–40 mm, stipitate for up to 6 mm, gasteriform part narrowly ellipsoid, of the perianth length, 5–9 mm ∅, pink, upper of the perianth white with green striations, inflated to the same ∅ as the lower part, lobes erect becoming erectly spreading, obtuse; Sti included or shortly exserted; Fr 18–25 mm, obtuse; Se 3–4  2–3 mm. G. croucheri ssp. pendulifolia (Van Jaarsveld) Zonneveld (Pl. Syst. Evol. 251: 225, 2005). Type: RSA, KwaZulu-Natal (Van Jaarsveld & al. 9838 [NBG]). — Distr: RSA (KwaZulu-Natal); quartzitic sandstone cliff faces in Eastern Valley Bushveld vegetation. I: Jaarsveld & Ward-Hilhorst (1994: 39, as G. croucheri).  Gasteria pendulifolia Van Jaarsveld (2001). [4] Differs from ssp. croucheri: L linear-lanceolate to lorate-falcate, to 45 cm, basally to 4.5 cm broad, epidermis often glaucous. G. croucheri ssp. pondoensis N. Crouch & al. (Bothalia 41(1): 183–184, ill., 2011). Type: RSA, Eastern Cape (Crouch & Styles 1149 [NH]). — Lit: Crouch & al. (2013). Distr: RSA (Eastern

759

Cape); on quartzitic sandstone cliff faces in Scarp Forest vegetation. [4] Differs from ssp. croucheri: L 20–150 cm; Per 42–50 mm. Both this subspecies and ssp. pendulifolia can have glaucous leaves. Zukulu & al. (2012: 12) report that the plants are used locally for protection against lightning strikes and misfortune through witchcraft. G. disticha (Linné) Haworth (Philos. Mag. Ann. Chem. 1827: 352, 1827). Type: [lecto — icono]: Commelin, Hort. Med. Amstel., t. 8, 1701. — Distr: RSA (Western Cape); Succulent Karoo, Nama Karoo and Renosterveld vegetations, flowers spring to mid-summer.  Aloe disticha Linné (1753). G. disticha var. disticha — Distr: RSA (Western Cape: Robertson Karoo); in the shade of Karoo shrubs, spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 63). Incl. Aloe linguiformis Miller (1768); incl. Aloe lingua var. angustifolia Aiton (1789)  Gasteria angustifolia (Aiton) Duval (1809)  Aloe angustifolia Salm-Dyck (1821) (nom. illeg., Art. 53.1)  Aloe angustifolia (Aiton) Salm-Dyck (1849) (nom. illeg., Art. 53.1)  Gasteria disticha var. angustifolia (Aiton) Baker (1880); incl. Aloe lingua var. crassifolia Aiton (1789)  Gasteria nigricans var. crassifolia (Aiton) Haworth (1821)  Aloe crassifolia (Aiton) Roemer & Schultes (1829); incl. Aloe lingua var. latifolia Haworth (1804)  Gasteria latifolia (Haworth) Haworth (1812); incl. Aloe lingua var. longifolia Haworth (1804)  Gasteria longifolia (Haworth) Duval (1809); incl. Aloe nigricans Haworth (1804)  Gasteria nigricans (Haworth) Duval (1809); incl. Aloe obscura Willdenow (1811) (nom. illeg., Art. 53.1); incl. Aloe conspurcata Salm-Dyck (1817)  Gasteria conspurcata (Salm-Dyck) Haworth (1827)  Gasteria disticha var. conspurcata (Salm-Dyck) Baker (1880); incl. Aloe conspurcata var. truncata Salm-Dyck (1817); incl. Aloe conspurcata var. unilateralis Salm-Dyck (1817); incl. Aloe lingua var. brevifolia Salm-Dyck (1817); incl. Aloe nigricans var. denticulata Salm-Dyck (1817)  Gasteria denticulata (Salm-Dyck) Haworth (1819);

760

incl. Aloe obtusifolia Salm-Dyck (1821)  Gasteria obtusifolia (Salm-Dyck) Haworth (1827); incl. Gasteria mollis Haworth (1821)  Aloe mollis (Haworth) Roemer & Schultes (1829); incl. Gasteria disticha var. major Haworth (1827); incl. Gasteria disticha var. minor Baker (1880). [1] Acaulescent, decumbent to erect, proliferating from the base to form small groups to 23 cm ∅; L distichous, 6–17  3–4.5 cm, lorate, erectly spreading, both faces with dense white spots arranged in irregular transverse bands, epidermis asperulous, margin irregularly undulating, tuberculate-crenulate, tip obtuse, rarely truncate, mucronate; juvenile L often patent or recurved, asperulous, truncate; Inf racemose, simple or with a pair of branches, 20–90 cm; Per 12–20 mm, stipitate for 2–3 mm, gasteriform part > of the perianth length, globose-ellipsoid or narrowly ellipsoid, 6 mm ∅, pink to reddish-pink, then constricted into a tube of 3–4 mm ∅, white with green striations; Fr oblong, 15–23  7 mm; Se 3–4  2–3 mm. G. disticha var. langebergensis Van Jaarsveld (Aloe 44(4): 99, ill., 2008). Type: RSA, Western Cape (Van Jaarsveld & Visser 19893 [NBG]). — Distr: RSA (Western Cape: Robertson Karoo: lower slopes of the Langeberg); spring-flowering. [1] Differs from var. disticha: Forming small clusters; L linear-lorate, 3–5  1–1.2 cm, margin denticulate. G. disticha var. robusta Van Jaarsveld (Aloe 44(4): 99, ill., 2008). Type: RSA, Western Cape (Smith s.n. [PRE]). — Distr: RSA (Western Cape: Gamka Karoo); mid-summer-flowering. [1] Differs from var. disticha: Proliferating to form small groups; L lorate, 5.5–10  4.5–6.5 cm. G. doreeniae Van Jaarsveld & A. E. van Wyk (Aloe 41(4): 81–82, ills., 2005). Type: RSA, Eastern Cape (Court 448 [NBG]). — Distr: RSA (Eastern Cape); on cliffs in Kowie Thicket vegetation, spring-flowering. [1] Acaulescent, proliferating and forming clusters; L distichous, lorate, 3.5–8  1.4–2.5 cm, surface asperulous, mottled with dark green and white spots; Inf racemose, 12–40 cm, usually unbranched;

E. Van Jaarsveld

Per 15–17 mm, globose-ovoid for the lower of its length, pink, uppermost pale pink with green striation; Fr 18 mm; Se 3–4  2–3 mm. Formerly classified as G. bicolor var. liliputana, e.g. by Jaarsveld & Ward-Hilhorst (1994). G. ellaphieae Van Jaarsveld (Cact. Succ. J. (US) 63(1): 3–7, ills., 1991). Type: RSA, Eastern Cape (Van Jaarsveld & al. 9904 [NBG]). — Distr: RSA (Eastern Cape); Gamtoos Thicket vegetation, in rock crevices; flowers early to mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 51). [4] Acaulescent, decumbent to erect, 1.5–4  5–16 cm, solitary or proliferating from the base to form small groups; L first distichous, becoming rosulate, 2–5  1–2 cm, triangular to triangularlanceolate and falcate, the inner erectly spreading, the outer recurved, lower face with an asymmetrical keel, both faces dark green with dense white tubercles arranged in irregular transverse bands, epidermis tuberculate, margin tuberculatedenticulate, tip acute or acuminate, recurved, mucronate; juvenile L distichous, lorate, ascending at first, becoming patent or recurved; Inf erectly spreading racemes, 25–40 cm, unbranched or branched in the upper and with 8–15 flowers; Per 22–27 mm, reddish-pink, stipitate for 2.5 mm, gasteriform for slightly > of the perianth length, 7.5 mm ∅, narrowly ellipsoid, above constricted into a tube 4.5 mm ∅, lobes erectly spreading, obtuse, white with central green striae; Sty included; Fr 12–15  5–6 mm; Se 3  2 mm. G. excelsa Baker (J. Linn. Soc., Bot. 18(109): 195, 1880). Type: RSA, Eastern Cape (Cooper s.n. [K]). — Distr: RSA (Eastern Cape); Valley Bushveld and Albany Thicket vegetations, rocky river valleys, flowers mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 43). Incl. Gasteria fuscopunctata Baker (1880); incl. Gasteria huttoniae N. E. Brown (1908); incl. Gasteria lutzii Von Poellnitz (1933). [4] Acaulescent, robust, decumbent to erect, solitary, 30–60  60–75 cm; L in a dense Ros, 10–40  10–18 cm, triangular to triangularlanceolate, erectly spreading to somewhat recurved, both faces dark green with indistinct white spots

Gasteria ASPHODELACEAE

arranged in transverse bands, which (rarely) may be barely visible, rarely striate, epidermis smooth, margin often very sharp, cartilaginous, serrulate, tip acute or obtuse, mucronate; juvenile L distichous, densely white-spotted, patent, lorate, tuberculate; Inf very large spreading panicles, 1–3 per plant, 1–1.9 m, branches erectly (rarely horizontally) spreading; Per 22–26 mm, stipitate for 2–3 mm, gasteriform part > of the perianth length, narrowly ellipsoid, tube constricted to 5 mm ∅, light pink (rarely white), 6–7 mm ∅ at the widest point, lobes white with green midstripes; Sty included; Fr 17–20  8–12 mm; Se 4–5  2–3 mm. Cumming (2003) counted the reproductive success of a population with 51 flowering-size plants. He found 116 racemes with 150–260 capsules each, with 10–85 seeds per capsule, resulting in the production of 1.132 million seeds in the study year. He observed only 4 seedling plants/year, resulting in an establishment rate of 1:300,000. Crook & al. (2009) found, based on casual observation, that some seeds remain enclosed at the bottom of open capsules, but further studies are needed to establish whether this is a general feature, since the capsules do not differ from those of other species of the genus. G. glauca Van Jaarsveld (Cact. Succ. J. (US) 70(2): 65–66, ills., 1998). Type: RSA, Eastern Cape (Van Jaarsveld & Welsh 14670 [PRE]). — Lit: Jaarsveld & Thomas (2003). Distr: RSA (Eastern Cape: Kouga River); sheer S-facing cliff-faces in Gamtoos Thicket vegetation, flowers in mid-summer. [4] Acaulescent, decumbent to erect, proliferating from the base to form dense clusters to 30 cm ∅; L rosulate, 5–7  1.5–1.8 cm, loratelanceolate, upper face slightly canaliculate in the upper , lower face somewhat convex with a distinct asymmetrical keel, both surfaces glaucous, epidermis tuberculate-asperulous, margins tuberculate-dentate, tip acute, mucronate; Inf simple racemes to 25 cm with 10–20 flowers in the upper ; Per 30–34 mm, reddish-pink, gasteriform basally for  of its length or slightly less, globose-ellipsoid, then constricted into a

761

tube 5 mm wide; St to 32 mm; Anth yellow, included; Ov yellowish-green, 6  2.5 mm; Fr oblong, 20  8 mm; Se 3  2 mm, black. Closely related to G. ellaphieae, which also occurs along the Kouga River. G. glomerata Van Jaarsveld (Bradleya 9: 100–104, ills., SEM-ills., 1991). Type: RSA, Eastern Cape (Van Jaarsveld & Sardien 11054 [NBG]). — Distr: RSA (Eastern Cape); on sheer sandstone cliff faces in Gamtoos Thicket vegetation, S aspects, spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 77). [1] Acaulescent, decumbent to erect, 1.5–4  2–8 cm, proliferating from the base to form dense globose clusters to 20 cm ∅; L distichous, 1.5–5  1.5–2.5 cm, lorate to widely ovate, the inner erectly spreading, the outer patent or recurved, biconvex in cross-section to almost terete, becoming flattened during the dry season, upper face often retuse in the upper , both faces glaucous, unspotted, epidermis minutely tuberculate, asperulous, margin entire, minutely crenulate-tuberculate in the upper , tip truncate or obtuse, mucronate; juvenile L distichous, lorate, ascending at first, becoming patent or recurved, asperulous, only slightly and obscurely tuberculate; Inf erectly spreading racemes, simple, 12–20 cm; Per 20–27 mm, gasteriform part slightly > of the perianth length, reddish-pink, globose to globose-ellipsoid, 6–9 ( 10) mm ∅, variable, then constricted into a tube 4 mm ∅; Fr 16  8 mm; Se 3  2 mm. G. koenii Van Jaarsveld (Haseltonia 23: 48, ills. (pp. 49–52), 2017). Type: RSA, Western Cape (Van Jaarsveld & al. 26922 [NBG]). — Distr: RSA (Western Cape: S foothills of the Osberg (Groot Swartberg)); in Eastern Valley Bushveld vegetation, 270 m; summer-flowering. [1] Acaulescent, decumbent, 30  to 35 cm ∅, proliferating from the base and forming clusters with 3–8 heads; R succulent, to 5 mm ∅; L rosulate, deltoid-lanceolate to linear, 1.5–3  2–4.5 cm, falcate and curving upwards, sometimes twisted and unevenly parallel, upper face flat to shallowly canaliculate, plane towards the apex, faintly to densely white-spotted, lower face

762

somewhat convex with a distinct eccentric keel, faintly to densely spotted, spots arranged in obscure transverse bands, both faces shiny, light to dark green, with smooth epidermis, margins minutely denticulate to almost entire, apex acute or subacute, sometimes acuminate, mucronate; juvenile L distichous, lorate, patent to ascending, epidermis similar to adult leaves, densely white-spotted, spots arranged in transverse bands, apex obtuse, mucronate; Inf racemose, to 70 cm, flowers secundly arranged and open at the same time, bracteate; floral Bra 7  2 mm, piliferous; Ped 5 mm, pink; Per 20–21 mm, gasteriform, stipitate for 2 mm; Tep fused except for the tips and margins of the inner ones, perianth tube subcylindrical, curved in the upper , the lower slightly decurved, inflated to 5–6 mm ∅, pink, upper white with green striations, tips erect becoming erectly spreading, obtuse; St 10–12 mm; Anth 2  1 mm, included; Ov 5–6  2–3 mm ∅, green; Sty 7 mm; Sti included, minute; Fr 22–30  6–8 mm ∅, narrowly obovoid, triangular in cross-section, obtuse at the apex; Se blackish. G. loedolffiae Van Jaarsveld (Bradleya 32: 44–46, ills. (pp. 45–48), 2014). Type: RSA, Eastern Cape (Van Jaarsveld & al. 24626 [NBG]). — Distr: RSA (Eastern Cape); sheer shale cliffs, Eastern Valley Bushveld vegetation, 270 m, summer-flowering. [3] Acaulescent, decumbent to ascending, to 38 cm ∅, solitary or proliferating from the base to form clusters of 3–5 heads; L rosulate (but distichous in juvenile plants), first ascending, then patent-spreading firm, lorate-lanceolate, 16.5–23  6–6.5 cm broad at the base, slightly falcate, adaxial face canaliculate, faintly whitespotted (spots in transverse bands in juvenile plants), abaxial face somewhat convex with a distinct asymmetrical keel, faintly spotted, both faces dull green with smooth, slightly minutely asperulous epidermis, margins minutely serrulate to almost entire, apex obtuse to subacute, mucronate; Inf ascending to flat-topped panicles (racemose in juvenile plants), to 1.15 m; Ped 4 mm, pink; Per 27–29 mm, stipitate for 3–4 mm, subcylindrical, curved in the middle, upper inflated

E. Van Jaarsveld

to 7 mm ∅; Tep fused for the greater part of their length, uniform dull pink; St 23–25 mm; Anth 3  1.5 mm, included; Ov 6.5  3 mm ∅, green; Sty 13 mm; Sti included, curved upwards, minute; Fr 16–25  7–10 mm, narrowly obovoid, triangular in cross-section, with obtuse apex; Se oblong, 5  3 mm, black. G. nitida (Salm-Dyck) Haworth (Philos. Mag. Ann. Chem. 1827: 359, 1827). Type: [neo — icono]: Salm-Dyck, Monogr. Gen. Aloes & Mesembr., fasc. 4, Aloe sect. 29, t. 17, 1842. — Distr: RSA (Eastern Cape); Kouga Sandstone Fynbos and Bisho Thornveld vegetations, flowers in mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 47).  Aloe nitida Salm-Dyck (1817); incl. Aloe nitida var. major Salm-Dyck (1817); incl. Aloe nitida var. minor Salm-Dyck (1817); incl. Aloe nitida var. obtusa Salm-Dyck (1817)  Gasteria obtusa (Salm-Dyck) Haworth (1827); incl. Aloe nitida var. grandipunctata Salm-Dyck (1821) (nom. illeg., Art. 52.1)  Gasteria nitida var. grandipunctata (Salm-Dyck) A. Berger (1908) (nom. illeg., Art. 52.1); incl. Aloe nitida var. parvipunctata Salm-Dyck (1821) (nom. illeg., Art. 52.1)  Gasteria nitida var. parvipunctata (Salm-Dyck) A. Berger (1908) (nom. illeg., Art. 52.1); incl. Aloe trigona var. obtusa Salm-Dyck (1821) (nom. illeg., Art. 52.1); incl. Aloe nitida Ker Gawler (1822) (nom. illeg., Art. 53.1); incl. Haworthia nigricans Haworth (1824); incl. Gasteria decipiens Haworth (1827)  Aloe decipiens (Haworth) Roemer & Schultes (1829); incl. Gasteria beckeri Schönland (1907); incl. Gasteria stayneri Von Poellnitz (1938). [4] Acaulescent, decumbent to erect, 6–20  5–28 cm, solitary or proliferating from the base to form small groups; R somewhat fusiform, succulent, to 12 mm ∅; L first distichous, becoming rosulate, 1.6–18  2.5–8 cm, triangular-lanceolate, rarely lanceolate-acuminate, erectly spreading with a distinct asymmetrical keel, both faces dark green, with faint to dense white spots arranged in irregular transverse bands, epidermis smooth and shiny, margin entire or indistinctly tuberculate, tip acute, mucronate; juvenile L distichous, lorate, 2–4  2–3.5 cm, epidermis

Gasteria ASPHODELACEAE

763

tuberculate, rarely smooth, dark green, not or rarely spotted; Inf erectly spreading branched lax panicles 0.2–1.2 m; Per 20–25 mm, bright reddish-pink, stipitate for 2–3 mm, gasteriform for slightly > of the perianth length, 5–8 mm ∅, narrowly ellipsoid, constricted above into a tube 4–5 mm ∅, lobes erectly spreading, obtuse, yellowish; Sty included; Fr oblong, 24–30  8 mm; Se 3–4  2 mm. G. pillansii Kensit (Trans. Roy. Soc. South Africa 1: 163, 1909). Type: RSA, Northern Cape (Pillans 833 [BOL, PRE]). — Distr: S Namibia, NW RSA (Northern Cape). G. pillansii var. ernesti-ruschii (Dinter & Von Poellnitz) Van Jaarsveld (Aloe 29(1): 17, 1992). Type: [lecto — icono]: Kakt. and. Sukk. (Berlin), 1938(2): ill. p. 36. — Distr: S Namibia, RSA (Northern Cape); Succulent Karoo vegetation, rocks, flowers mid-summer to autumn. I: Jaarsveld & Ward-Hilhorst (1994: 61). – Fig. 5.  Gasteria ernesti-ruschii Dinter & Von Poellnitz (1938). [2] Differs from var. pillansii: L 2–7 cm; Inf 6–30 cm; Per 25–30 mm; St shortly exserted. G. pillansii var. hallii Van Jaarsveld (Aloe 44 (4): 94, ill., 2008). Type: RSA (Van Jaarsveld & Duncan 7912 [NBG]). — Distr: RSA (Northern Cape: Richtersveld); succulent shrubland, rocks, autumn-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 2, as G. pillansii var. ernesti-ruschii). [2] Differs from var. pillansii: L 2–4  1.5–2 cm; Fl shorter and in autumn; St shortly exserted. G. pillansii var. pillansii — Distr: RSA (Northern Cape: Namaqualand); Succulent Karoo vegetation, flowers in mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 59). – Figs. 6 and 7. Incl. Gasteria neliana Von Poellnitz (1930). [2] Acaulescent, decumbent to erect, 5–20  6–40 cm, proliferating from subterranean stolons and forming dense groups; L distichous, 2–20  1.5–5 cm, lorate, erectly spreading or patent, both faces spotted with immersed tubercles in obscure transverse bands, epidermis asperulous, rarely with few domed tubercles, margin cartilaginous,

Fig. 5 Gasteria pillansii var. ernesti-ruschii. (Copyright: U. Eggli)

tuberculate or crenulate, tip obtuse to acute, mucronate; Inf racemose, 1–3 per plant, spreading and slightly curved, 0.6–1.2 ( 1.65) m, rarely with a pair of branches; Fl laxly arranged in the upper ; Per 25–50  6–8 mm, stipitate for 3 mm, obscurely gasteriform basally for or less of the length, globose-ellipsoid to slightly constricted above the ovary, above gradually enlarging and rarely clavate, gasteriform part pink, tube white with green striations; St included or exserted to 5 mm; Fr oblong, 15–23  7 mm; Se 4–5  2.5 mm. G. polita Van Jaarsveld (Cact. Succ. J. (US) 73 (3): 127–129, ills., 2001). Type: RSA, Western Cape (Van Jaarsveld & Kok 13742 [NBG]). — Distr: RSA (Western Cape); Tsitsikamma Sandstone Fynbos vegetation, rocky outcrops, springflowering. I: Retief (2015). [4] Acaulescent, solitary or proliferating to form small groups, 12  25 cm; L rosulate, ascending,

764

E. Van Jaarsveld

Fig. 6 Gasteria pillansii var. pillansii. (Copyright: E. Van Jaarsveld)

Fig. 7 Gasteria pillansii var. pillansii. (Copyright: E. Van Jaarsveld)

becoming spreading, strap-shaped, 6–12  3–4 cm, apex obtuse to subacute, mucronate, adaxial face canaliculate, abaxial face somewhat convex with a distinct asymmetrical keel, both faces dark green mottled with white spots in transverse pattern, epidermis smooth, shiny, margin tuberculatecrenate; juvenile L distichous, lorate, patent to ascending, smooth and shiny, apex obtuse, mucronate; Inf racemose, to 60 ( 100) cm, spreading, simple or sometimes with 2 branches; Per 35 mm, stipitate for 4 mm, gasteriform basally, narrowly ellipsoid over of the length, pink, upper white with green striation; Fr oblong, 24–25  4–6 mm; Se 4–5  2 mm, black. A new population with overall larger plants (inflorescences to 1 m) with somewhat earlier flowering time was recently discovered (Baard 2012a, Baard 2012b).

form small groups; L in a dense rosette, 24–36  2.5–4 cm, erectly spreading, often falcate, linearensiform to linear-acuminate, with an indistinct asymmetrical keel, both faces dark green with dense white spots in transverse bands, epidermis smooth, margin cartilaginous, serrulate, tip acute or acuminate, mucronate; juvenile L distichous, erectly spreading, lorate, acuminate, distinctly tuberculate, mucronate; Inf erect to erectly spreading lax panicles, 0.35–1.5 m, rarely simple or side branches erectly spreading; Per 18–25 mm, stipitate for 2 mm, gasteriform part > of the perianth length, globose-ellipsoid, reddish-pink, 6–7 mm ∅, then constricted into a tube of 4–4.5 mm ∅, light to dark pink, lobes white, rarely pink, erectly spreading, obtuse, with green median stripes; Sty included; Fr oblong, 12–27  7 mm; Se 2–5  2–3 mm.

G. pulchra (Aiton) Haworth (Synops. Pl. Succ., 86, 1812). Type: [lecto — icono]: Miller, Gard. Dict., 1759, t. 292. — Distr: RSA (Eastern Cape); Gamtoos and Groot Thicket vegetations, on conglomerate, spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 45).  Aloe maculata var. pulchra Aiton (1789)  Aloe pulchra (Aiton) Haworth (1804); incl. Aloe obliqua De Candolle (1802) (nom. illeg., Art. 53.1); incl. Aloe pulchra Jacquin (1805) (nom. illeg., Art. 53.1); incl. Gasteria poellnitziana H. Jacobsen (1954) (nom. inval., Art. 36.1). [4] Acaulescent, decumbent to erect, 20–36  20–36 cm, solitary or proliferating from the base to

G. rawlinsonii Obermeyer (Flow. Pl. Afr. 43 (3/4): t. 1701 + text, 1976). Type: RSA, Eastern Cape (Rawlinson s.n. [PRE 34421]). — Distr: RSA (Eastern Cape); Groot Thicket vegetation, sheer shady quartzitic sandstone cliff faces; spring-flowering. I: Jaarsveld & Ward-Hilhorst (1994: 79). [1] Caulescent, pendulous, proliferating from the base; stems leafy, to 1 m, rarely branched, internodes 1–2 cm; L distichous or rosulate, 3–8  1–2.5 cm, linear, lorate, slightly falcate, tips recurved, lower face convex, without keel, both faces green, unspotted or with faint white spots, epidermis asperulous, margin sparingly denticulate, or sometimes unarmed, prickles

Gasteria ASPHODELACEAE

765

part narrowly to globosely ellipsoid, making up > of the perianth length, 5–8 mm ∅, pink, above constricted to a tube 4 mm ∅, white with green striation; Sty included; Fr oblong, 19–23  7 mm; Se 3–4  2 mm.

Fig. 8 Gasteria retusa. (Copyright: E. Van Jaarsveld)

turning black with age, tip obtuse, mucronate; Inf racemose, 10–50 cm; Bra 5  2 mm at the base; Per variable, 16–25 mm, stipitate for 1–3 mm, gasteriform part > of the length, globoseellipsoid or globose, pink, then constricted into a tube 4–6 mm ∅, tube pink or white, occasionally with green striations; Fr 18 mm, oblong-ovoid; Se 3–4 mm. A slow growing cliff-dweller with two forms grading into each other: One has the leaves remaining distichous and short inflorescences; the other is with rosulate leaves and longer inflorescences and a globose gasteriform perianth part. G. retusa (Van Jaarsveld) Van Jaarsveld (Aloe 44(4): 100, ills., 2008). Type: RSA, Western Cape (Van Jaarsveld & Stayner 4656 [NBG]). — Distr: RSA (Western Cape): Robertson Karoo vegetation, on shale, spring-flowering. I: Jaarsveld & Weber (1994: as G. carinata var.). – Fig. 8.  Gasteria carinata var. retusa Van Jaarsveld (1992). [4] Acaulescent, decumbent to erect, proliferating from the base to form small groups 3–17  10–25 cm; L distichous, 3–15 ( 17)  1–3.5 cm, erectly spreading, lorate, both faces light to dark green, with raised semitranslucent white dome-shaped tubercles in transverse bands, margin cartilaginous often undulating, tuberculate-crenulate, rarely denticulate, tip retuse, truncate or obtuse, mucronate; juvenile L similar to adult leaves; Inf simple ascending racemes, 15–45 cm; Ped 8–9 mm; Per 17–22 mm, stipitate for 1–2 mm, gasteriform

G. thunbergii N. E. Brown (Bothalia 1: 140, 1923). Type: RSA, Western Cape (Thunberg 8595 [UPS]). — Distr: RSA (Western Cape: Gouritz River Valley); Southern Cape Valley Thicket vegetation, on conglomerate, flowers mainly in autumn. I: Jaarsveld (2008: 93).  Gasteria carinata var. thunbergii (N. E. Brown) Van Jaarsveld (1998);  Aloe disticha Thunberg (1785) (nom. illeg., Art. 53.1). [4] Acaulescent, decumbent to erect, proliferating from the base to form small groups 6–20  10–30 cm; L distichous, 3–20  1–3 cm, ascending, acerose-subulate to linear-triangular, upper face deeply channelled, both faces green, with raised white to greenish tubercles in transverse bands, margin cartilaginous, denticulate, tip mucronate; juvenile L similar to adult leaves; Inf simple ascending racemes, 30–45 cm; Per variable, 18–20 mm, gasteriform at the base for  of the perianth length, narrowly ellipsoid, pink, lobes white, rarely pink, erectly spreading, obtuse, with green median stripes; Sty included; Fr 20–22  6 mm; Se oblong, 6  2 mm. G. tukhelensis Van Jaarsveld (Bothalia 35(2): 164–165, ills., 2005). Type: RSA, KwaZuluNatal (Van Jaarsveld & al. 17996 [NBG]). — Distr: RSA (KwaZulu-Natal); cliffs along the Tukhela River, in Valley Bushveld vegetation, summer-flowering. [3] Acaulescent, decumbent, to 25  70 cm ∅; L rosulate, deltoid-lanceolate, 12–25  3–5 cm, falcate and curving upwards, adaxial face canaliculate, abaxial face somewhat convex with an asymmetrical keel, faintly spotted in obscure transverse bands, both surfaces shiny, dark green, margin minutely denticulate to almost entire, apex acute to subacute, mucronate; juvenile L distichous, lorate, patent to ascending, epidermis tuberculate; Inf racemose, to 56 cm, with up to 11 secundly arranged pendent flowers; Per pink, 40–43 mm, curved in the middle, basally only

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very weakly elongate-gasteriform; Fr 23–32 mm; Se 5–7  2–3 mm. G. vlokii Van Jaarsveld (Cact. Succ. J. (US) 58 (4): 170–174, ills., 1987). Type: RSA, Western Cape (Vlok 880 [NBG]). — Distr: RSA (Western Cape); South Swartberg Sandstone Fynbos vegetation, quartzitic sandstones, flowers in mid-summer. I: Jaarsveld & Ward-Hilhorst (1994: 53). [4] Acaulescent, proliferating from the base to form small groups to 14 cm ∅; R succulent, to 8 mm ∅, fusiform; L distichous, ultimately becoming rosulate, 5–9  2–3 cm broad at the base, patent, lorate, lanceolate to triangular, falcate, with an asymmetrical keel in the upper , both faces green with dense white spots in obscure transverse bands, epidermis asperulous, margin acute, tuberculate, crenulate, becoming continuous towards the tip, tip acute or obtuse with an asymmetrical mucro; juvenile L lorate, tip obtuse, mucronate; Inf racemose, curved and spreading, 30–84 cm, rarely with a pair of branches; Per 29–33 mm, stipitate for 2–3 mm, gasteriform part slightly > of the perianth length, 6–7 mm ∅, dark reddish-pink, above constricted into a tube of 4 mm ∅, lobes white with green median striations; Fr 15–18  6 mm; Se oblong, 3–5  2 mm.

References Aymerich, P. & Gustamante, L. (2015) Nuevas citas de plantas alóctonas de origen ornamental en el litoral meridional de Cataluña. Bouteloua 20: 22–41, ills. Baard, J. (2012a) Discovery on the Garden Route. Another population of the rare Gasteria polita found. Veld Fl. (1975+) 98(1): 20–21, ills., map. Baard, J. (2012b) Gasteria polita, an endangered succulent from South Africa. CactusWorld 30(3): 159–161, ills., map. Crook, I. [& al. 2009], Minors, D. M. & Minors, C. J. (2009) Observations on the seed dispersal strategy of Gasteria excelsa. Haworthiad 23(2): 52–54, ills. Crouch, N. R. [& al. 2000], Smith, G. F., Symmons, R. & Tomalin, M. (2000) Gasteria croucheri — the magic impundu of the Zulu. Brit. Cact. Succ. J. 18(2): 70–78, ills., map. Crouch, N. R. [& al. 2013], Smith, G. F. & Condy, G. (2013) Gasteria croucheri subsp. pondoensis. Flow. Pl. Afr. 63: 22–30, t. 2284, ill., map.

E. Van Jaarsveld Cumming, D. (2003) The army needs you! A matter of recruitment in Gasteria excelsa Baker. Alsterworthia Int. 3(3): 21. Daru, B. H. [& al. 2013], Manning, J. C., Boatwright, J. S., Maurin, O., Mclean, N., Schaefer, H., Kuzmina, M. & Bank, M. van der (2013) Molecular and morphological analysis of subfamily Alooideae (Asphodelaceae) and the inclusion of Chortolirion in Aloe. Taxon 62(1): 62–76, ills. URL: http://www.jstor.org/stable/24389313. Grace, O. M. & Rønsted, N. (2013) Unlocking the secrets of DNA in the classification of aloes. Aloe 50(1-2): 22–23. Jaarsveld, E. J van (1992) The genus Gasteria, a synoptic review. Aloe 29(1): 5–30, ills., maps, key, (2): 49 [erratum]. Jaarsveld, E. J van (1998) A new taxon and new combinations in the Gasteria carinata complex. Cact. Succ. J. (US) 70(2): 65–71, ills., map, key. Jaarsveld, E. J van (2008) The genus Gasteria: A synoptic review (new taxa and combinations). Aloe 44(4): 84–103, ills. Jaarsveld, E. J van (2009) New Gasteria cultivars. Aloe 45(3): 52–55, ills. Jaarsveld, E. J van & Thomas, V. (2003) Gasteria glauca. Flow. Pl. Afr. 58: 16–21, t. 2184, map. Jaarsveld, E. J van & Ward-Hilhorst, E. (1994) Gasterias of South Africa. A new revision of a major succulent group. Vlaeberg (ZA): Fernwood Press in association with the National Botanical Institute. Jaarsveld, E. J van & Weber, R. (1994) Gasteria carinata var. retusa. Flow. Pl. Afr. 53: 86–89, t. 2097, map. Manning, J. C. [& al. 2014a], Boatwright, J. S., Daru, B. H., Maurin, O. & Bank, M. van der (2014) A molecular phylogeny and generic classification of Asphodelaceae subfamily Alooideae: A final resolution of the prickly issue of polyphyly in the Alooids. Syst. Bot. 39(1): 55–74, key. https://doi.org/10.1600/03636 4414X678044 Mays, H. (2009) Hybrids & cultivars of the succulent Asphodelaceae. Volume 1. Aloe, Astroloba, Bulbine, Gasteria & nothogenera. St. Michaels on Wyre (GB): Harry Mays. Raimondo, D. [& al. 2009], Staden, L. von, Foden, W., Victor, J. E., Helme, N. A., Turner, R. C., Kamundi, D. A. & Manyama, P. A. (eds.) (2009) Red List of South African plants 2009. Pretoria (ZA): South African National Biodiversity Institute SANBI (= Strelitzia No. 25). Retief, K. (2015) Gasterias at Nature’s Valley. Aloe 51(2): 41–45, ills. Zonneveld, B. J. M. & Jaarsveld, E. J van (2005) Taxonomic implications of genome size for all species of the genus Gasteria Duval (Aloaceae). Pl. Syst. Evol. 251: 217–227, maps. https://doi.org/10.1007/s00606-0040244-x Zukulu, S. [& al. 2012], Dold, T., Abbott, T. & Raimondo, D. (2012) Medicinal and charm plants of Pondoland. Pretoria (ZA): South African National Biodiversity Institute SANBI.

Gastroloba ASPHODELACEAE U. Eggli

Gastroloba D. M. Cumming (Bull. Afr. Succ. Pl. Soc. 9: 36, 1974). — Alooideae.  Gasteria  Astroloba. Rowley (1982: 76) lists two known combinations, one of them formally named as G. apicroides (Baker) G. D. Rowley ( Gasteria apicroides Baker). Two further cultivars are described and illustrated by Mays (2009: 59).

References Mays, H. (2009) Hybrids & cultivars of the succulent Asphodelaceae. Volume 1. Aloe, Astroloba, Bulbine, Gasteria & nothogenera. St. Michaels on Wyre (GB): Harry Mays. Rowley, G. D. (1982) Intergeneric hybrids in succulents. Nation. Cact. Succ. J. 37(1): 2–6, (2): 45–49, (3): 76–80, (4): 119, ills.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_68

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Haworthia ASPHODELACEAE M. B. Bayer and E. Van Jaarsveld

Haworthia Duval (Pl. Succ. Horto Alencon., [7], 1809). Type: Aloe pumila var. arachnoidea Linné. — Alooideae — Lit: Bayer (1982: ill. synopsis); Scott (1985: ill. synopsis); Breuer & Metzing (1997: typification); Breuer (1998a: bibliography); Breuer (1998b: synopsis); Bayer (1999: revision); Battista (2009: ill. synopsis); Schulz (2009: ill. synopsis); Mays (2013: cultivar synopsis); Zonneveld (2015: sectional classification); Gildenhuys & Klopper (2016: classification Haworthiopsis); Gildenhuys (2017: illustrated synopsis Haworthiopsis). Distr: S Namibia, RSA (predominantly Western Cape, Eastern Cape, but also Northern Cape, Free State, KwaZulu-Natal and Mpumalanga), Swaziland, S Moçambique; mainly Succulent Karoo, Nama Karoo and Albany Thicket vegetations, and to a lesser extent savanna, mostly in rocky places in the shade of grasses or shrubs. Etym: For Adrian H. Haworth (1768–1833), English zoologist and botanist and succulent plant specialist. Incl. Catevala Medikus (1786) (nomen rejiciendum, Art. 56.1). Type: Aloe retusa Linné [Pfeiffer,

M. B. Bayer (*) Hermanus, South Africa e-mail: [email protected] E. Van Jaarsveld Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa e-mail: [email protected]

Nomencl. Bot., 1871–74 (cf. P. V. Heath, Calyx 4(3): 83, 1994)]. Incl. Apicra Willdenow (1811). Type: not typified. Incl. Aprica D. Dietrich (1840). Type: not typified. Incl. Tulista Rafinesque (1840). Type: Aloe pumila var. margaritifera Linné. Incl. Haworthiopsis G. D. Rowley (2013). Type: Haworthia coarctata Haworth. Incl. Hawiopsis G. D. Rowley (2014). Incl. Tulworthia G. D. Rowley (2014). Incl. Tulworthiopsis G. D. Rowley (2014). Dwarf rosulate perennials, acaulescent to caulescent, solitary or proliferating to form dense tight clusters; R fibrous to succulent, sometimes fusiform; Ros sessile or caulescent, 2–15 cm ∅, flat to globose or elongate; L erect, spreading to recurved, sometimes highly reduced, laxly to densely spirally arranged, rarely distichous or somewhat tristichous, triangular-lanceolate to linear, usually firm, often drying back from the tip and becoming papery, surfaces often tuberculate and variously marked, glabrous to rarely pubescent, often with translucent windows, and with various patterns, markings and reticulations, very pale green, blue-green to dark (blackish-) green, upper face flat, or channelled to convex, lower face convex, often keeled towards the tip, margin entire, denticulate, ciliate, with bristly Sp or tuberculate, tip acute, or acuminate to mucronate, sometimes retuse or truncate; Inf 1–5 racemes from near the rosette centre, erect, simple or branched, firm to wiry, bracteate, peduncle brownish-green, often

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_56

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with a powdery bloom; Bra small; Fl small, zygomorphic, bilabiate, 5–50 per inflorescence, 15 mm, tube curved or straight, slightly swollen towards the base; Tep 6, white to pink, often with central green or brownish striations, tips spreading to recurved with green, brownish or occasionally yellow median colouration; Fil 6, included; Ov oblong to oblong-ovoid, 3-chambered, with 6 grooves; Sty subulate; Sti apical, minute, trilobate or rounded; Fr erect oblong-ovoid woody capsules, dehiscing longitudinally; Se black to grey, compressed, irregularly angled or flattish. Classification: Treutlein & al. (2003b) found evidence that Haworthia in the traditional circumscription is not monophyletic, and this was corroborated by Ramdhani & al. (2009) and Ramdhani & al. (2011), as well as Daru & al. (2013) and Manning & al. (2014a) in their detailed studies of the whole subfamily Alooideae. These last-mentioned authors recovered three well-supported clades of Haworthia that conform to the traditionally accepted subgenera, with the exception of H. koelmaniorum, which occupies an enigmatic position separate from all other Haworthias. The typical subgen. Haworthia has been found in a polytomy amongst Aloe sect. Kumara, Aloe sect. Macrifolia and a clade composed of the “true Aloes” and the remaining alooid genera. Subgen. Robustipedunculares is sister a clade that includes Aloe aristata, A. variegata and the genus Astroloba, and subgen. Hexangulares is sister to Gasteria. Rowley (2013a) formalized this phylogeny by accepting the genus Tulista for subgen. Robustipedunculares, and by describing the new genus Haworthiopsis for subgen. Hexangulares. Tulista in the sense of Rowley (l.c.) also includes Astroloba, Poellnitzia and Aloe aristata, a circumscription that is based on the phylogeny presented by Daru & al. (2013). Manning & al. (2014a) also accept the break-up of Haworthia into the three genera just mentioned, but recircumscribe Tulista by excluding Aloe aristata (recognized as monotypic genus Aristaloe). The cytogenetic work by Zonneveld (2015) als supports breaking up Haworthia s.l. Editorial comment by U. Eggli: Despite the good support for many branches in the phylogenies of Daru & al. (2013) and Manning & al. (2014a), it

M. B. Bayer and E. Van Jaarsveld

is argued that it is still premature to split Haworthia s.l., until a full and complete picture of the alooid genera is available — even though a good proportion of Aloe species have already been included in the phylogenetic studies cited above, taxa so far omitted could well cause surprising new interpretations when they are finally also investigated. Haworthia s.l. has its centre of diversity in the Western Cape. Species definition is notoriously difficult because many species are highly variable and phenotypically plastic. To account for this Bayer (2002a: 59), Vosa (2005) and Bayer (2007a: 162–164) formulated a “superspecies” concept. It is well known that there are few reproductive barriers and natural hybrids are common within all three subgenera. In addition both H. pumila and H. minima hybridize naturally with Astroloba corrugata. [Editorial comment by U. Eggli: In fact, Ramdhani & al. (2011) proposed ancient hybridization (also with species from other alooid genera) to explain the phylogeny that resulted from their DNA study. These authors also found evidence that interspecific hybridization is still on-going, esp. in subgen. Haworthia, and they argue that the absence of reproductive barriers supports such an assumption]. It is commonly known that environmental and growing conditions determine physical appearance of Haworthias to a great degree, and Bayer & Dold (2003) discuss the role of geology and habitat in determining ecotypes. The difficulties to diagnostically circumscribe taxa in Haworthia have led to differing views on how many and which species should be recognized, culminating in the vast over-splitting as advocated by Hayashi (2008) (4 subgenera, 546 species of which 234 were only provisionally named) or Breuer (2010) and Breuer (2011) (3 subgenera, 366 species in 108 “aggregates”, also with numerous provisional names). Here, species definitions take into account morphological data supplemented with data from geographical distribution, with due notion of sympatric occurrences. We follow the classification concept of Bayer & Manning (2012a) and Bayer & Manning (2012b), which are based on the earlier synoptical work of Bayer (1982), the monograph of Bayer (1999), and subsequent new evidence

Haworthia ASPHODELACEAE

(Bayer 2002a, Bayer 2006, Bayer 2007a, Bayer 2008, Bayer 2009, Bayer 2010a, Bayer 2012a, Bayer 2012b). The following three subgenera are recognized: [1] Subgen. Haworthia: Base of the Fl triangular or rounded-triangular, tube obclavate, curved; OTep free; Sty upcurved; Se irregularly angled. — 37 species. [2] Subgen. Hexangulares Uitewaal ex M. B. Bayer 1971: Base of the Fl 6-angular, gradually narrowing to the junction with the Ped; tube obcapitate, curved; OTep partly fused to the ITep; Sty straight; Se irregularly angled. — 17 species. [3] Subgen. Robustipedunculares Uitewaal ex M. B. Bayer 1971: Base of the Fl 6-angular, rounded, abruptly joined to the Ped; OTep partly fused to the ITep, tube obcapitate, straight; Sty straight; Se flattish. — 4 species. Ecology: North & al. (2008) report contractile roots for Haworthia, and this helps to explain the growth habit of the retuse-leaved species. Contraction is the result of radial expansion and longitudinal shrinkage of cells in the inner and/or middle root cortex layers. Pollination Biology: Smith & al. (2002) found an interesting difference in nectar sugar concentrations: Species of subgen. Haworthia have 60% sucrose. The significance of this difference is unknown, as our knowledge of pollinators of Haworthia is very incomplete. Bayer (2013b) reports an unidentified anthophorid solitary bee as frequent visitor, and Bayer (2013c) presents photographs of the bombylid solitary bee fly Australoechus hirtus on Haworthia flowers. In addition, a lycaenid butterfly is also sometimes visiting Haworthia flowers (Bayer, pers. obs.) Grenier (2013) also illustrates Australoechus and in addition Amegilla as flower visitors of Haworthia species and observed that pollinators do not discriminate between three concurrently flowering species. Marx (2013b) observed a tabanid horse-fly

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(probably Philoliche aethiopica) as regular pollinator of H. arachnoidea. Horticulture: Haworthias are popular amongst succulent plant collectors, and many can tolerate relatively shaded growing conditions. Propagation is usually from offsets, but also possible from detached whole leaves or in some taxa even from detached roots, as well as from seed or via tissue culture. In the list of names of unresolved application below, many are of horticultural derivation. Over the course of the years, numerous forms have been selected, propagated, and named as cultivars, esp. in Japan (Hayashi 2013), where special oddities are highly prized. Some species occasionally produce variegated offspring, and some of these have also been formally named as cultivars. Hildyard (2016a) has a short synopsis on variegated forms in the genus. Acknowledgment: We are indebted to Sean Gildenhuys for his help with placing some of the synonyms. The following names are of unresolved application but are referred to this genus: Aloe anomala Haworth (1804)  Apicra anomala (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4); Aloe aspera Haworth (1804)  Apicra aspera (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Astroloba aspera (Haworth) Uitewaal (1947); Aloe cylindrica Lamarck ex Poiret (1810); Aloe cylindrica var. rigida Lamarck (1783)  Aloe rigida (Lamarck) De Candolle (1799)  Apicra rigida (Lamarck) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthia rigida (Lamarck) Haworth (1821)  Haworthiopsis rigida (Lamarck) Gildenhuys & Klopper (2016); Aloe expansa Haworth pro sp. (1804)  Haworthia expansa (Haworth) Haworth (1812)  Aloe rigida var. expansa (Haworth) Salm-Dyck (1836)  Haworthia rigida var. expansa (Haworth) Baker (1880)  Haworthiopsis expansa (Haworth) Gildenhuys & Klopper (2016); Aloe expansa var. major Haworth (1804); Aloe glabrata var. concolor Salm-Dyck (1849)  Haworthia

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glabrata var. concolor (Salm-Dyck) Baker (1880); Aloe glabrata var. perviridis Salm-Dyck (1849)  Haworthia glabrata var. perviridis (Salm-Dyck) Baker (1880); Aloe hybrida SalmDyck pro sp. (1817)  Haworthia hybrida (Salm-Dyck) Haworth (1821)  Haworthiopsis hybrida (Salm-Dyck) Gildenhuys & Klopper (2016); Aloe pseudorigida Salm-Dyck pro sp. (1817)  Apicra pseudorigida (Salm-Dyck) Haworth (1819)  Haworthia pseudorigida (SalmDyck) Haworth (1819)  Haworthia tortuosa var. pseudorigida (Salm-Dyck) A. Berger (1908)  Haworthiopsis pseudorigida (Salm-Dyck) Gildenhuys & Klopper (2017); Aloe pumila Thunberg (1785) (nom. illeg., Art. 53.1); Aloe radula var. major Salm-Dyck (1817); Aloe radula var. media Salm-Dyck (1817); Aloe radula var. minor Salm-Dyck (1817); Aloe rigida var. paulo-major Salm-Dyck (1817); Aloe rugosa Salm-Dyck (1834)  Haworthia rugosa (Salm-Dyck) Baker (1880); Aloe rugosa var. laetevirens Salm-Dyck (1834); Aloe rugosa var. perviridis Salm-Dyck (1834)  Haworthia rugosa var. perviridis (Salm-Dyck) A. Berger (1908); Aloe semimargaritifera var. glabrata Salm-Dyck (1834); Aloe semimargaritifera var. major Salm-Dyck (1817)  Haworthia semimargaritifera var. major (Salm-Dyck) Haworth (1819)  Haworthia margaritifera subvar. major (Haworth) A. Berger (1908); Aloe semimargaritifera var. minor SalmDyck (1817)  Haworthia semimargaritifera var. minor (Salm-Dyck) Haworth (1819); Aloe semimargaritifera var. multipapillosa Salm-Dyck (1834)  Haworthia margaritifera subvar. multipapillosa (Salm-Dyck) A. Berger (1908); Aloe subalbicans Salm-Dyck (1854)  Haworthia margaritifera var. subalbicans (Salm-Dyck) A. Berger (1908); Aloe subalbicans var. acuminata Salm-Dyck (1854)  Haworthia margaritifera subvar. acuminata (Salm-Dyck) A. Berger (1908); Aloe subalbicans var. laevior Salm-Dyck (1854)  Haworthia margaritifera subvar. laevior (Salm-Dyck) A. Berger (1908); Aloe subattenuata Salm-Dyck (1834)  Haworthia subattenuata (Salm-Dyck) Baker (1880); Aloe subfasciata Salm-Dyck (1825) (nom. illeg., Art. 52.1)  Haworthia subfasciata (Salm-Dyck) Baker (1880) (nom. illeg., Art. 52.1); Aloe subrigida

M. B. Bayer and E. Van Jaarsveld

Roemer & Schultes fil. pro sp. (1829)  Haworthiopsis subrigida (Roemer & Schultes fil.) Gildenhuys & Klopper (2016); Aloe subulata Salm-Dyck (1822); Aloe subulata Salm-Dyck (1829) (nom. illeg., Art. 52.1?)  Haworthia subulata (Salm-Dyck) Baker (1880); Aloe tortuosa var. major Salm-Dyck (1837)  Haworthia tortuosa var. major (Salm-Dyck) Baker (1896)  Haworthiopsis major (Salm-Dyck) Gildenhuys & Klopper (2016); Apicra aspera var. major Haworth (1819) (incorrect name, ICN Art. 11.4)  Astroloba aspera var. major (Haworth) Uitewaal (1947)  Haworthia aspera var. major (Haworth) Parr (1971); Apicra patula Willdenow (1811) (incorrect name, ICN Art. 11.4); Haworthia affinis Baker (1880)  Haworthia bilineata var. affinis (Baker) Von Poellnitz (1938); Haworthia altilinea Haworth (1824)  Aloe altilinea (Haworth) Roemer & Schultes fil. (1829)  Haworthia mucronata var. altilinea (Haworth) Halda (1997); Haworthia altilinea fa. acuminata Von Poellnitz (1938)  Haworthia mucronata fa. acuminata (Von Poellnitz) Von Poellnitz (1940); Haworthia altilinea fa. minor Triebner (1938)  Haworthia mucronata fa. minor (Triebner) Von Poellnitz (1940); Haworthia altilinea fa. subglauca Von Poellnitz (1938)  Haworthia mucronata fa. subglauca (Von Poellnitz) Von Poellnitz (1940); Haworthia altilinea var. bicarinata Triebner (1938)  Haworthia mucronata var. bicarinata (Triebner) Von Poellnitz (1940); Haworthia altilinea var. brevisetata Von Poellnitz (1937); Haworthia altilinea var. setulifera Triebner & Von Poellnitz (1938)  Haworthia mucronata var. setulifera (Triebner & Von Poellnitz) Von Poellnitz (1940); Haworthia altilinea var. typica Von Poellnitz (1937) (nom. inval., ICN Art. 24.3); Haworthia argyrostigma Baker (1896) (nom. inval., Art. 32.1)  Haworthia attenuata var. argyrostigma (Baker) A. Berger (1908) (nom. inval., Art. 32.1); Haworthia aspera Haworth (1812); Haworthia asperula Haworth (1824)  Aloe asperula (Haworth) Roemer & Schultes fil. (1829)  Haworthia retusa ssp. asperula (Haworth) Halda (1997)  Haworthia retusa var. asperula (Haworth) Halda (1997)  Haworthia magnifica var. asperula (Haworth) Breuer (2016) (incorrect name, ICN Art. 11.4); Haworthia baccata G. G. Smith (1944); Haworthia

Haworthia ASPHODELACEAE

bijliana Von Poellnitz (1929)  Haworthia setata var. bijliana (Von Poellnitz) Von Poellnitz (1938); Haworthia bilineata Baker (1880); Haworthia cassytha Baker pro sp. (1896)  Haworthiopsis cassytha (Baker) Gildenhuys & Klopper (2016); Haworthia coarctata var. sampaiana Resende (1938)  Haworthia sampaiana (Resende) Resende pro sp. (1940)  Haworthiopsis sampaiana (Resende) Gildenhuys & Klopper (2016); Haworthia coarctatoides Resende & Viveiros (1948); Haworthia columnaris Baker (1889)  Haworthia pilifera var. columnaris (Baker) Von Poellnitz (1938)  Haworthia obtusa var. columnaris (Baker) Uitewaal (1948); Haworthia confusa Von Poellnitz (1933)  Haworthia minima var. confusa (Von Poellnitz) Von Poellnitz (1938) (incorrect name, ICN Art. 11.4)  Haworthia tenera var. confusa (Von Poellnitz) Uitewaal (1948); Haworthia curta Haworth pro sp. (1819)  Aloe curta (Haworth) Sprengel (1825)  Haworthia tortuosa var. curta (Haworth) Baker (1896)  Haworthiopsis curta (Haworth) Gildenhuys & Klopper (2016); Haworthia cuspidata Haworth (1819)  Aloe cuspidata (Haworth) Roemer & Schultes fil. (1829); Haworthia fergusoniae Von Poellnitz (1930); Haworthia ferox Von Poellnitz (1933); Haworthia ferox var. armata Von Poellnitz (1937); Haworthia fukuyae M. Hayashi (2007); Haworthia gracilidelineata Von Poellnitz (1933)  Haworthia bilineata var. gracilidelineata (Von Poellnitz) Von Poellnitz (1938)  Haworthia cymbiformis fa. gracilidelineata (Von Poellnitz) Pilbeam (1983); Haworthia helmiae Von Poellnitz (1937)  Haworthia unicolor var. helmiae (Von Poellnitz) M. B. Bayer (1976)  Haworthia aristata var. helmiae (Von Poellnitz) Pilbeam (1983) (nom. inval., ICN Art. 41.4)  Haworthia aristata ssp. helmiae (Von Poellnitz) Halda (1997); Haworthia henriquesii Resende (1941); Haworthia icosiphylla Baker (1880); Haworthia janseana Uitewaal (1940); Haworthia krausiana Hort. Haage & Schmidt (s.a.) (nom. inval., ICN Art. 29.1, 38.1a); Haworthia krausii hort. ex J. R. Brown (1957); Haworthia lisbonensis Resende pro sp. (1946)  Haworthiopsis lisbonensis (Resende) Gildenhuys & Klopper (2016); Haworthia longifolia Farden (1939); Haworthia multifaria Haworth (1824) 

773

Aloe multifaria (Haworth) Roemer & Schultes fil. (1829); Haworthia obscura Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); Haworthia ollasonii G. R. Hayes (1971) (nom. inval., ICN Art. 39.1, 40.1); Haworthia pearsonii C. H. Wright (1907)  Haworthia arachnoidea ssp. pearsonii (C. H. Wright) Halda (1997)  Haworthia arachnoidea var. pearsonii (C. H. Wright) Halda (1997); Haworthia pellucens var. delicatula A. Berger (1908)  Haworthia translucens var. delicatula (A. Berger) Von Poellnitz (1938); Haworthia perplexa Von Poellnitz (1938); Haworthia platinosa M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1); Haworthia polyphylla Baker (1880)  Haworthia altilinea var. polyphylla (Baker) Von Poellnitz (1938)  Haworthia mucronata var. polyphylla (Baker) Von Poellnitz (1940); Haworthia pseudogranulata Von Poellnitz (1937); Haworthia radula var. asperior Haworth (1821); Haworthia radula var. laevior Haworth (1821); Haworthia radula var. magniperlata Haworth (1821); Haworthia reinwardtii var. minor hort. ex Baker (1880); Haworthia resendeana Von Poellnitz pro sp. (1938)  Haworthiopsis resendeana (Von Poellnitz) Gildenhuys & Klopper (2016); Haworthia revendettii Uitewaal (1940)  Haworthiopsis revendettii (Uitewaal) Gildenhuys & Klopper (2016); Haworthia rubrobrunnea Von Poellnitz pro sp. (1940)  Haworthiopsis rubrobrunnea (Von Poellnitz) Gildenhuys & Klopper (2016); Haworthia ryderiana Von Poellnitz (1937); Haworthia semiglabrata Haworth (1819)  Aloe semiglabrata (Haworth) Roemer & Schultes fil. (1829); Haworthia semimargaritifera var. multiperlata Haworth (1819)  Aloe semimargaritifera var. multiperlata (Haworth) Roemer & Schultes fil. (1829)  Haworthia margaritifera subvar. multiperlata (Haworth) Von Poellnitz (1938); Haworthia sessiliflora Baker (1896); Haworthia setata var. subinermis Von Poellnitz (1936); Haworthia similis M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1); Haworthia stiemiei Von Poellnitz (1938); Haworthia tisleyi Baker (1880); Haworthia tortella Haworth pro sp. (1819)  Haworthia tortuosa var. tortella (Haworth) Baker (1896)  Haworthiopsis tortella (Haworth) Gildenhuys & Klopper (2016); Haworthia triebneriana var.

774

lanceolata Triebner & Von Poellnitz (1938); Haworthia triebneriana var. nitida Von Poellnitz (1940); Haworthia uitewaaliana Von Poellnitz (1939); Haworthia vitris M. Hayashi (2005) (nom. inval., ICN Art. 40.7); Haworthia walmsleyi Hort. Haage & Schmidt (s.a.) (nom. inval., ICN Art. 29.1, 38.1a); Haworthia yarrowa M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1). H. angustifolia Haworth (Philos. Mag. J. 66: 283, 1825). Type [neo]: RSA, Eastern Cape (Bruyns 1653 [NBG 120 017]). — Distr: RSA (Eastern Cape).  Haworthia chloracantha ssp. angustifolia (Haworth) Halda (1997)  Haworthia chloracantha var. angustifolia (Haworth) Halda (1997). H. angustifolia var. altissima M. B. Bayer (Haworthia Revisited, 26, ill. (p. 27), 1999). Type: RSA, Eastern Cape (Smith 5220 [NBG]). — Distr: RSA (Eastern Cape: Grahamstown Distr.); in Bisho Thornveld and savanna vegetations.  Haworthia altissima (M. B. Bayer) M. Hayashi (2000). [1] Differs from var. angustifolia: Ros proliferous; L slender, erect, to 15  1 cm, finely denticulate along margins and keel, colour tending to greyish-green rather than darkish green. H. angustifolia var. angustifolia — Distr: RSA (Eastern Cape); in savanna. I: Bayer (1982: fig. 1a); Scott (1985: 55). Incl. Aloe stenophylla Roemer & Schultes fil. (1829); incl. Haworthia albanensis Schönland (1912)  Haworthia angustifolia var. albanensis (Schönland) Von Poellnitz (1937); incl. Haworthia angustifolia var. grandis G. G. Smith (1943)  Haworthia angustifolia fa. grandis (G. G. Smith) Pilbeam (1983); incl. Haworthia grandis M. Hayashi (2000) (nom. inval., ICN Art. 39.1, 40.1); incl. Haworthia anna M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1); incl. Haworthia cocksia M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1); incl. Haworthia viridula M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1). [1] Ros to 4 cm ∅, proliferous, stemless; L 10–40, slender, erect, to 10  1 cm, lanceolate-acuminate, somewhat flaccid, brownish to

M. B. Bayer and E. Van Jaarsveld

dark green, margins and keel finely denticulate; Inf to 20 cm, lax; Fl 8–10, white to dull pinkishwhite. H. angustifolia var. baylissii (C. L. Scott) M. B. Bayer (Haworthia Revisited, 27, ill., 1999). Type: RSA (Scott 796 [PRE, NBG]). — Distr: RSA (Eastern Cape: Oudekraal); in Albany Thicket vegetation. I: Bayer (1982: Fig. 1b, as fa.); Scott (1985: 102, as H. baylissii).  Haworthia baylissii C. L. Scott (1968)  Haworthia angustifolia fa. baylissii (C. L. Scott) M. B. Bayer (1982)  Haworthia chloracantha ssp. baylissii (C. L. Scott) Halda (1997). [1] Differs from var. angustifolia: L deltoid, recurving. H. angustifolia var. paucifolia G. G. Smith (J. South Afr. Bot. 14: 48, ill. (p. 58), 1948). Type: RSA, Eastern Cape (Smith 6819 [NBG]). — Distr: RSA (Eastern Cape: Frazer’s Camp to Kaffirdrift); in savanna.  Haworthia angustifolia fa. paucifolia (G. G. Smith) Pilbeam (1983)  Haworthia paucifolia (G. G. Smith) M. Hayashi (2010). [1] Differs from var. angustifolia: L only 3–4, to 3 cm. H. arachnoidea (Linné) Duval (Pl. Succ. Horto Alencon., 7, 1809). Type: [lecto — icono]: Commelin, Praeludia Bot., t. 27, 1703. — Lit: Bayer (2003b); Bayer (2004b); Bayer (2010g). Distr: RSA (Northern Cape, Western Cape, Eastern Cape); mainly in Succulent Karoo vegetation.  Aloe pumila var. arachnoidea Linné (1753)  Aloe arachnoidea (Linné) Burman fil. (1768)  Catevala arachnoidea (Linné) Medikus (1786)  Apicra arachnoidea (Linné) Willdenow (1811) (incorrect name, ICN Art. 11.4); incl. Aloe arachnoides Thunberg (1785); incl. Aloe arachnoidea var. communis Aiton (1789) (nom. inval., ICN Art. 26.1). H. arachnoidea var. arachnoidea — Distr: RSA (Western Cape: Little Karoo). I: Bayer (1982: Fig. 2). – Fig. 1. Incl. Haworthia arachnoidea var. minor Haworth (1819); incl. Haworthia joubertii M. Hayashi

Haworthia ASPHODELACEAE

775

Bayer (1976); incl. Haworthia araneoides M. Hayashi (2010) (nom. inval., ICN Art. 39.1, 40.1). [1] Differs from var. arachnoidea: Ros smaller, to 6 cm ∅; L softer in texture.

Fig. 1 Haworthia arachnoidea var. arachnoidea. (Copyright: E. J. Van Jaarsveld)

(2005) (nom. inval., ICN Art. 40.7)  Haworthia arachnoidea var. joubertii (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7); incl. Haworthia laxa M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia arachnoidea var. laxa (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7); incl. Haworthia limbata M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia arachnoidea var. limbata (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7); incl. Haworthia gilva Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia isomorpha Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] R slender, succulent; Ros stemless, variable in size from 6 to exceptionally 12 cm ∅, solitary or forming small clusters; L 25–80, dense, incurving, uniformly light to dark green, not translucent and only occasionally faintly reticulate, flattened and often drying grey-white to brownish at the tips forming a protective cover, triangular-lanceolate to ovate-lanceolate, 2–7  1–1.5 cm, keeled, margin and keels with translucent bristly Sp to 12 mm, tip acuminate-aristate; Inf to 30 cm; Fl 20–30, white. H. arachnoidea var. aranea (A. Berger) M. B. Bayer (Haworthia Revisited, 30, ill. (p. 31), 1999). Type: [lecto — icono]: A. Berger in Engler, A. & Prantl, K. (eds.), Pflanzenr. 4(38 = Heft 33): 114, Fig. 39A–E, 1908. — Distr: RSA (Western Cape: Little Karoo); dry Mountain Fynbos vegetation. I: Bayer (1982: Fig. 3, as H. aranea).  Haworthia bolusii var. aranea A. Berger (1908)  Haworthia aranea (A. Berger) M. B.

H. arachnoidea var. namaquensis M. B. Bayer (Haworthia Revisited, 31, ills., 1999). Type: RSA, Northern Cape (Bayer 1674 [NBG]). — Distr: RSA (Northern Cape, Western Cape); Succulent Karoo vegetation.  Haworthia namaquensis (M. B. Bayer) Breuer (2010). [1] Differs from var. arachnoidea: Ros to 6 cm ∅; L paler green. H. arachnoidea var. nigricans (Haworth) M. B. Bayer (Haworthia Revisited, 32, ill., 1999). Type [neo]: RSA (Bayer 2419 [NBG]). — Lit: Bayer (2002b). Distr: RSA (Western Cape: Little Karoo); Succulent Karoo vegetation. I: Scott (1985: 100, as H. helmiae).  Haworthia setata var. nigricans Haworth (1821)  Aloe setata var. nigricans (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia venteri Von Poellnitz (1939)  Haworthia unicolor var. venteri (Von Poellnitz) M. B. Bayer (1976); incl. Haworthia scottii Breuer (2003); incl. Haworthia nigrata M. Hayashi (2006); incl. Haworthia apta Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia formosa Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia kuromisa Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia regens Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Differs from var. arachnoidea: L with dark purplish coloration. H. arachnoidea var. scabrispina M. B. Bayer (Haworthia Revisited, 34, ills., 1999). Type: RSA, Western Cape (Bayer 2105 [NBG]). — Distr: RSA (Western Cape); Nama Karoo vegetation.  Haworthia scabrispina (M. B. Bayer) Breuer (2010); incl. Haworthia matjiesta Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Differs from var. arachnoidea: Ros roundish, raised above ground-level; L with firm rigid brownish spines.

776

H. arachnoidea var. setata (Haworth) M. B. Bayer (Haworthia Revisited, 34, ills. (p. 35), 1999). Type: RSA, Western Cape (Makrill s.n. [[lecto — icono]: K [drawing]]). — Distr: RSA (Western Cape, W Eastern Cape: Little Karoo). I: Scott (1985: 69, as H. setata).  Haworthia setata Haworth (1819)  Aloe setata (Haworth) Roemer & Schultes fil. (1829)  Haworthia arachnoidea ssp. setata (Haworth) Halda (1997); incl. Haworthia setata var. major Haworth (1821)  Aloe setata var. major (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia setata var. media Haworth (1821)  Aloe setata var. media (Haworth) Roemer & Schultes fil. (1829); incl. Aloe setosa Roemer & Schultes fil. (1829); incl. Haworthia gigas Von Poellnitz (1933)  Haworthia setata var. gigas (Von Poellnitz) Von Poellnitz (1938)  Haworthia arachnoidea var. gigas (Von Poellnitz) M. Hayashi (2000); incl. Haworthia minima var. major Von Poellnitz (1938) (incorrect name, ICN Art. 11.4)  Haworthia tenera var. major (Von Poellnitz) Uitewaal (1948); incl. Haworthia pectinis M. Hayashi (2003)  Haworthia tretyrensis var. pectinis (M. Hayashi) Breuer (2016); incl. Haworthia tretyrensis Breuer (2003); incl. Haworthia angiras M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia arachnoidea var. angiras (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7); incl. Haworthia cangoensis M. Hayashi (2005) (nom. inval., ICN Art. 40.7); incl. Haworthia kogmanensis M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia cangoensis var. kogmanensis (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 35.1, 40.7); incl. Haworthia royalis M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia cangoensis var. royalis (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 35.1, 40.7); incl. Haworthia candida M. Hayashi (2006)  Haworthia aranea var. candida (M. Hayashi) Breuer (2016). [1] Differs from var. arachnoidea: Ros variable; L white-spined. H. aristata Haworth (Suppl. Pl. Succ., 51, 1819). Type: K [lecto — icono]. — Distr: RSA (Eastern Cape); Albany Thicket vegetation. – Fig. 2.

M. B. Bayer and E. Van Jaarsveld

Fig. 2 Haworthia aristata. (Bayer 7697b: N of Swartkop, Kaboega.) (Copyright: M. B. Bayer)

 Aloe aristata (Haworth) Roemer & Schultes fil. (1829) (nom. illeg., ICN Art. 53.1); incl. Haworthia denticulata Haworth (1821)  Aloe denticulata (Haworth) Roemer & Schultes fil. (1829)  Aloe altilinea var. denticulata (Haworth) Salm-Dyck (1842)  Haworthia altilinea var. denticulata (Haworth) Von Poellnitz (1938); incl. Haworthia lapis Breuer & M. Hayashi (2004); incl. Haworthia dura M. Hayashi (2005) (nom. inval., ICN Art. 40.7); incl. Haworthia rava M. Hayashi (2005) (nom. inval., ICN Art. 40.7)  Haworthia lapis var. rava (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7). [1] Ros stemless, proliferating slowly, to 6 cm ∅; L slender, erect, incurved, dark green, hardly translucent and faintly reticulated, margins and keel entire or minutely spined; Inf to 15 cm, lax; Fl 10–15, white. Growing in close vicinity with H. cooperi at times, with occasional occurrence of intermediate plants (Bayer 2010b). H. attenuata (Haworth) Haworth (Synops. Pl. Succ., 92, 1812). Type [neo]: RSA, Eastern Cape (Perry 660 [NBG 144 672]). — Distr: RSA (Eastern Cape).  Aloe attenuata Haworth (1804)  Apicra attenuata (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthia pumila ssp.

Haworthia ASPHODELACEAE

attenuata (Haworth) Halda (1997)  Haworthiopsis attenuata (Haworth) G. D. Rowley (2015). H. attenuata var. attenuata — Distr: RSA (Eastern Cape); Albany Thicket vegetation. I: Bayer (1982: Fig. 46); Gildenhuys (2017: 15–16, as Haworthiopsis). Incl. Haworthia clariperla Haworth (1826)  Aloe clariperla (Haworth) Roemer & Schultes fil. (1829)  Aloe attenuata var. clariperla (Haworth) Salm-Dyck (1836)  Haworthia attenuata var. clariperla (Haworth) Baker (1880)  Haworthia attenuata fa. clariperla (Haworth) M. B. Bayer (1976); incl. Haworthia fasciata var. caespitosa A. Berger (1908)  Haworthia attenuata var. caespitosa (A. Berger) R. S. Farden (1939)  Haworthia attenuata fa. caespitosa (A. Berger) Pilbeam (1983) (nom. inval., ICN Art. 41.4); incl. Haworthia britteniana Von Poellnitz (1937)  Haworthia attenuata var. britteniana (Von Poellnitz) Von Poellnitz (1937)  Haworthia attenuata fa. britteniana (Von Poellnitz) M. B. Bayer (1982); incl. Haworthia attenuata var. deltoidea R. S. Farden (1939); incl. Haworthia attenuata var. linearis R. S. Farden (1939); incl. Haworthia attenuata var. minissima R. S. Farden (1939); incl. Haworthia attenuata var. odonoghueana R. S. Farden (1939); incl. Haworthia attenuata var. inusitata R. S. Farden (1939) (nom. inval., ICN Art. 39.1); incl. Haworthia attenuata var. uitewaaliana R. S. Farden (1939) (nom. inval., ICN Art. 39.1). [2] Ros stemless, proliferous, to 13 cm tall and 10 cm ∅; L to 13  1.5 cm, attenuate, spreading, lanceolate-deltoid, surfaces scabrid with distinct raised non-confluent tubercles; Inf sparsely branched, 24–28 cm, lax; Fl tube obcapitate, Tep revolute. H. attenuata var. glabrata (Salm-Dyck) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 37, 2012). Type: [neo — icono]: Salm-Dyck, Monogr. Gen. Aloes & Mesembr. 3: Aloe t. 7 [sect. 6: 13], 1840. — Distr: RSA (Eastern Cape: former Transkei); Eastern Valley Bushveld vegetation. I: Bayer (1982: Fig. 50, as H. glabrata); Gildenhuys (2017: 17, as Haworthiopsis).  Aloe glabrata Salm-Dyck (1834)  Haworthia glabrata (Salm-Dyck) Baker (1880) 

777

Haworthiopsis attenuata var. glabrata (Salm-Dyck) G. D. Rowley (2015). [2] Differs from var. attenuata: L scabrid and more turgid. H. attenuata var. radula (Jacquin) M. B. Bayer (Haworthia Revisited, 167, ill., 1999). Type: [lecto — icono]: Jacquin, Pl. Hort. Schoenbr. 4: t. 422, 1804. — Distr: RSA (Eastern Cape: Baviaanskloof); Gamtoos Thicket vegetation. I: Bayer (1982: Fig. 56, as H. radula); Gildenhuys (2017: 18, as Haworthiopsis).  Aloe radula Jacquin (1804)  Apicra radula (Jacquin) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthia radula (Jacquin) Haworth (1812)  Haworthia pumila ssp. radula (Jacquin) Halda (1997)  Haworthiopsis attenuata var. radula (Jacquin) G. D. Rowley (2015); incl. Haworthia radula var. pluriperlata Haworth (1821). [2] Differs from var. attenuata: L with many minute crowded white tubercles. H. bayeri J. D. Venter & S. A. Hammer (Cact. Succ. J. (US) 69(2): 75–76, ill., 1997). Type: RSA, Western Cape (Stayner s.n. in Karoo Garden 164/69 [NBG]). — Distr: RSA (Western Cape: Little Karoo); Succulent Karoo vegetation. Incl. Haworthia uniondalensis hort. (s.a.) (nom. inval., ICN Art. 29.1); incl. Haworthia hayashii M. Hayashi (2002)  Haworthia bayeri var. hayashii (M. Hayashi) Breuer (2016); incl. Haworthia bayeri var. scabrifolia Breuer (2002) (nom. inval., ICN Art. 39.1, 40.1); incl. Haworthia indigoa M. Hayashi (2005); incl. Haworthia jadea M. Hayashi (2005)  Haworthia bayeri var. jadea (M. Hayashi) Breuer (2016); incl. Haworthia laeta M. Hayashi (2005); incl. Haworthia truterorum Breuer & Marx (2011)  Haworthia indigoa var. truterorum (Breuer & Marx) Breuer (2016). [1] Ros stemless, to 8 cm ∅; L 15–20, retuse, dark brownish-green to blackish-green, slightly scabrid, keels and margins with minute Sp or smooth, tip rounded and not pointed, the end-area opaque, cloudy-transparent, with sparse reticulate patterning or longitudinal lines; Inf to 30 cm; Fl 15–25. Here belongs H. willowmorensis in the sense of Scott (1985). The recently described H. truterorum

778

M. B. Bayer and E. Van Jaarsveld

is provisionally synonymized here and appears to fall between H. bayeri and H. emelyae. H. blackburniae W. F. Barker (J. South Afr. Bot. 3: 93, 1937). Type: RSA, Western Cape (Reynolds 1842 [NBG, BOL, PRE]). — Distr: RSA (Western Cape); Succulent Karoo and Fynbos vegetations. Incl. Haworthia blackburniana hort. (s.a.) (nom. inval., ICN Art. 61.1). H. blackburniae var. blackburniae — Distr: RSA (Western Cape: mountains of the C Little Karoo). I: Bayer (1982: Fig. 6). R fusiform; Ros stemless, basally 1–1.5 cm ∅, clumping; L 10–15, erect, long, slender, to 40 cm  3–5 mm, bright green to brownish-green or dark greyish-green, upper face channelled, margins glabrous or finely spined; Inf to 30 cm; Fl 15–20, white with green veins. H. blackburniae var. derustensis M. B. Bayer (Haworthia Revisited, 41, ill., 1999). Type: RSA, Western Cape (Vlok s.n. in Venter 93/24 [NBG]). — Distr: RSA (Western Cape: Oudtshoorn Distr.).  Haworthia derustensis (M. B. Bayer) M. Hayashi (2000)  Haworthia graminifolia var. derustensis (M. B. Bayer) Breuer (2016). [1] Differs from var. blackburniae: Ros robust, to 1.8 cm ∅; L very long, to 45 cm, to 3 mm broad, brownish-green at the base, green above. H. blackburniae var. graminifolia (G. G. Smith) M. B. Bayer (Haworthia Revisited, 42, ills., 1999). Type: RSA, Western Cape (Smith 5222 [NBG]). — Distr: RSA (Western Cape: Swartberg Mts.). I: Bayer (1982: Fig. 16, as H. graminifolia).  Haworthia graminifolia G. G. Smith (1942)  Haworthia blackburniae ssp. graminifolia (G. G. Smith) Halda (1997). [1] Differs from var. blackburniae: L 2 mm. H. bolusii var. bolusii — Distr: RSA (interior of the Eastern Cape). I: Scott (1985: 72). Incl. Haworthia odetteae Breuer (2003); incl. Haworthia odyssei M. Hayashi (2005) (nom.

Haworthia ASPHODELACEAE

inval., ICN Art. 40.7)  Haworthia odetteae var. odyssei (M. Hayashi) Breuer (2016) (nom. inval., ICN Art. 40.7); incl. Haworthia capillaris M. Hayashi (2006); incl. Haworthia sapphaia M. Hayashi (2007). [1] Ros 4–8 cm ∅, slowly proliferating; L oblong-lanceolate, incurved, translucent bluishgreen, margins and keel with Sp >2 mm; Inf robust, to 30 cm; Fl broad and flat across the base of the tube. H. bolusii var. pringlei (C. L. Scott) M. B. Bayer (Haworthiad 16(2): 62, 2002). Type: RSA, Eastern Cape (Pringle s.n. in Scott 8970 [PRE]). — Distr: RSA (Eastern Cape). I: Scott (1994b: as H. pringlei).  Haworthia pringlei C. L. Scott (1994)  Haworthia decipiens var. pringlei (C. L. Scott) M. B. Bayer (1999); incl. Haworthia aquamarina M. Hayashi (2003); incl. Haworthia hastata M. Hayashi (2005)  Haworthia pringlei var. hastata (M. Hayashi) Breuer (2016); incl. Haworthia hisui M. Hayashi (2005)  Haworthia cooperi var. hisui (M. Hayashi) Breuer (2016); incl. Haworthia lazulis M. Hayashi (2005) (nom. inval., ICN Art. 40.7). [1] Differs from var. bolusii: L  lanceolate, incurved and erect to suberect, bright green, margins and keel with white Sp. H. bruynsii M. B. Bayer (J. South Afr. Bot. 47: 789, 1981). Type: RSA, Eastern Cape (Rossouw 456 [NBG]). — Distr: RSA (Eastern Cape: Great Karoo); Nama Karoo vegetation. I: Bayer (1982: Fig. 47); Gildenhuys (2017: 63, as Haworthiopsis).  Haworthia retusa var. bruynsii (M. B. Bayer) Halda (1997)  Haworthiopsis bruynsii (M. B. Bayer) G. D. Rowley (2013). [2] Ros stemless, solitary, to 6 cm ∅; L 5–11, 2–3 cm, brownish-green, with flat retuse end-area, opaque and slightly scabrid with small raised tubercles; Inf simple, slender; Fl distant, slender, tube obcapitate; Tep fused, tips revolute. H. chloracantha Haworth (Revis. Pl. Succ., 57, 1821). Type [lecto]: RSA, Western Cape (Bowie s.n. [[lecto — icono]: K [drawing]]). — Distr: RSA (Western Cape); Fynbos vegetation.

779

 Aloe chloracantha (Haworth) Roemer & Schultes fil. (1829). H. chloracantha var. chloracantha — Distr: RSA (Western Cape). I: Bayer (1982: Fig. 8a). [1] Ros 2.5–4 cm ∅, proliferous; L 18–25, 0.5–1.5 cm long, erectly spreading, light green, faintly reticulate, firm to slightly scabrid, triangular in cross-section, margins and keel with translucent Sp to 0.3 mm; Inf lax, to 25 cm; Fl small. H. chloracantha var. denticulifera (Von Poellnitz) M. B. Bayer (Haworthia Handb., 112, 1976). Type: B [lecto: unpubl. ill.]. — Distr: RSA (Western Cape: Mosselbay). I: Bayer (1982: Fig. 8b).  Haworthia angustifolia var. denticulifera Von Poellnitz (1937)  Haworthia denticulifera (Von Poellnitz) M. Hayashi (2000); incl. Haworthia angustifolia var. liliputana Uitewaal (1953). [1] Differs from var. chloracantha: L 2.5– 3.5 cm long, dark green. H. chloracantha var. subglauca Von Poellnitz (Kakteenkunde 9: 135, 1937). Type [neo]: RSA, Western Cape (Hurling & Neil s.n. [BOL]). — Distr: RSA (Western Cape: Mosselbay). I: Bayer (1982: Fig. 8c).  Haworthia subglauca (Von Poellnitz) M. Hayashi (2000). [1] Differs from var. chloracantha: L bluishgreen, Sp larger and sparser. H. coarctata Haworth (Philos. Mag. J. 64: 301, 1824). Type [neo]: RSA, Eastern Cape (Smith 7092 [NBG 68473]). — Distr: RSA (Eastern Cape); Fynbos and thickets.  Aloe coarctata (Haworth) Roemer & Schultes fil. (1829)  Haworthia reinwardtii ssp. coarctata (Haworth) Halda (1997)  Haworthia reinwardtii var. coarctata (Haworth) Halda (1997)  Haworthiopsis coarctata (Haworth) G. D. Rowley (2013)  Haworthiopsis reinwardtii var. coarctata (Haworth) Breuer (2016). H. coarctata fa. greenii (Baker) M. B. Bayer (Haworthia Revisited, 172, ill., 1999). Type: RSA, Eastern Cape (Cooper s.n. [K]). — Distr: RSA

780

(Eastern Cape: Grahamstown). I: Bayer (1982: Fig. 48b, as var.).  Haworthia greenii Baker (1880)  Haworthia coarctata var. greenii (Baker) M. B. Bayer (1973)  Haworthia reinwardtii var. greenii (Baker) Halda (1997)  Haworthiopsis reinwardtii var. greenii (Baker) Breuer (2016); incl. Haworthia peacockii Baker (1880); incl. Haworthia greenii fa. minor Resende (1943); incl. Haworthia greenii fa. bakeri Resende (1943) (nom. illeg., Art. 52.1, 26.1). [2] Differs from var. coarctata: L without tubercles. H. coarctata var. adelaidensis (Von Poellnitz) M. B. Bayer (Haworthia Revisited, 172, ills., 1999). Type: RSA, Eastern Cape (Armstrong s.n. [B [lecto: unpubl. ill.]]). — Distr: RSA (Eastern Cape: Grahamstown). I: Bayer (1982: Fig. 48d, as ssp.); Gildenhuys (2017: 21, as Haworthiopsis).  Haworthia reinwardtii var. adelaidensis Von Poellnitz (1940)  Haworthia coarctata ssp. adelaidensis (Von Poellnitz) M. B. Bayer (1973)  Haworthia adelaidensis (Von Poellnitz) Breuer (2010)  Haworthiopsis coarctata var. adelaidensis (Von Poellnitz) G. D. Rowley (2013); incl. Haworthia reinwardtii var. riebeekensis G. G. Smith (1944); incl. Haworthia reinwardtii var. bellula G. G. Smith (1945)  Haworthia coarctata fa. bellula (G. G. Smith) Pilbeam (1983). [2] Differs from var. coarctata: Ros smaller, stems to 15 cm, 3 cm ∅; L 3.2  1 cm. H. coarctata var. coarctata — Distr: RSA (Eastern Cape). I: Bayer (1982: Fig. 48a); Gildenhuys (2017: 19–20, as Haworthiopsis). Incl. Haworthia chalwinii Marloth & A. Berger (1906)  Haworthia reinwardtii var. chalwinii (Marloth & A. Berger) Resende (1943)  Haworthia coarctata fa. chalwinii (Marloth & A. Berger) Pilbeam (1983); incl. Haworthia fallax Von Poellnitz (1933)  Haworthia reinwardtii var. fallax (Von Poellnitz) Von Poellnitz (1937); incl. Haworthia reinwardtii var. conspicua Von Poellnitz (1937)  Haworthia coarctata fa. conspicua (Von Poellnitz) Pilbeam (1983); incl. Haworthia reinwardtii var. pseudocoarctata Von Poellnitz (1940)  Haworthia coarctata fa. pseudocoarctata

M. B. Bayer and E. Van Jaarsveld

(Von Poellnitz) Resende (1943)  Haworthia greenii fa. pseudocoarctata (Von Poellnitz) Resende & Pinto-Lopes (1946); incl. Haworthia coarctata fa. major Resende (1943); incl. Haworthia coarctata var. kraussii Resende (1943); incl. Haworthia fulva G. G. Smith (1943); incl. Haworthia greenii var. silvicola G. G. Smith (1943); incl. Haworthia reinwardtii var. committeesensis G. G. Smith (1943); incl. Haworthia coarctata var. haworthii Resende (1943) (nom. illeg., ICN Art. 52.1, 26.1); incl. Haworthia reinwardtii var. huntsdriftensis G. G. Smith (1944); incl. Haworthia musculina G. G. Smith (1948). [2] Ros to 12 cm ∅, caulescent, proliferating; L numerous, to 7  2 cm, ratio stem diameter : leaf width = 1:1.7, erect-spreading or incurved, scabrid, both faces brownish-green, usually with rounded tubercles; Inf simple or occasionally compound, to 60 cm; Fl tube obcapitate; Tep revolute. Not always easily separated from H. reinwardtii and its forms. The ratio between stem diameter and leaf width is a good help. H. coarctata var. tenuis (G. G. Smith) M. B. Bayer (Nation. Cact. Succ. J. 28: 80, 1973). Type: RSA, Eastern Cape (Smith 3420 [NBG]). — Distr: RSA (Eastern Cape: Alexandria). I: Bayer (1982: Fig. 48c); Gildenhuys (2017: 22, as Haworthiopsis).  Haworthia reinwardtii var. tenuis G. G. Smith (1948)  Haworthia tenuis (G. G. Smith) Breuer (2010)  Haworthiopsis coarctata var. tenuis (G. G. Smith) G. D. Rowley (2013)  Haworthiopsis reinwardtii var. tenuis (G. G. Smith) Breuer (2016). [2] Differs from var. coarctata: Ros with elongated narrow stems to 40 cm, 2.5 cm ∅; L 3.5  0.8 cm. H. cooperi Baker (Refug. Bot. 4: t. 233 + text, 1871). Type: RSA, Eastern Cape (Cooper s.n. [K]). — Distr: RSA (Eastern Cape). H. cooperi var. cooperi — Distr: RSA (Eastern Cape); mainly Fynbos, grassland and Albany Thicket vegetations. I: Bayer (1982: Fig. 10a); Bayer (2010c).

Haworthia ASPHODELACEAE

Incl. Haworthia vittata Baker (1871); incl. Haworthia subularis M. Hayashi (2003); incl. Haworthia pallens Breuer & M. Hayashi (2004)  Haworthia caerulea var. pallens (Breuer & M. Hayashi) Breuer (2016); incl. Haworthia calva M. Hayashi (2005)  Haworthia venetia var. calva (M. Hayashi) Breuer (2016); incl. Haworthia cooperi var. cymbiformoides M. B. Bayer (2008) (nom. inval., ICN Art. 36.1b); incl. Haworthia elegans Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia foeda Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia turcosa Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia yocans Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Ros to 12 cm ∅, often proliferous, stemless; L 20–40, fleshy, swollen, erect, oblonglanceolate, quickly tapering, acuminate or truncate, bluish-green, slightly translucent, with veins usually reddening and leaves becoming purplish in exposed situations, marginal Sp 10 cm ∅; L more suberect to spreading. H. retusa (Linné) Duval (Pl. Succ. Horto Alencon., 7, 1809). Type: [lecto — icono]: Commelin, Horti Med. Amstelod. 2: 11, t. 6, 1701. — Distr: RSA (Western Cape); Fynbos vegetation.  Aloe retusa Linné (1753)  Catevala retusa (Linné) Medikus (1786)  Apicra retusa (Linné) Willdenow (1811) (incorrect name, ICN Art. 11.4). H. retusa var. longibracteata (G. G. Smith) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 36, 2012). Type: RSA, Western Cape (Dekenah 18 in Smith 5378 [NBG, PRE]). — Distr: RSA (Western Cape). I: Scott (1985: 127, as H. longibracteata).  Haworthia longibracteata G. G. Smith (1945)  Haworthia retusa fa. longibracteata (G. G. Smith) Pilbeam (1983)  Haworthia turgida var. longibracteata (G. G. Smith) M. B. Bayer (1999). [1] Differs from var. retusa: L erect to suberect, ovate-lanceolate. H. retusa var. nigra (M. B. Bayer) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 36, 2012). Type: RSA, Western Cape (Smith 5753 [NBG]). — Lit: Bayer (2004a); Bayer (2004c); Bayer (2005); all as H. mutica var. Distr: RSA (Western Cape: Heidelberg area).  Haworthia mutica var. nigra M. B. Bayer (1999)  Haworthia silviae var. nigra (M. B. Bayer) M. Hayashi (2000); incl. Haworthia chromutica M. Hayashi (2000) (nom. inval., ICN Art. 39.1, 40.1); incl. Haworthia quimutica Breuer (2011) (nom. inval., ICN Art. 38.1a). [1] Differs from var. retusa: Ros more proliferous; L greener and more translucent.

807

H. retusa var. retusa — Distr: RSA (Western Cape: Riversdale). I: Bayer (1982: Fig. 33a). Incl. Haworthia fouchei Von Poellnitz (1940)  Haworthia retusa fa. fouchei (Von Poellnitz) Pilbeam (1983)  Haworthia retusa var. fouchei (Von Poellnitz) Breuer (2016); incl. Haworthia retusa var. densiflora G. G. Smith (1946); incl. Haworthia retusa var. multilineata G. G. Smith (1946)  Haworthia retusa fa. multilineata (G. G. Smith) Pilbeam (1983)  Haworthia multilineata (G. G. Smith) C. L. Scott (1985); incl. Haworthia retusa var. solitaria G. G. Smith (1946)  Haworthia solitaria (G. G. Smith) C. L. Scott (1973); incl. Haworthia geraldii C. L. Scott (1965)  Haworthia retusa fa. geraldii (C. L. Scott) Pilbeam (1983) (nom. inval., ICN Art. 41.5); incl. Haworthia retusa var. quimutica Hayashi (2001); incl. Haworthia subretusa Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Ros stemless, rarely slowly proliferating, to 12 cm ∅; L 10–15, turgid, rigid, with pronouncedly retuse end-areas, 8  2 cm, brownish or green and rarely with purplish hue, variously lined and windowed, surface and usually also margins and keel without spines or tubercles, tips pointed; Inf robust, to 30 cm; Fl 20–30, closely spaced, white with greenish-brown veins. Possible hybridization with H. mirabilis is mentioned by Bayer (2012d). H. retusa var. suberecta (Von Poellnitz) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 36, 2012). Type [neo]: RSA, Western Cape (Bayer s.n. in Karoo Garden 631/69 [NBG]). — Distr: RSA (Western Cape). I: Scott (1985: 126, as H. dekenahii).  Haworthia turgida var. suberecta Von Poellnitz (1938)  Haworthia turgida fa. suberecta (Von Poellnitz) Pilbeam (1983)  Haworthia suberecta (Poellnitz) Breuer (2010); incl. Haworthia turgida var. subtuberculata Von Poellnitz (1938); incl. Haworthia turgida var. pallidifolia G. G. Smith (1946)  Haworthia turgida fa. pallidifolia (G. G. Smith) Pilbeam (1983)  Haworthia pallidifolia (G. G. Smith) M. Hayashi (2010)  Haworthia suberecta var.

808

pallidifolia (G. G. Smith) Breuer (2016); incl. Haworthia pseuda Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia reflexa Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia rodinii Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Differs from var. retusa: L strongly mottled, tips slightly truncate and rounded. H. retusa var. turgida (Haworth) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 36, 2012). Type [neo]: RSA, Western Cape (Bayer 2420 [NBG 132378]). — Distr: RSA (Western Cape). I: Bayer (1982: Fig. 40, as H. turgida).  Haworthia turgida Haworth (1819)  Aloe turgida (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia laetevirens Haworth (1819)  Aloe laetevirens (Haworth) Link (1822); incl. Haworthia caespitosa Von Poellnitz (1937)  Haworthia turgida fa. caespitosa (Von Poellnitz) Pilbeam (1983); incl. Haworthia caespitosa fa. subplana Von Poellnitz (1938); incl. Haworthia caespitosa fa. subproliferans Von Poellnitz (1938). [1] Differs from var. turgida: Ros partially stemless, proliferous, 5–10 cm ∅; L 20–40, ovate-lanceolate, 4  1.2 cm, turgid, often as thick as broad, recurved or slightly retuse, margins and keel lightly spined. H. rossouwii Von Poellnitz (Kakteenkunde 1938: 75, ill., 1938). Type: RSA, Western Cape (Rossouw s.n. in Triebner 1059 [B [lecto, ill. later publ. l.c. p. 75]]). — Lit: Bayer (2001). Distr: RSA (Western Cape: Bredasdorp to Heidelberg); mainly in Fynbos vegetation. H. rossouwii var. calcarea (M. B. Bayer) M. B. Bayer (Aloe 38(1–2): 36, 2001). Type: RSA, Western Cape (Burgers 1648 [NBG]). — Distr: RSA (Western Cape: Bredasdorp Distr.). I: Bayer (2001).  Haworthia mirabilis var. calcarea M. B. Bayer (1999)  Haworthia calcarea (M. B. Bayer) M. Hayashi (2000)  Haworthia mundula var. calcarea (M. B. Bayer) Breuer (2016). [1] Differs from var. rossouwii: Ros with fewer leaves; L short, erect, with short retuse end-area.

M. B. Bayer and E. Van Jaarsveld

H. rossouwii var. elizeae (Breuer) M. B. Bayer (Haworthia Update Vol. 2, 154, 2006). Nom. inval., ICN Art. 41.5. Type: RSA, Western Cape (de Vries 563 in Breuer 6936 [Res. Inst. Evol. Biol. Tokyo]). — Distr: RSA (Western Cape: W of Swellendam).  Haworthia elizeae Breuer (2003) (nom. inval., ICN Art. 41.5); incl. Haworthia esterhuizenii Breuer (2002) (nom. inval., ICN Art. 41.5). [1] Differs from var. rossouwii: Ros smaller, more globose. Unfortunately, there is no valid name available for this taxon. H. rossouwii var. minor (M. B. Bayer) M. B. Bayer (Haworthia Update Vol. 7, Part 4, 36–37, 2012). Type: RSA, Western Cape (Bayer s.n. in Karoo Garden 36/70 [NBG]). — Distr: RSA (Western Cape: Bredasdorp Distr.).  Haworthia heidelbergensis var. minor M. B. Bayer (1999); incl. Haworthia rooivleiensis Breuer (2010). [1] Differs from var. rossouwii: Ros much smaller, to 3 cm ∅; L light yellowish-green, well-spined. H. rossouwii var. petrophila (M. B. Bayer) M. B. Bayer (Aloe 38(1–2): 36, 2001). Type: RSA, Western Cape (Burgers 2158 [NBG]). — Distr: RSA (Western Cape: Bromberg, Stormsvlei).  Haworthia variegata var. petrophila M. B. Bayer (1999)  Haworthia petrophila (M. B. Bayer) M. Hayashi (2000). [1] Differs from var. rossouwii: Ros very proliferous; L slender, shorter and incurved, strongly spined. H. rossouwii var. rossouwii — Distr: RSA (Western Cape: SW of Heidelberg to Bredasdorp); mainly in Fynbos vegetation. I: Bayer (1982: Fig. 36, as H. serrata). Incl. Haworthia serrata M. B. Bayer (1973)  Haworthia chloracantha var. serrata (M. B. Bayer) Halda (1997)  Haworthia rossouwii var. serrata (M. B. Bayer) Breuer (2016). [1] Ros stemless, rarely proliferating, to 7 cm ∅; L 20–30, narrow, 6  1 cm, acuminate, bright

Haworthia ASPHODELACEAE

yellowish-green with translucent lines above, margins and keel spiny; Inf robust, to 40 cm; Fl 20–30, white with green venation. H. scabra Haworth (Suppl. Pl. Succ., 58, 1819). Type: K [lecto — icono]: publ. Cact. Succ. J. (US) 52: 274, 1980. — Distr: RSA (Western Cape, Eastern Cape); mainly in Fynbos and Succulent Karoo vegetation.  Aloe scabra (Haworth) Roemer & Schultes fil. (1829)  Haworthiopsis scabra (Haworth) G. D. Rowley (2013). H. scabra var. lateganiae (Von Poellnitz) M. B. Bayer (Haworthia Revisited, 195, ills., 1999). Type: B [lecto — icono]: photograph publ. Desert Pl. Life 9: 103, 1937. — Distr: RSA (Western Cape: Little Karoo, E of Oudtshoorn). I: Bayer (1982: Fig. 60b, as H. starkiana var.); Gildenhuys (2017: 56, as Haworthiopsis).  Haworthia lateganiae Von Poellnitz (1937)  Haworthia starkiana var. lateganiae (Von Poellnitz) M. B. Bayer (1976)  Haworthiopsis scabra var. lateganiae (Von Poellnitz) G. D. Rowley (2013). [2] Differs from var. scabra: L long and slender, smooth and shiny, dark green. H. scabra var. morrisiae (Von Poellnitz) M. B. Bayer (Haworthia Handb., 137, 1976). Type: B [lecto — icono]: ill., publ. Kakteenkunde 1937: 132. — Distr: RSA (Western Cape: Little Karoo, Schoemanspoort). I: Bayer (1982: Fig. 58b); Gildenhuys (2017: 57, as Haworthiopsis).  Haworthia morrisiae Von Poellnitz (1937)  Haworthiopsis scabra var. morrisiae (Von Poellnitz) G. D. Rowley (2013). [2] Differs from var. scabra: L surfaces minutely scabrid, tubercles small and confluent. H. scabra var. scabra — Distr: RSA (Eastern Cape: Baviaanskloof). I: Bayer (1982: Fig. 58a); Gildenhuys (2017: 53–55, as Haworthiopsis). Incl. Haworthia tuberculata Von Poellnitz (1931)  Haworthia scabra var. tuberculata (Von Poellnitz) Halda (1997); incl. Haworthia tuberculata var. acuminata Von Poellnitz (1938); incl. Haworthia tuberculata var. subexpansa Von

809

Poellnitz (1938); incl. Haworthia tuberculata var. sublaevis Von Poellnitz (1938); incl. Haworthia tuberculata var. angustata Von Poellnitz (1940); incl. Haworthia scabra var. johanii M. Hayashi (2001)  Haworthia johanii (M. Hayashi) Breuer (2010)  Haworthiopsis scabra var. johanii (M. Hayashi) Breuer (2016); incl. Haworthia plettens Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [2] Ros stemless, slowly proliferating, to 16 cm tall, 6 cm ∅; L 12–25, to 1.6  2.2 cm, triangular-lanceolate, attenuate, almost as thick as wide, dull green covered with dust, incurved, surfaces scabrid or smooth, with or without distinct raised non-confluent concolorous tubercles; Inf sparsely branched, lax, to 48 cm; Fl 15–20, tube obcapitate, curved; ITep revolute. H. scabra var. starkiana (Von Poellnitz) M. B. Bayer (Haworthia Revisited, 197, ills., 1999). Type: RSA, Western Cape (Taylor s.n. [B [lecto — icono]: unpubl. photograph]). — Distr: RSA (Western Cape: Little Karoo, Schoemanspoort). I: Bayer (1982: Fig. 60a, as H. starkiana); Gildenhuys (2017: 58–59, as Haworthiopsis).  Haworthia starkiana Von Poellnitz (1933)  Haworthia starkiana var. starkiana (1976)  Haworthia scabra ssp. starkiana (Von Poellnitz) Halda (1997)  Haworthiopsis scabra var. starkiana (Von Poellnitz) G. D. Rowley (2013); incl. Haworthia taylorii W. F. Barker ms. (s.a.) (nom. inval., ICN Art. 29.1); incl. Haworthia smitii Von Poellnitz (1938)  Haworthia pumila var. smitii (Von Poellnitz) Halda (1997)  Haworthiopsis scabra var. smitii (Von Poellnitz) Gildenhuys & Klopper (2016). [2] Differs from var. scabra: Ros larger, forming clumps; L smooth, shiny, without tubercles, yellowish-green. Gildenhuys (2017: 58) also accepts var. smitii, which he terms a “very distint taxon”. H. semiviva (Von Poellnitz) M. B. Bayer (Haworthia Handb., 153, 1976). Type: B [lecto — icono]: photograph publ. Succulenta 22: 25, 1940. — Distr: RSA (Northern Cape, Western Cape); Nama Karoo vegetation. I: Bayer (1982: Fig. 35).

810

 Haworthia bolusii var. semiviva Von Poellnitz (1938)  Haworthia arachnoidea var. semiviva (Von Poellnitz) Halda (1997); incl. Haworthia sphaeroidea Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia victoria Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Ros stemless, rarely proliferating, 5–6 cm ∅; L 30–40, broadly ovate, 6  1.5 cm, thin, incurved, pale green, translucent and usually drying up from the tips; Inf 20–30 cm; Fl 30–35, white with green venation, broad and flat across the upper base of the tube. H. sordida Haworth (Revis. Pl. Succ., 51, 1821). Type: [neo — icono]: Salm-Dyck, Monogr. Gen. Aloes & Mesembr. 7: Aloe t. 1 (Sect. 7: 2), 1863. — Distr: RSA (Eastern Cape); mainly Albany Thicket vegetation.  Aloe sordida (Haworth) Roemer & Schultes fil. (1829)  Haworthia scabra ssp. sordida (Haworth) Halda (1997)  Haworthia scabra var. sordida (Haworth) Halda (1997)  Haworthiopsis sordida (Haworth) G. D. Rowley (2013). H. sordida var. lavrani C. L. Scott (Cact. Succ. J. (US) 53(3): 124-126, ill., 1981). Type: RSA, Eastern Cape (Hechter s.n. [PRE 61124]). — Distr: RSA (Eastern Cape: Little Karoo). I: Scott (1985: 8); Gildenhuys (2017: 67, as Haworthiopsis).  Haworthia scabra var. lavrani (C. L. Scott) Halda (1997)  Haworthia lavrani (C. L. Scott) Breuer (2010)  Haworthiopsis sordida var. lavrani (C. L. Scott) G. D. Rowley (2013). [2] Differs from var. sordida: Ros smaller, to 5 cm ∅; L shorter, recurved. H. sordida var. sordida — Distr: RSA (Eastern Cape: E Little Karoo). I: Bayer (1982: Fig. 59); Gildenhuys (2017: 64–66, as Haworthiopsis). Incl. Haworthia agavoides Zantner & Von Poellnitz (1938)  Haworthia sordida var. agavoides (Zantner & Von Poellnitz) G. G. Smith (1950)  Haworthiopsis sordida var. agavoides (Zantner & Von Poellnitz) Breuer (2016).

M. B. Bayer and E. Van Jaarsveld

[2] Ros stemless, rarely proliferating, to 15 cm tall, 8 cm ∅; L 6–15, to 15  2 cm, lanceolatedeltoid, attenuate, erect, dark grey to blackishgreen, surfaces scabrid with indistinct slightly raised non-confluent tubercles, margins obtuse; Inf sparsely branched, lax, to 40 cm; Fl tube straight; ITep revolute. H. springbokvlakensis C. L. Scott (J. South Afr. Bot. 36: 287–288, 1970). Type: RSA, Eastern Cape (Scott 245 [PRE]). — Distr: RSA (Eastern Cape: E Little Karoo); Albany Thicket vegetation. I: Bayer (1982: Fig. 37).  Haworthia retusa var. springbokvlakensis (C. L. Scott) Halda (1997); incl. Haworthia groenewaldii Breuer (2011). [1] Ros stemless, solitary, to 10 cm ∅; L 8–12, turgid, 6  1.5 cm, very rounded and retuse with translucent end-area and several short longitudinal lines, smooth; Inf 20–25 cm; Fl white with brownish venation. H. tauteae Archibald pro sp. (Flow. Pl. Afr. 25: t. 992 + text, 1946). Type: RSA, Western Cape (Taute s.n. [[lecto — icono]: l.c. t. 992]).  Haworthiopsis tauteae (Archibald) Gildenhuys & Klopper (2016). This is the naturally occurring hybrid H. viscosa  H. scabra. — [U. Eggli] H. tessellata Haworth (Philos. Mag. J. 64: 300, 1824). Type: K [lecto — icono]: ill. publ. in Cact. Succ. J. (US): 50: 75, 1978. — Distr: S Namibia, RSA (Northern Cape, Free State); Nama Karoo and Succulent Karoo vegetation. I: Bayer (1982: Fig. 61c, as H. venosa ssp.); Hildyard (2016b); Gildenhuys (2017: 44–45, as Haworthiopsis). – Fig. 12.  Aloe tessellata (Haworth) Roemer & Schultes fil. (1829)  Haworthia venosa ssp. tessellata (Haworth) M. B. Bayer (1982)  Haworthia venosa var. tessellata (Haworth) Halda (1997)  Haworthiopsis tessellata (Haworth) Boatwright & J. C. Manning (2014); incl. Haworthia parva Haworth (1824)  Aloe parva (Haworth) Roemer & Schultes fil. (1829)  Haworthia tessellata var. parva (Haworth) Baker (1880); incl. Haworthia tessellata var. inflexa Baker (1880); incl. Haworthia

Haworthia ASPHODELACEAE

811

[2] Description as for H. venosa but Ros stemless; L 8–10, subdeltoid, variously patterned. Gildenhuys (2017: 45) also accepts H. crousii at varietal level (under Haworthiopsis), which differs by being more robust and less proliferous than var. tessellata.

Fig. 12 Haworthia tessellata. (Copyright: U. Eggli)

engleri Dinter (1914)  Haworthia tessellata var. engleri (Dinter) Von Poellnitz (1938); incl. Haworthia pseudotessellata Von Poellnitz (1929); incl. Haworthia tessellata var. tuberculata Von Poellnitz (1936); incl. Haworthia minutissima Von Poellnitz (1939)  Haworthia tessellata var. minutissima (Von Poellnitz) Viveiros (1949); incl. Haworthia tessellata var. elongata van Woerden (1940); incl. Haworthia tessellata fa. brevior Resende & Von Poellnitz (1942); incl. Haworthia tessellata fa. longior Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. coriacea Resende & Von Poellnitz (1942)  Haworthia coriacea (Resende & Von Poellnitz) Breuer (2010); incl. Haworthia tessellata var. luisieri Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. obesa Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. palhinhiae Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. simplex Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. stepheniana Resende & Von Poellnitz (1942); incl. Haworthia tessellata var. velutina Resende & Von Poellnitz (1942); incl. Haworthia tessellata fa. major J. R. Brown (1947); incl. Haworthia crousii M. Hayashi (2001)  Haworthiopsis tessellata var. crousii (M. Hayashi) Gildenhuys & Klopper (2016); incl. Haworthia crausii M. Hayashi (2001) (nom. inval., ICN Art. 61.1)  Haworthiopsis tessellata var. crausii (M. Hayashi) Gildenhuys & Klopper (2016) (nom. inval., Art. 61.1); incl. Haworthia helensis Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a); incl. Haworthia mediata Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a).

H. transiens (Von Poellnitz) M. Hayashi (Haworthia Study 3: 13, 2000). Type: B [lecto — icono]: unpubl. photograph. — Distr: RSA (Eastern Cape: Baviaanskloof, Langkloof); Fynbos and Albany Thicket vegetations. I: Bayer (1982: Fig. 11d, as H. cymbiformis var.).  Haworthia planifolia var. transiens Von Poellnitz (1938)  Haworthia cymbiformis var. transiens (Von Poellnitz) M. B. Bayer (1976); incl. Haworthia cymbiformis var. brevifolia Triebner & Von Poellnitz (1938); incl. Haworthia cymbiformis var. multifolia Triebner (1938)  Haworthia cymbiformis fa. multifolia (Triebner) Pilbeam (1983); incl. Haworthia cymbiformis var. translucens Triebner & Von Poellnitz (1938); incl. Haworthia diaphana M. Hayashi (2006); incl. Haworthia klipensis Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [1] Ros similar to H. cymbiformis and to H. cooperi var. picturata; L obtuse, surface reticulate with translucent areas between the veins. H. truncata Schönland (Trans. Roy. Soc. South Africa 1: 391, 1910). Type: RSA, Western Cape (Britten s.n. [K]). — Distr: RSA (Western Cape: Little Karoo); Succulent Karoo vegetation. Mak (2003) presents a synopsis of hybrids involving this species as one parent. H. truncata var. maughanii (Von Poellnitz) B. Fearn (Nation. Cact. Succ. J. 21(1): 28–29, 1966). Type [neo]: RSA, Western Cape (Malherbe s.n. in NBG 307/40 [NBG 68307]). — Distr: RSA (Western Cape: Little Karoo, Calitzdorp). I: Bayer (1982: Fig. 24, as H. maughanii).  Haworthia maughanii Von Poellnitz (1933). [1] Differs from var. turgida: L multifarious, round in cross-section.

812

M. B. Bayer and E. Van Jaarsveld

 Haworthia hemicrypta (M. B. Bayer) M. Hayashi (2000). [1] Differs from var. variegata: Plants moderately variegated to plain; L long and slender, tending to arch out and then curve inwards.

Fig. 13 Haworthia truncata var. truncata. (Copyright: E. J. Van Jaarsveld)

H. truncata var. truncata — Distr: RSA (Western Cape: C Little Karoo). I: Bayer (1982: Fig. 39). – Fig. 13. Incl. Haworthia truncata fa. crassa Von Poellnitz (1938); incl. Haworthia truncata fa. normalis Von Poellnitz (1938); incl. Haworthia truncata fa. tenuis Von Poellnitz (1938)  Haworthia truncata var. tenuis (Von Poellnitz) M. B. Bayer (1976); incl. Haworthia truncata var. minor Breuer (2003); incl. Haworthia papillaris Breuer (2010). [1] Ros stemless, slowly proliferating; L distichous, 10–12, abruptly truncate with flat to slightly corrugated and subpellucid end-areas, 1.2–4 cm wide, 0.3–1 cm thick, dark grey-green, scabrid with minute tubercles; Inf to 20 cm; Fl 20–30, white with brownish veins. H. variegata L. Bolus (J. Bot. 67: 137, 1929). Type: RSA, Western Cape (Ferguson s.n. [BOL]). — Distr: RSA (Western Cape); Fynbos vegetation.  Haworthia chloracantha ssp. variegata (L. Bolus) Halda (1997)  Haworthia chloracantha var. variegata (L. Bolus) Halda (1997). [1] Ros stemless, proliferous, to 4 cm ∅; L 30–40, erect, slender lanceolate, dark green, variegated, margins and keel spined; Inf slender, to 35 cm, lax; Fl 15–20, greenish-white with brownish venation. H. variegata var. hemicrypta M. B. Bayer (Haworthia Revisited, 158, ills., 1999). Type: RSA, Western Cape (Burgers 2582 [NBG]). — Distr: RSA (Western Cape: Bredasdorp).

H. variegata var. modesta M. B. Bayer (Haworthia Revisited, 159, ills., 1999). Type: RSA, Western Cape (Bayer 2551 [NBG]). — Distr: RSA (Western Cape: Bredasdorp).  Haworthia modesta (M. B. Bayer) M. Hayashi (2000). [1] Differs from var. variegata: L broader and shorter and with less conspicuous spination. H. variegata var. variegata — Distr: RSA (Western Cape: Riversdale). [1] Plants strongly variegated. H. venosa (Lamarck) Haworth (Revis. Pl. Succ., 51, 1821). Type: [lecto — icono]: Commelin, Praeludia Bot. t. 29, 1703. — Distr: RSA (Northern Cape, Western Cape: lower Breede and Gouritz river valleys); Fynbos and Albany Thicket vegetations. I: Bayer (1982: Fig. 61a); Gildenhuys (2017: 46, as Haworthiopsis).  Aloe venosa Lamarck (1783)  Haworthiopsis venosa (Lamarck) G. D. Rowley (2013); incl. Aloe recurva Haworth (1804)  Apicra recurva (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthia recurva (Haworth) Haworth (1812)  Haworthia venosa ssp. recurva (Haworth) M. B. Bayer (1976); incl. Aloe tricolor Haworth (1804)  Apicra tricolor (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4); incl. Haworthia distincta N. E. Brown (1876); incl. Haworthia venosa var. oertendahlii Hjelmquist (1943); incl. Haworthia irmiae Breuer (2010) (nom. inval., ICN Art. 36.1b, 38.1a). [2] Ros usually stemless, slowly proliferating with offsets or stolons, to 3 cm tall and 5 cm ∅; L 12–20, to 10  1.5 cm, ovate-deltoid, spreading to recurved, upper surface smooth, reticulate, lower surface usually slightly scabrid; Inf sparsely branched, lax, to 35 cm; Fl 15–20, tube obcapitate; ITep revolute.

Haworthia ASPHODELACEAE

Fig. 14 Haworthia viscosa. (Copyright: E. J. Van Jaarsveld)

H. viscosa (Linné) Haworth (Synops. Pl. Succ., 90, 1812). Type: [lecto — icono]: Commelin, Praeludia Bot., 82, t. 31, 1703. — Distr: RSA (Western Cape, Eastern Cape: Little Karoo); Succulent Karoo and Albany Thicket vegetations. I: Bayer (1982: Fig. 62); Gildenhuys (2017: 60–61, as Haworthiopsis). – Fig. 14.  Aloe viscosa Linné (1753)  Apicra viscosa (Linné) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthiopsis viscosa (Linné) G. D. Rowley (2013)  Tulista viscosa (Linné) G. D. Rowley (2013) (nom. inval., ICN Art. 36.2); incl. Aloe triangularis Medikus (1784); incl. Aloe tortuosa Haworth pro sp. (1804)  Apicra tortuosa (Haworth) Willdenow (1811) (incorrect name, ICN Art. 11.4)  Haworthia tortuosa (Haworth) Haworth (1812)  Haworthiopsis tortuosa (Haworth) Gildenhuys & Klopper (2016); incl. Aloe pseudotortuosa Salm-Dyck (1817)  Haworthia pseudotortuosa (SalmDyck) Haworth (1819)  Haworthia viscosa var. pseudotortuosa (Salm-Dyck) Baker (1880)  Haworthia viscosa fa. pseudotortuosa (SalmDyck) Pilbeam (1983) (nom. inval., ICN Art. 41.5); incl. Haworthia asperiuscula Haworth (1819)  Aloe asperiuscula (Haworth) SalmDyck (1840)  Haworthia viscosa fa. asperiuscula (Haworth) Pilbeam (1983) (nom. inval., ICN Art. 41.5)  Haworthiopsis viscosa var. asperiuscula (Haworth) Breuer (2016); incl. Haworthia concinna Haworth (1819)  Aloe concinna (Haworth) Roemer & Schultes fil. (1829)  Haworthia viscosa var. concinna

813

(Haworth) Baker (1880); incl. Haworthia cordifolia Haworth (1819)  Aloe cordifolia (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia indurata Haworth (1821)  Aloe indurata (Haworth) Roemer & Schultes fil. (1829)  Aloe viscosa var. indurata (Haworth) Salm-Dyck (1836)  Haworthia viscosa var. indurata (Haworth) Baker (1880); incl. Haworthia viscosa var. major Haworth (1821)  Aloe viscosa var. major (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia viscosa var. minor Haworth (1821)  Aloe viscosa var. minor (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia viscosa var. parvifolia Haworth (1821)  Aloe viscosa var. parvifolia (Haworth) Roemer & Schultes fil. (1829); incl. Haworthia torquata Haworth (1827)  Aloe torquata (Haworth) Salm-Dyck (1836)  Haworthia viscosa var. torquata (Haworth) Baker (1880)  Haworthia viscosa fa. torquata (Haworth) Pilbeam (1983) (nom. inval., ICN Art. 41.5); incl. Aloe subtortuosa Roemer & Schultes fil. (1829); incl. Haworthia viscosa var. caespitosa Von Poellnitz (1938); incl. Haworthia viscosa var. subobtusa Von Poellnitz (1938)  Haworthia viscosa fa. subobtusa (Von Poellnitz) Pilbeam (1983) (nom. inval., ICN Art. 41.5); incl. Haworthia beanii G. G. Smith (1944)  Haworthia viscosa fa. beanii (G. G. Smith) Pilbeam (1983)  Haworthiopsis viscosa var. beanii (G. G. Smith) Breuer (2016); incl. Haworthia beanii var. minor G. G. Smith (1944); incl. Haworthia asperiuscula var. subintegra G. G. Smith (1945); incl. Haworthia viscosa var. cougaensis G. G. Smith (1945); incl. Haworthia viscosa var. viridissima G. G. Smith (1945); incl. Haworthia asperiuscula var. patagiata G. G. Smith (1946); incl. Haworthia viscosa var. quaggaensis G. G. Smith (1948); incl. Haworthia viscosa ssp. dereki-clarki Halda (1998); incl. Haworthia viscosa var. variabilis Breuer (2003)  Haworthia variabilis (Breuer) Breuer (2010)  Haworthiopsis variabilis (Breuer) Zonneveld (2014)  Haworthiopsis viscosa var. variabilis (Breuer) Gildenhuys & Klopper (2016). [2] Ros caulescent, proliferous, to 8 cm ∅, to 30 cm tall; L 20–60, to 5  1.5 cm, deltoid,

814

closely arranged in 3 rows, spreading, surfaces scabrid, tips spreading, pungent; Inf sparsely branched, lax, 15–20 cm; Fl 8–15, tube obcapitate, curved; ITep revolute. Gildenhuys (2017: 62, with ills.) also accepts var. variabilis as “one of the many known variations” of H. viscosa. H. vlokii M. B. Bayer (Haworthia Revisited, 160–161, ills., 1999). Type: RSA, Western Cape (Vlok s.n. in Venter 91/2 [NBG]). — Distr: RSA (Western Cape: Little Karoo, Swartberg Mts.); Fynbos vegetation. [1] Ros acaulescent, 4–5 cm ∅, proliferous; L spreading to suberect, surface opaque, with inconspicuous whiter dots towards the tips, margins and keel with short spines; Inf simple, 30–45 cm; Fl white and brownish-pink. H. wittebergensis W. F. Barker (J. South Afr. Bot. 8: 245, 1942). Type: RSA, Western Cape (Pieterse s.n. [NBG 68214]). — Distr: RSA (Western Cape: Laingsburg: Witteberg Mts.); mainly Fynbos vegetation. I: Bayer (1982: Fig. 43). – Fig. 15. [1] Ros stemless, slowly proliferating, to 3 cm ∅; L 20–30, long and slender, 15  0.8 cm, attenuate, conspicuously amplexicaul, greygreen with white spines on margins and keel, coriaceous; Inf slender, to 30 cm, lax; Fl 15–20, laxly arranged, white with green venation. H. woolleyi Von Poellnitz (Repert. Spec. Nov. Regni Veg. 42: 269–270, 1937). Type: B [lecto, ill.

M. B. Bayer and E. Van Jaarsveld

publ. Cact. J. (Croydon) 7: 3, 1938]. — Distr: RSA (Eastern Cape: E Little Karoo: Steytlerville); Albany Thicket vegetation. I: Bayer (1982: Fig. 63); Gildenhuys (2017: 47, as Haworthiopsis).  Haworthia venosa ssp. woolleyi (Von Poellnitz) Halda (1997)  Haworthiopsis woolleyi (Von Poellnitz) G. D. Rowley (2013)  Haworthiopsis venosa var. woolleyi (Von Poellnitz) Breuer (2016). [1] Description as for H. tessellata but Ros stemless; L 20–25, slender, attenuate. H. zantneriana Von Poellnitz (Cact. J. (Croydon) 5: 35, in clavi, 1935). Type: B [lecto, ill. publ. Desert Pl. Life 9: 90, 1937]. — Distr: RSA (Eastern Cape); Albany Thicket vegetation. I: Bayer (1982: Fig. 45).  Haworthia chloracantha var. zantneriana (Von Poellnitz) Halda (1997). [1] Ros stemless, proliferous, 5–6 cm ∅; L 20–40, lanceolate, acuminate, 6  1.2 cm, attenuate, soft, glabrous, spreading, pale green, usually with pellucid white longitudinal marks; Inf slender, to 25 cm; Fl sparsely arranged, 20–30, white with green venation. H. zantneriana var. minor M. B. Bayer (Haworthia Revisited, 164, ills., 1999). Type: RSA, Eastern Cape (Bayer 1702 [NBG]). — Distr: RSA (Eastern Cape: Willowmore, S Karoo).  Haworthia inspida Breuer (2010). [1] Differs from var. zantneriana: Ros small, to 5 cm ∅; L unmarked. H. zantneriana var. zantneriana — Distr: RSA (Eastern Cape: Willowmore, Steytlerville). [1] Plants to 8 cm tall; L with markings.

References

Fig. 15 Haworthia wittebergensis. (Copyright: E. J. Van Jaarsveld)

Battista, L. (2009) Il genere Haworthia. Piante Grasse 29 (Suppl.): 1–52, ills. map. Bayer, M. B. (1976) Haworthia handbook. A guide to the species, with identification keys and illustrations. Kirstenbosch (ZA): National Botanic Gardens of South Africa. Bayer, M. B. (1982) The new Haworthia handbook. A revised guide to the literature of the genus, with discussion of the species, identification keys and colour

Haworthia ASPHODELACEAE illustrations. Kirstenbosch (ZA): National Botanic Gardens of South Africa. Bayer, M. B. (1999) Haworthia revisited. A revision of the genus. Hatfield (ZA): Umdaus Press. Bayer, M. B. (2001) Haworthia rossouwii V.Poelln. and the demise of H. serrata Bayer. Aloe 38(1-2): 31–36, ills. Bayer, M. B. (2002a) Haworthia update. Essays on Haworthia — Volume 1. Hatfield (ZA): Umdaus Press. Bayer, M. B. (2002b) An odd population of H.[aworthia] arachnoidea var. nigricans. Aloe 38(3-4): 76–79, ills. Bayer, M. B. (2003a) Whereto Haworthia limifolia ? Aloe 40(2): 41–51, ills., map. Bayer, M. B. (2003b) Some of the interplay of Haworthia arachnoidea and H. mucronata. Haworthiad 17(3): 94–99, ills., map. Bayer, M. B. (2004a) Still on about H.[aworthia] mutica var. nigra. Haworthiad 18(1): 21–33, ills., map. Bayer, M. B. (2004b) Haworthia arachnoidea and H. nortieri. Haworthiad 18(2): 44–51, ills. Bayer, M. B. (2004c) Continuing the saga of Haworthia mutica var. nigra. Haworthiad 18(2): 52–57, ills. Bayer, M. B. (2005) Completing the saga of H. mutica var. nigra? Haworthiad 19(1): 7–13, ills. Bayer, M. B. (2006) Haworthia update. Essays on Haworthia. Volume 2. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2007a) Haworthia update. Essays on Haworthia. Volume 3. Part 1 / Part 2. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2007b) Comments on Haworthia mortonii I. Breuer. Alsterworthia Int. 7(3): 25–27, ills. Bayer, M. B. (2008) Haworthia update. Essays on Haworthia. Volume 4. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2009) Haworthia update. Essays on Haworthia, volume 5. Part 1 / Part 2. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2010a) Haworthia update. Essays on Haworthia. Volume 6. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2010b) Variants of Haworthia aristata at Kaboega. Haworthiad 24(1): 20–23, ills. Bayer, M. B. (2010c) Klip River plants in cultivation. Haworthiad 24(2): 36–37, ills. Bayer, M. B. (2010d) A case of mistaken identity. Haworthiad 24(2): 40–41, ill. Bayer, M. B. (2010e) Haworthia minima ‘opalina’. Haworthiad 24(3): 64–65, ills. Bayer, M. B. (2010f) Haworthia kingiana and H. minima variants. Haworthiad 24(3): 68–74, ills. Bayer, M. B. (2010g) Comments on Haworthiad 24 No. 2, 2010. Haworthiad 24(3): 79–86, ills. Bayer, M. B. (2012a) Haworthia update. Essays on Haworthia, Volume 7 (part 1 / 2 / 3 / 4). St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays. Bayer, M. B. (2012b) Haworthia update. Essays on Haworthia, Volume 8. St. Michaels on Wyre (GB): Alsterworthia International / Harry Mays.

815 Bayer, M. B. (2012c) New finds in Haworthia. Cact. Succ. J. (US) 84(1): 41–50, ills., map. https://doi.org/ 10.2985/0007-9367-84.1.41 Bayer, M. B. (2012d) Just what do we do with names for Haworthia ? CactusWorld 30(4): 211–214, ills. Bayer, M. B. (2013a) Plants in my collection 11: Haworthia decipiens. Haworthiad 27(2): 42–43, ills. Bayer, M. B. (2013b) The Haworthia pollinator. Alsterworthia Int. 13(1): 30, ills. Bayer, M. B. (2013c) Another Haworthia pollinator. Alsterworthia Int. 13(1): 31, ills. Bayer, M. B. & Dold, T. (2003) Ecotypes in Haworthia. Aloe 40(1): 10–15, ills., maps. Bayer, M. B. & Manning, J. C. (2012a) A rationalization of names in Haworthia. A list of species with new combinations and new synonyms. Alsterworthia Int. 12(1): 7–17. Bayer, M. B. & Manning, J. C. (2012b) The Haworthia nomenclator: A list of accepted species with some guidelines for infraspecific names. Redditch (GB): Haworthia Society. Biko’o, A. A. [& al. 2011], Plessis, D. G. C. du & Myburgh, W. J. (2011) Population size, structure and habitat features of Haworthia koelmaniorum var. mcmurtryi, an endemic plant from Mpumalanga Province, South Africa. Koedoe 53(1): 1–8, ill., map. https://doi.org/10.4102/koedoe.v53i1.1025 Bosch, P. J. A. (2004) The geological, pedological and climatical influence on variety generation in Haworthia koelmaniorum var. koelmaniorum and H. koelmaniorum var. mcmurtryi. Aloe 41(2-3): 56–60, ills., map. Breuer, I. (1998a) The world of Haworthias. Volume 1. Bibliography and annotated index. Niederzier / Homburg (DE): Ingo Breuer / Arbeitskreis für Mammillarienfreunde. Breuer, I. (1998b) Die Gattung Haworthia im Überblick. Schumannia 2: 3–74, ills., maps. Breuer, I. (2010) The genus Haworthia. Book 1. St. Michaels on Wyre (GB): Alsterworthia International. Breuer, I. (2011) The genus Haworthia. Book 2, part 1 / part 2. St. Michaels on Wyre (GB): Alsterworthia International. Breuer, I. & Metzing, D. (1997) Types of names accepted in Haworthia (Aloaceae). Taxon 46(1): 3–14. https:// doi.org/10.2307/1224286 Coopoosamy, R. M. & Naidoo, K. K. (2011) Screening of traditional utilized Haworthia limifolia for antibacterial and antifungal properties. J. Med. Pl. Res. 5(1): 109–113. Craib, C. (2007) The autoecology and status of Haworthia koelmaniorum variety mcmurtryi. Aloe 44(1): 22–27, ills. Daru, B. H. [& al. 2013], Manning, J. C., Boatwright, J. S., Maurin, O., Mclean, N., Schaefer, H., Kuzmina, M. & Bank, M. van der (2013) Molecular and morphological analysis of subfamily Alooideae (Asphodelaceae) and the inclusion of Chortolirion in Aloe. Taxon 62(1): 62–76, ills. http://www.jstor.org/stable/24389313 Gildenhuys, S. D. (2017) Haworthiopsis — an illustrated taxonomy. Aloe 53(1): 2–78, ills., map. Gildenhuys, S. D. & Klopper, R. R. (2016) A synoptic review and new infrageneric classification for the genus Haworthiopsis (Xanthorrhoeaceae: Asphodeloideae).

816 Phytotaxa 265(1): 1–26, map, key. https://doi.org/ 10.11646/phytotaxa.265.1.1 Grenier, F. (2013) Haworthia taxonomy: A practical approach. Aloe 50(3): 110-115, ills. Guillot, D. & Laguna Lumbreras, E. (2012) Algunas especies y formas horticolas escapadas de cultivo o pertenecientes a cultivos abandonados presentes en la Comunidad Valenciana (citas y aspectos históricos). Bouteloua 9: 47–55, ills. Hayashi, M. (2002) Some new species of [Haworthia] section Retusae. Haworthia Study 7: 14, ills. Hayashi, M. (2008) A system for Haworthia. Haworthia Study 19: 10–15. Hayashi, M. (2013) Total list of Haworthia cultivars. Haworthia cultivar names published in Japan since 1905. Haworthia Study 28: [i]–[vi], 1–271 pp., ills. Hildyard, B. (2016a) Variegates. Haworthiad 30(1): 15–20, ills. Hildyard, B. (2016b) Haworthia tessellata Haw. (Haworthiopsis tessellata). Haworthiad 30(3): 73–80, map, ills. Jacobsen, H. (1960) A handbook of succulent plants. London (GB): Blandford Press. Jarvis, C. (2007) Order out of chaos. Linnean plant names and their types. London (GB): The Linnean Society of London / Natural History Museum, London. Mak, H. C. K. (2003) Haworthia truncata hybrids — a preliminary survey (part 1) / part 2. Haworthiad 17(2): 63–74, (3): 103–114, ills. Manning, J. C. (2013) Haworthia margaritifera / pumila. Alsterworthia Int. 13(1): 6, ill. Manning, J. C. [& al. 2014a], Boatwright, J. S., Daru, B. H., Maurin, O. & Bank, M. van der (2014) A molecular phylogeny and generic classification of Asphodelaceae subfamily Alooideae: A final resolution of the prickly issue of polyphyly in the Alooids. Syst. Bot. 39(1): 55–74, key. https://doi.org/10.1600/0363 64414X678044 Marx, G. (2008a) Haworthia magnifica var. splendens revisited. Aloe 44(3): 70–74, ills. Marx, G. (2008b) Haworthia marxii S. D. Gildenhuys — an update. Haworthiad 22(3): 74–80, ills. Marx, G. (2009a) Haworthia emelyae, the ‘Klein Karoo Kliprosie’. Aloe 46(4): 76–95, ills., map. Marx, G. (2009b) A few notes on Haworthia devriesii Breuer. Alsterworthia Int. 9(1): 10–12, ills. Marx, G. (2011a) Haworthia uitewaaliana von Poellnitz. Aloe 48(2): 28–29, ills. Marx, G. (2011b) Haworthia splendens: Undeniably magnificent, conceivably mirabiloid, but a rarity in need of protection. Haworthiad 25(1): 3–9, ills. Marx, G. (2013a) Haworthia mirabilis (Haw.) Haw. — a medley of mystery and misinterpretation. Aloe 50(3): 98–109, ills. Marx, G. (2013b) Haworthia pollination and pollinators. Haworthiad 27(3): 59–62, ills. Marx, G. (2014a) Two new populations of Haworthia emelyae. Aloe 51(1): 20–24, ills. Marx, G. (2014b) Haworthias — nature’s naughty cultivars. A dated update regarding a few fascinating and

M. B. Bayer and E. Van Jaarsveld vindicatable retusoids. Cact. Succ. J. (US) 86(4): 136–148, ills. https://doi.org/10.2985/015.086.0401 Marx, G. (2016a) Haworthia correcta Poelln. corrected. Aloe 52(1): 18–23, ills. Marx, G. (2016b) The quick brown fox jumps over the lazy dog. Haworthiad 30(2): 31–37, ills. Mays, H. (2013) Hybrids & cultivars of the succulent Asphodelacea [sic!]. Volume 2. Haworthia. St. Michaels on Wyre (GB): Harry Mays. Molteno, S. & Smith, G. F. (2017) Notes on morphological variation in, and the biogeography of, Tulista kingiana Gideon F. Sm. & Molteno [sic!] (Asphodelaceae: Alooideae) from the south-central Western Cape Province of South Africa. Bradleya 35: 46–57, ills., map. https://doi.org/10.25223/brad.n35.2017.a5 North, G. B. [& al. 2008], Brinton, E. K. & Garrett, T. Y. (2008) Contractile roots in succulent monocots: Convergence, divergence and adaptation to limited rainfall. Pl. Cell Environm. 31: 1179–1189, ills. https://doi.org/ 10.1111/j.1365-3040.2008.01832.x Ramdhani, S. [& al. 2009], Bayer, M. B., Cowling, R. M. & Barker, N. P. (2009) Molecular phylogenetic studies in Haworthia Duval (Asphodelaceae). Alsterworthia Int. 9(3): 13–17, ills. Ramdhani, S. [& al. 2011], Barker, N. P. & Cowling, R. M. (2011) Revisiting monophyly in Haworthia Duval (Asphodelaceae): Incongruence, hybridization and contemporary speciation. Taxon 60(4): 1001–1014. https://www.jstor.org/stable/41317322 Rowley, G. D. (2013a) Generic concepts in the Alooideae — part 3. The phylogenetic story. Alsterworthia Int. Spec. Issue 10: 3–6. Schulz, R. (2009) Haworthia for the collector. San Bruno (US): Schulz Publishing. Scott, C. L. (1977) The identity of Haworthia arachnoidea (L.) Duval. Cact. Succ. J. (US) 49(5): 205–208. Scott, C. L. (1980) Haworthia aristata Haw. Nation. Cact. Succ. J. 35(1): 11–13. Scott, C. L. (1985) The genus Haworthia (Liliaceae), a taxonomic revision. Johannesburg (ZA): Aloe Books. Scott, C. L. (1994a) A new species of Haworthia, H. reddii. Cact. Succ. J. (US) 66(4): 182–183, ill. Scott, C. L. (1994b) A new species of Haworthia (Asphodelaceae) — H. pringlei. Bradleya 12: 103–104. https://doi.org/10.25223/brad.n12.1994.a7 Scott, C. L. (1996) A new species of Haworthia (Asphodelaceae) — H. venusta. Bradleya 14: 87–88, ills. https://doi.org/10.25223/brad.n14.1996.a12 Scott, M. (2011) General notes on the phenotypes and morphological features of Haworthia maxima (Haw.) Duval. Aloe 48(2): 42–45, ills. Smith, G. F. [& al. 2002], Wyk, B-E. van, Steyn, E. M. A. & Breuer, I. (2002) Infrageneric classification of Haworthia (Aloaceae): Perspectives from nectar sugar analysis. Syst. & Geogr. Pl. 71(2): 391–397. www.jstor. org/stable/3668687 Treutlein, J. [& al. 2003b], Smith, G. F., Wyk, B-E. van & Wink, M. (2003) Evidence for the polyphyly of Haworthia (Asphodelaceae subfamily Alooideae;

Haworthia ASPHODELACEAE Asparagales) inferred from nucleotide sequences of rbcL, matK, ITS1 and genomic fingerprinting with ISSR-PCR. Pl. Biol. (Stuttgart) 5(5): 513–521. https:// doi.org/10.1055/s-2003-44793 Vosa, C. G. (2005) On the classification of some species of the genus Haworthia, subgenus Haworthia (Asphodelaceae). Caryologia 57(4): 395–399, ills. https://doi.org/10.1080/ 00087114.2004.10589422 Vries, V de. (2010) Haworthia vincentii — east of Albertinia. Haworthiad 24(2): 38–39, ills. Zietsman, P. C. & Smith, G. F. (2010) Notes on African plants. Asphodelaceae. Occurence of Haworthia

817 bolusii var. blackbeardiana in the Free State, South Africa. Bothalia 40(1): 58, ill. http://hdl.handle. net/2263/16110 Zonneveld, B. J. M. (2015) Nuclear genome sizes of 343 accessions of wild collected Haworthia and Astroloba (Asphodelaceae, Alooideae), compared with the genome sizes of Chortolirion, Gasteria and 83 Aloe species. Pl. Syst. Evol. 301(3): 931–953. https://doi.org/10.1007/s00606-014-1127-4 Scott, C. L. (1995) A new species of Haworthia (Asphodelaceae) - H. joeyae. Bradleya 13: 80-81, ills. https://doi.org/10.25223/brad.n13.1995.a

Kniphofia ASPHODELACEAE G. F. Smith and E. Figueiredo

Kniphofia Moench (Methodus, 631, 1794). Type: Kniphofia alooides Moench [nom. illeg.,  Kniphofia uvaria (Linné) Oken]. — Asphodeloideae — Lit: Baker (1896: 275, Flora Capensis); Berger (1908: 31–32, synopsis); Codd (1968: revision RSA); Marais (1973: revision tropical Africa); Kativu (1996: 28, revision Flora Zambesiaca area). Distr: Mostly continental Africa but extending with 1 species each to Madagascar and the Arabian Peninsula (Yemen), esp. widespread in S Africa in all countries except Namibia and Botswana. Etym: For Johann Jeremias (Hieronymus) Kniphof (1704–1765), German physician, naturalist and botanist. Incl. Notosceptrum Bentham (1883). Type: Kniphofia benguellensis Baker [see Codd, Taxon 34: 510, 1985 on the typification]. Herbaceous perennials, tufted or solitary from a thick, simple or branched rhizome, rarely somewhat caulescent, up to 2 ( 4) m high; L soft (mesophytic), rarely succulent (K. typhoides), rosulate, rarely distichous, linear, long-tapered to the apex,

G. F. Smith (*) · E. Figueiredo Department of Botany, Nelson Mandela University, Port Elizabeth, Eastern Cape, South Africa Centre for Functional Ecology, Departamento de Ciências da Vida, Calçada Martim de Freitas, Universidade de Coimbra, Coimbra, Portugal e-mail: [email protected]; epnfigueiredo@gmail. com

V-shaped in cross section, usually carinate, margins smooth or minutely serrulate; Inf a terminal, dense to lax, many-flowered, subspicate, simple or very rarely branched raceme; Bra scarious, persistent, longer than the short pedicels; Ped inconspicuously articulated;. Fl spreading or pendulous; Per deciduous, tube campanulate to cylindrical, lobes short, white, yellow, brownish or various shades of red; St as long as or longer than the perianth at anthesis; Ov sessile, with many axile ovules; Sty filiform, usually finally exserted; Sti capitate, minute; Fr a capsule, globose to ovoid, often triquetrous, chartaceous when dry; Se somewhat flattened, acutely 3-angled or winged; endosperm fleshy. — Cytology: x = 6. In one of the first comprehensive treatments of Kniphofia, Berger (1908) recognised 67 species and some varieties. He upheld the small genus Notosceptrum based on floral characters, but Codd (1967) merged it with Kniphofia. For Kniphofia in South Africa, Codd (1968) recognised 45 species and some subspecies, while Marais (1973) treated 22 tropical African species. Kativu (1996) and Kativu (2001: 26–33) included eight species for the Flora Zambesiaca area, while Whitehouse (2002) recorded eight species of Kniphofia from the Flora of Tropical East Africa region. In the most recently published regional treatment of the genus, Codd (2005) documented 48 species and 6 subspecies from southern Africa (Swaziland, Lesotho and South Africa). In total, Kniphofia consists of about 70 species with a predominantly African distribution, single outliers occuring on

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_57

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Fig. 1 Kniphofia typhoides. (Copyright: G. Fritz)

Madagascar and in Yemen. In terms of number of species, the genus is best represented in southern Africa. Ramdhani & al. (2006) and Ramdhani & al. (2009) discussed the phylogeny of Kniphofia. The genus essentially consists of herbaceous perennials that prefer moist habitats, where they often occur in vast numbers in dense colonies. Several species of Kniphofia, commonly and in the gardening trade known as red-hot pokers, are of considerable horticultural value in parts of the mild-temperate Northern Hemisphere where, for example in the United Kingdom, numerous very attractive hybrids are offered for sale. Some of the species and cultivars have value as cut flowers, again more so in Europe than in its native Africa. Succulence has evolved in a single South African species of the genus: K. typhoides Codd (Flow. Pl. Afr. 36(3–4): t. 1424 + text, 1964). Type: RSA, KwaZulu-Natal (K. Saunders s.n. in Medley Wood 3895 [K]). —

Distr: RSA (KwaZulu-Natal, Mpumalanga, Gauteng, North-West, Free State); almost invariably found in heavy black clay soils in usually moist depressions, watercourses and seepage areas. – Fig. 1.  Notosceptrum natalense Baker (1896). Plants with one to several stems from a short, thick rhizome; L  distichous, glaucous, 35–65  0.8–2 cm, shallowly channelled above, keeled below, often spirally twisted, succulent, margins smooth; Inf peduncle 40–70 cm; raceme cylindrical, very dense, 15–30  1.5–2.5 cm, rounded at the apex; buds erect, dark brown; Fl ascending to spreading, distinctly sugar-scented; Bra obovate or subrotund to ovate, 4–6 mm, rounded at the apex, margin entire to minutely denticulate; Ped up to 0.5 mm; Per subcampanulate or shortly cylindrical, 4.5–6.5 mm long, brown to purplish-brown, lobes ovate to rounded, 1.5 mm, not spreading; St exserted by 3–4 mm at anthesis; Fr ovoid, 5–6 mm.

Kniphofia ASPHODELACEAE

Smith & al. (1997) were the first to point out that K. typhoides is a typical leaf succulent. In its natural habitat it flowers from late summer to early autumn (February to March).

References Baker, J. G. (1896) Liliaceae. In: Thiselton-Dyer, W. T. (ed.): Flora Capensis; 6(2): 253–384. London (GB): L. Reeve & Co. http://biodiversitylibrary.org/page/ 712288 Berger, A. (1908) Liliaceae — Asphodeloideae — Aloineae. In: Engler, A. (ed.): Das Pflanzenreich IV.38 (Heft 33). Leipzig (DE): Wilhelm Engelmann. http://www.biodiversitylibrary.org/bibliography/72397#/ Codd, L. E. (1967) The status of the genus Notosceptrum Benth. (Liliaceae). Bot. Not. 120: 41–45. Codd, L. E. (1968) The South African species of Kniphofia. Bothalia 9(3–4): 363–513. https://doi.org/ 10.4102/abc.v9i3/4.1790 Codd, L. E. (2005) Kniphofia. In: Germishuizen, G. & Momberg, B. A. (eds.): Flora of southern Africa, 5(1: Fasc. 2): Asphodelaceae (first part). Pretoria (ZA): South African National Biodiversity Institute.

821 Kativu, S. (1996) Asphodelaceae of the Flora Zambesiaca area. Kirkia 16: 27–53. Kativu, S. (2001) Asphodelaceae. In: Pope, G. V. (ed.): Flora Zambesiaca; 12(3): 25–48, keys. Richmond (GB): Royal Botanic Gardens Kew. Marais, W. (1973) A revision of the tropical species of Kniphofia (Liliaceae). Kew Bull. 28: 465–483. https:// doi.org/10.2307/4108891 Ramdhani, S., Barker, N. P. & Baijnath, H. (2006) Phylogenetics of the genus Kniphofia Moench (Asphodelaceae). In: Ghazanfar, S. A. & Beentje, H. J. (eds.): Taxonomy and Ecology of African Plants: their conservation and sustainable use (Proceedings of the 17th AETFAT Congress); pp. 559–573. Richmond (GB): Royal Botanic Gardens, Kew. Ramdhani, S., Barker, N. P. & Baijnath, H. (2009) Rampant non-monophyly of species in Kniphofia Moench (Asphodelaceae) suggests a recent Afromontane radiation. Taxon 58: 1141–1152. https://www.jstor.org/sta ble/27757008 Smith, G. F. [& al. 1997], Jaarsveld, E. J. van, Arnold, T. H., Steffens, F. E., Dixon, R. D. & Retief, J. A. (eds.) (1997) List of Southern African succulent plants. Pretoria (ZA): Umdaus Press. Whitehouse, C. M. (2002) Asphodelaceae. In: Beentje, H. J. (ed.): Flora of Tropical East Africa. (Rotterdam (NL)/Brookfield (US): A. A. Balkema.

Maysara ASPHODELACEAE U. Eggli

Maysara D. M. Cumming (Haworthiad 13(3): 115, 1999). — Alooideae — Lit: Cumming (1999). Etym: For Harry Mays (fl. 1999), English succulent plant enthusiast and at the time editor of the journal “Haworthiad”; plus the suffix ‘-ara’, indicating plurigeneric hybrids. Incl. Rowleyara D. M. Cumming (1999) (nom. illeg., Art. 53.1).

= Astroloba  Gasteria  Haworthia. The only known combination has not been named formally.

References Cumming, D. M. (1999) New nothogenera proposed for four trigeneric hybrids within Aloaceae. Haworthiad 13(1): 20–21.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_69

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Poellneria ASPHODELACEAE U. Eggli

Poellneria G. D. Rowley (Nation. Cact. Succ. J. 28(1): 7, 1973). — Alooideae = Poellnitzia  Gasteria. No formally named taxa are known, and Rowley (1982: 49) lists a single combination with an unidentified Gasteria as the other parent.

References Rowley, G. D. (1982) Intergeneric hybrids in succulents. Nation. Cact. Succ. J. 37(1): 2–6, (2): 45–49, (3): 76–80, (4): 119, ills.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_70

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Poellnitzia ASPHODELACEAE G. F. Smith

Poellnitzia Uitewaal (Succulenta 22: 61, 1940). Type: Apicra rubriflora L. Bolus. — Alooideae — Lit: Smith (1994: taxonomic history); Smith (1995: monograph). Distr: RSA (Western Cape: Worcester-Robertson Karoo). Etym: For Joseph Karl L. A. Von Poellnitz (1896–1945), German agriculturist and botanist in Thüringen, strongly interested in succulent plant systematics. Herbaceous perennials, caulescent, proliferous, stems to 25 cm, 1 cm ∅; Ros of 4-ranked leaves in spiralling rows; L thick, hard, squarrose-imbricate, dark green, 2–4  2 cm broad near the base, to 5 mm thick, margin minutely scabrid, apex pungent-acuminate; Inf racemose, simple, born horizontally, to 50 cm, sterile part bracteate, with secund erect flowers; Ped short, erect, persistent; Tep orange to red with dark green tips, basally united into a narrow elongate tube 20 mm long and 3 mm ∅ with the upper slightly decurved, free parts of the tepals closely adhering, apically spoon-shaped, connivent; St 6, inserted in the perianth tube, 18 mm; Fil light green; Anth yellow, dorsifixed, dehiscing longitudinally, introrse; Ov superior, green, sessile, 6–7 mm, 3 mm ∅; Sty white, straight, G. F. Smith (*) Department of Botany, Nelson Mandela University, Port Elizabeth, Eastern Cape, South Africa Centre for Functional Ecology, Departamento de Ciências da Vida, Calçada Martim de Freitas, Universidade de Coimbra, Coimbra, Portugal e-mail: [email protected]

capitate, 12 mm; Fr 3-locular capsules, cylindrical, apically retuse, dehiscing loculicidally, 16 mm, 3–4 mm ∅; Se angled, dark brown to black, shortly winged, 4 mm. —Cytology: 2n = 14 (Smith 1991). On vegetative morphological grounds, this monotypic genus shows affinities with some representatives of Aloe, Astroloba and Haworthia, and the molecular phylogeny studies of Daru & al. (2013) and Manning & al. (2014a) indeed place Poellnitzia as sister to Astroloba species. However, the flower morphology is unique in the family in that the dark green free portions of the tepals are connivent with the very tips scarcely separated. Cumming (2006) found that Poellnitzia does not produce hybrids with Astroloba, Haworthia or Aloe, and in crosses with Gasteria, seed only forms on Gasteria, but never on Poellnitzia. Cumming (2006) therefore suggests that Poellnitzia is best treated as a monotypic genus. However, further investigations of Poellnitzia and its relatives may place it in one of the other Alooideae genera. In this regard, Manning & Smith (2000) suggested the placement of Poellnitzia in Astroloba, as the tepals are imbricate as in species of Astroloba, and the flower characters represent a bird-pollination syndrome. P. rubriflora is restricted to the WorcesterRobertson Karoo centre of endemism in South Africa (Wyk & Smith 2001). P. rubriflora (L. Bolus) Uitewaal (Succulenta 22: 61, 1940). Type: RSA, Western Cape (Smith

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_58

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Fig. 1 Poellnitzia rubriflora. (Copyright: U. Eggli)

s.n. [BOL 45213]). — Distr: RSA (Western Cape: Worcester-Robertson Karoo: Robertson, Bonnievale and McGregor districts); predominantly winter rainfall areas, karroid shrublands, 400–800 m. I: Smith (1994: 74); Smith & al. (1995). – Figs. 1 and 2.  Apicra rubriflora L. Bolus (1920)  Haworthia rubriflora (L. Bolus) Parr (1971)  Aloe rubriflora (L. Bolus) G. D. Rowley (1981)  Astroloba rubriflora (L. Bolus) G. F. Smith & J. C. Manning (2000) (incorrect name, ICN Art. 11.4)  Tulista rubriflora (L. Bolus) G. D. Rowley (2013); incl. Apicra jacobseniana Von Poellnitz (1939)  Poellnitzia rubriflora var. jacobseniana (Von Poellnitz) Uitewaal (1955)  Haworthia rubriflora var. jacobseniana (Von Poellnitz) Parr (1972) (nom. inval., ICN Art. 41.5). Description as for the genus.

References Cumming, D. M. (2006) Poellnitzia Uitewaal: Inferred relationships from limited hybridization. Alsterworthia Int. 6(1): 13–16. Daru, B. H. [& al. 2013], Manning, J. C., Boatwright, J. S., Maurin, O., Mclean, N., Schaefer, H., Kuzmina, M. &

G. F. Smith

Fig. 2 Poellnitzia rubriflora. (Copyright: U. Eggli) Bank, M. van der (2003) Molecular and morphological analysis of subfamily Alooideae (Asphodelaceae) and the inclusion of Chortolirion in Aloe. Taxon 62(1): 62–76, ills. http://www.jstor.org/ stable/24389313. Manning, J. C. & Smith, G. F. (2000) The genus Poellnitzia included in Astroloba. Bothalia 30(1): 53. http://www. abcjournal.org/index.php/ABC/article/view/540. Manning, J. C. [& al. 2014a], Boatwright, J. S., Daru, B. H., Maurin, O. & Bank, M. van der (2014) A molecular phylogeny and generic classification of Asphodelaceae subfamily Alooideae: A final resolution of the prickly issue of polyphyly in the Alooids. Syst. Bot. 39(1): 55–74, key. https://doi.org/10.1600/ 036364414X678044. Smith, G. F. (1991) The chromosomes of Chortolirion and Poellnitzia (Asphodelaceae: Alooideae). Bothalia 21(2): 171–175, ills. http://abcjournal.org/index.php/ABC/arti cle/view/884. Smith, G. F. (1994) Taxonomic history of Poellnitzia Uitewaal, a unispecific genus of Alooideae (Asphodelaceae). Haseltonia 2: 74–78, ills. Smith, G. F. (1995) FSA contributions 3: Asphodelaceae/ Aloaceae, 1028000 Poellnitzia. Bothalia 25(1): 35–36, ill., map. http://abcjournal.org/index.php/ABC/article/ view/709. Smith, G. F., Wyk, B-E. van & Condy, G. (1995) Poellnitzia rubriflora. Flow. Pl. Afr. 54: 94–98, t. 2119, map. Wyk, A. E. (B.) van & Smith, G. F. (2001) Regions of floristic endemism in Southern Africa. A review with emphasis on succulents. Hatfield (ZA): Umdaus Press.

Trachyandra ASPHODELACEAE E. Van Jaarsveld

Trachyandra Kunth (Enum. Pl. 4: 573, 1843). Type: Anthericum hispidum Linné. — Asphodeloideae — Lit: Obermeyer (1962: revision RSA). Distr: S Africa (esp. SW RSA) extending to tropical Africa (Somalia, Ethiopia, Yemen), 1 species in Madagascar. Etym: Gr. ‘trachys’, rough; and Gr. ‘aner, andros’, male; for the rough filaments. Incl. Lepicaulon Rafinesque (1837) (nomen rejiciendum, Art. 56.1). Type: Anthericum squameum Linné fil. Incl. Obsitila Rafinesque (1837) (nomen rejiciendum, Art. 56.1). Type: Anthericum hispidum Linné [lectotype, designated by Merrill, Ind. Raf., 92, 1949 (ICBN 2006: 283)]. Incl. Dilanthes Salisbury (1866). Type: Dilanthes revoluta Salisbury [typification by inference, only element included]. Incl. Liriothamnus Schlechter (1924). Type: Anthericum involucratum Baker. Plants variable, perennial, glabrous to glandularpubescent rosulate leaf succulents, acaulescent, geophytic or shrubby to 2 m tall; Rstock a tuberous caudex or as erect rhizome; R fusiform to fibrous, terete; stem erect, woody and covered with leaf

bases; L dimorphic or uniform, lamina flat or triangular in cross-section to terete, often linear-lorate, rarely canaliculate, straight or undulating; Inf racemose, occasionally paniculate, peduncle terete, bracteose; Fl solitary, spreading to pendulous; Per rotate, often becoming recurved; Tep 6 in 2 whorls, white to mauve, OTep narrower than ITep; St 6, adnate to the tepal bases; Fil scabrid; Anth dorsifixed, introrse; Ov globose, superior, 3-locular; Sty terete; Sti capitate; Fr loculicidal globose capsules, drooping; Se dark grey to black, smooth to papillate, becoming glutinous. A genus of 65 species (Whitehouse 2002), most of which are geophytes with some claim to succulence. Trachyandra has a wide distribution in S Africa but shows a concentration of species in the winter rainfall regions of RSA. They are seldom cultivated but are easy and should be kept dry during the summer months. T. adamsonii is the largest in the genus, and has a shrub-like growth form superficially resembling climbing species of Aloe such as A. ciliaris. Most if not all species exhibit considerable succulence. Some species show distinct basal caudices, esp. those of Sect. Liriothamnus, while those of the other sections are usually without a caudex. Only a limited number of species is in cultivation, and for this reason, only a small selection is covered below by way of example.

E. Van Jaarsveld (*) Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_59

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The genus is divided into 3 sections according to Obermeyer (1962: 717): [1] Sect. Liriothamnus (Schlechter) Obermeyer 1962: Plants glabrous; outer L not reduced to scales; Inf mostly simple. [2] Sect. Trachyandra: Plants glabrous; outer L reduced to scales; Inf mostly branched. [3] Sect. Glandulifera Obermeyer 1962: Plants glandular-pubescent. T. adamsonii (Compton) Obermeyer (Bothalia 7 (4): 720, 1962). Type: RSA, Western Cape (Compton s.n. in NBG 318/22 [NBG]). — Distr: RSA (Northern Cape, Western Cape); dry Olifants Sandstone Fynbos and Succulent Karoo vegetations, on quartzitic sandstones.  Liriothamnus adamsonii Compton (1931). [1] Erect, sparingly branched, woody, rosulate shrubs to 2 m; caudex depressed, globose above and tapering into the stem, flat below with grey bark; R succulent, terete, 5 mm ∅; Br 1–2 cm ∅, grey with peeling leaf remains, becoming smooth at the base; L glaucous, 16–30  1.5–3.5 cm, striate, crowded at the stem tips, linear-lanceolate, flat, lower face slightly keeled, upper face basally channelled, margin denticulate, apex acuminate; Inf ascending, racemose, to 50 cm, with a pair of side branches at the base; Bra triangular to ovateacuminate, 2–10  1–3 mm; Ped erect, 6–11 mm; Fl white with yellow eye at the tepal bases, to 30 mm ∅; OTep 11  2.5 mm, linear-lanceolate; ITep 10  3.5 mm; St 8 mm; Ov oblong, 2 mm; Fr erect, ovoid, 12 mm; Se grey, oblong, 3 mm. T. aridimontana J. C. Manning (South Afr. J. Bot. 56(1): 1–5, ills., 1990). Type: RSA, Northern Cape (Oliver & al. 478 [PRE]). — Distr: RSA (Northern Cape: Richtersveld); Rosyntjieberg Succulent Shrubland and Succulent Karoo vegetations. [1] Erect sparingly branched rosulate deciduous succulent herbs to 35 cm; caudex depressed, globose above and tapering into the stem, flat below with grey bark; R succulent, terete, 3 mm ∅; Br 0.4–1.5 cm ∅, bark grey; L glaucous, crowded at the stem tips, 3 – 6, linear-lanceolate, 3–12  0.3–0.7 cm ∅, subterete but

E. Van Jaarsveld

dorsiventrally flattened, lower face convex, upper face flat to canaliculate, margin denticulate, apex mucronate; Inf ascending, racemose, to 22 cm; Bra ovate to lanceolate-acuminate, to 8 mm; Ped patent, 1–5 mm; Fl white, 15–20 mm ∅; OTep linear, 11–13  2–3.5 mm; ITep ovate, 11–13  3–5.5 mm; St 8–9 mm; Ov ovoid, 1.5 mm; Sty 7–8 mm; Fr erect, narrowly ovoid, 8–12  4 mm; Se black, oblong, 3  1.5 mm. T. ciliata (Linné fil.) Kunth (Enum. Pl. 4: 585, 1843). Type: RSA, “Cape Prov.” (Thunberg s.n. [UPS]). — Distr: Namibia, RSA (Northern Cape, Western Cape, Eastern Cape); Sandstone Fynbos and Renosterveld vegetations. – Fig. 1.  Anthericum ciliatum Linné fil. (1781)  Bulbine ciliata (Linné fil.) Link (1821); incl. Anthericum longifolium Jacquin (1786)  Phalangium longifolium (Jacquin) Poiret (1804)  Trachyandra longifolia (Jacquin) Kunth (1843); incl. Anthericum canaliculatum Aiton (1789)  Phalangium canaliculatum (Aiton) Poiret (1804)  Bulbine canaliculata (Aiton) Sprengel (1825)  Trachyandra canaliculata (Aiton) Kunth (1843); incl. Anthericum vespertinum Jacquin (1804)  Phalangium vespertinum (Jacquin) Poiret (1804)  Trachyandra vespertina (Jacquin) Kunth (1843); incl. Anthericum blepharophoron Roemer & Schultes (1829)  Trachyandra blepharophora (Roemer & Schultes) Kunth (1843); incl. Trachyandra bracteosa Kunth (1843); incl. Anthericum recurvatum Dinter (1931); incl.

Fig. 1 Trachyandra ciliata. (Copyright: E. Van Jaarsveld)

Trachyandra ASPHODELACEAE

Anthericum hamatum Von Poellnitz (1942); incl. Anthericum maculatum Von Poellnitz (1942); incl. Anthericum pilosiflorum Von Poellnitz (1942); incl. Anthericum pilosiflorum var. subpapillosum Von Poellnitz (1942); incl. Anthericum spongiosum Von Poellnitz (1942). [2] Acaulescent fast-growing geophytes, rosulate, caudex absent; R fleshy, terete, spreading; L linear, to 1 m, to 2.5 cm ∅, upper face flat, lower face distinctly keeled, softly succulent, margins ciliate; Inf racemose, ascending-spreading, to 50 cm, with a pair of side branches at the base; peduncle pubescent, becoming glabrous; Bra to 10 mm, cymbiform, auriculate, younger bracts imbricate, subulate; Fl white, translucent, 20 mm ∅, Tep recurved; OTep linear, 10  2 mm; ITep ovate, 10  3 mm; St 8–9 mm; Ov globose, 1.5 mm; Sty 7–8 mm; Fr globose, to 14 mm ∅; Se black, rough. T. falcata (Linné fil.) Kunth (Enum. Pl. 4: 486, 1843). Type: RSA, “Cape Prov.” (Thunberg s.n. [UPS]). — Distr: S Namibia, RSA (Northern Cape, Western Cape); Succulent Karoo vegetation, sandy areas, winter- to spring-flowering. Anthericum falcatum Linné fil. (1782)  Bulbine falcata (Linné fil.) Schultes fil. (1829); incl. Chlorophytum drepanophyllum Baker (1897)  Anthericum drepanophyllum (Baker) Schlechter (1907); incl. Anthericum weissianum Dinter (1931). [2] Ascending, evergreen, from a reduced rhizome, to 60 cm tall; R numerous, spreading, swollen during spring; shoots incl. leaves and peduncle base surrounded by membranous scales; L up to 4, distichous, 30  2–3 cm, flat, semi-succulent, falcately curved, glabrous or shortly pubescent, margins minutely ciliate; Inf ascending sparingly branched racemes (or rarely unbranched) longer than the leaves, 30–50 cm, peduncle  of the same length as the flowering part, hairy at first becoming glabrous; lower Bra amplexicaul, short, forming a wide ascending collar around the axis, apiculate; fertile Bra widely ovate, subulate, membranous, white with a brown patch near the tip, closely imbricate in bud; Ped 10–15 mm; Fl white to pale mauve; Tep 12 mm; Fil scabrid; Ov with 8–10 ovules; Fr

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turbinate, 10–12 mm, apiculate; Se grey with raised hyaline margins. Well known as “veldkool”; the young soft inflorescences are eaten in the same way as asparagus and the species is an important food plant of the Khoi. T. involucrata (Baker) Obermeyer (Bothalia 7 (4): 721, 1962). Type: RSA, Northern Cape (Drège 2681 [K, L, PRE [photo]]). — Distr: RSA (Northern Cape); Succulent Karoo, rocks.  Anthericum involucratum Baker (1876)  Liriothamnus involucratus (Baker) Schlechter (1924). [1] Erect sparingly branched rosulate deciduous succulent gnarled shrublets to 60 cm; caudex depressed, globose above and tapering into the stem, flat below with grey bark; R succulent, terete, 3 mm ∅; Br 5–15 mm ∅, bark grey; L glaucous, crowded at the stem tips, 3–6, linear-lanceolate, 17  0.6 cm ∅, subterete, canaliculate, denticulate at the base, apex mucronate; Inf ascending, racemose, to 26 cm; Bra ovate to lanceolate-acuminate, to 7 mm, clasping; Ped recurved in fruit, 5–10 mm; Fl white, 15–20 mm ∅; OTep linear, 10–14  2–2.5 mm; ITep ovate, 10–14  3–4 mm; St 8–9 mm; Ov oblong to globose, 1.5 mm; Sty 7–10 mm; Fr erect, narrowly ovoid, 10–19  2–3 mm; Se black, ovoid, 3  1.5 mm. T. tabularis (Baker) Obermeyer (Bothalia 7 (4): 730, 1962). Type: RSA, Western Cape (Bolus 4726 [K, BOL]). — Distr: RSA (Western Cape: mainly Table Mountain); Fynbos, sandstone cliffs, 500–1000 m, summer-flowering.  Anthericum tabulare Baker (1897); incl. Anthericum glabrum Adamson (1941); incl. Anthericum palustre Adamson (1941). [2] First solitary, then dividing to form dense clusters; rhizome herbaceous, subterranean, wiry, to 30 cm; R yellow; outer L reduced to whitishtranslucent papery scales; L 1–8, succulent, bright green and reddish at the base, pendent, terete in young plants, becoming flat in mature plants, 75  0.25–0.7 cm, surface smooth, faintly striate, slightly translucent, margin entire to obscurely denticulate; Inf solitary or loose panicles up to

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5-branched, peduncle arcuate, to 50 cm, curved upwards; Ped 5–7 mm; Fl white, sweetly scented,  30 mm ∅ at anthesis; Tep 14 mm, with reddishbrown midribs; OTep oblanceolate; ITep linear to oblanceolate; Fr globose, 5 mm ∅; Se black, angular, 2.5–3 mm ∅. T. tortilis (Baker) Obermeyer (Bothalia 7(4): 745, 1962). Type: RSA, Western Cape (Schlechter 4846 [Z, PRE]). — Lit: Jaarsveld (1991); Boatwright & Manning (2010). Distr: RSA (Northern Cape, Western Cape); Succulent Karoo, rocks.  Anthericum tortile Baker (1904); incl. Anthericum salteri Leighton (1938); incl. Anthericum oocarpum Schlechter ex Von Poellnitz (1942). [2] Acaulescent geophytes to 15 cm tall with a subterranean tuber crowned with broad scale leaves; R swollen, fused to the elongate tuber, spreading, terete; L 3–6, linear, glaucous, widely spreading, transversely and plicately folded, 6–10  0.2 cm, glabrous or sparingly pubescent, flat, margin entire; Inf lax ascending divaricate panicles to 9.5 cm, with up to 5 pairs of side

E. Van Jaarsveld

branches; peduncle pubescent becoming glabrous, basally arcuate; Bra to 3 mm, ovate-lanceolate, cymbiform, auriculate; Ped to 5 mm; Fl white to pale pink, 15 mm ∅; all Tep linear-obovate, 5  2 mm; St 2–3 mm; Ov globose, 0.75 mm ∅; Sty 2–3 mm; Fr linear-ovoid, 7 mm; Se ridged.

References Boatwright, J. S. & Manning, J. C. (2010) Notes on the genus Trachyandra (Asphodelaceae: Asphodeloideae) 1: A review of the T. thyrsoidea group (section Trachyandra), including three new species from the Northern Cape. South Afr. J. Bot. 76(3): 499–510, ills., map, key. https://doi.org/10.4102/abc.v40i1.183. Jaarsveld, E. J. van (1991) The subfamily Asphodeloideae with specific reference to Trachyandra tortilis. Cact. Succ. J. (US) 63(4): 196–199, ills. Obermeyer, A. A. (1962) A revision of the South African species of Anthericum, Chlorophytum and Trachyandra. Bothalia 7(4): 669–767, keys, ills. https://doi.org/ 10.4102/abc.v7i4.1679. Whitehouse, C. M. (2002) Asphodelaceae. In: Beentje, H. J. (ed.): Flora of Tropical East Africa. Rotterdam (NL) / Brookfield (US): A. A. Balkema.

Part VII The Family Bromeliaceae

Bromeliaceae U. Eggli and E. J. Gouda

Including Tillandsiaceae Wilbread. Perennial small to large herbs or rarely somewhat shrubby, terrestrial or often epiphytic with caulescent or sessile small to large Ros; L spiral (rarely distichous), simple, entire or serrate, basally often broadened and  sheathing, often leathery or succulent, usually covered with peltate, shortly stalked, whitish, multicellular scales; Inf terminal (and rosettes often monocarpic), seemingly lateral (Hechtia p.p.) or more rarely lateral (Dyckia), pedunculate or sessile, simple or compound panicles, racemes, spikes or congested heads; Bra often brightly coloured; Fl bisexual or rarely unisexual and plants monoecious (Cryptanthus p.p., Dyckia p.p.) or dioecious (Catopsis, Hechtia), regular or almost so; Per a perigone, with 3 outer Tep (referred to as sepals, usually smaller and green, rarely petaloid) and 3 inner free or basally united Tep (referred to as petals), sometimes with a basallateral appendage (termed scale or nectary scale) or longitudinal callosities (termed lateral folds in some literature); St 6 in 2 series; Fil united to each other or to the tepals, or free; Ov of 3 carpels,

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] E. J. Gouda Curator University Botanic Gardens, Utrecht, The Netherlands e-mail: [email protected]

superior to inferior, with septal nectaries and axile placentation, with 2 to many ovules per locule; Fr longitudinally dehiscing 3-parted capsules, or fleshy berries, rarely the individual fruits of an inflorescence united into a fleshy collective fruit (Ananas); Se various, often winged or plumose, embryo small with copious endosperm. Order: Poales Important Literature: Smith & Downs (1974: Fl. Neotropica, Pitcairnioideae); Smith & Downs (1977: dto., Tillandsioideae); Smith & Downs (1979: dto., Bromelioideae); Böhme (1988: nectary and ovary anatomy); Rauh (1990: synopsis cult. taxa); Baensch (1994: synopsis cult. taxa); Smith & Till (1998: family synopsis); Benzing (2000: ecology); Butcher (2000: updated key to genera); Crayn & al. (2004: CAM origin); Barfuss & al. (2005: phylogeny Tillandsioideae); Givnish & al. (2007: phylogeny); Horres & al. (2007: systematics Bromelioideae); Schulte & Zizka (2008: phylogeny Bromelioideae); Givnish & al. (2011: phylogeny & biogeography); Gouda & al. (2012+: digital encyclopedia); Silvestro & al. (2014: evolution Bromelioideae); Givnish & al. (2014: radiation & evolution); Aguilar-Rodríguez & al. (2014: synopsis bat pollination); Gitaí & al. (2014: cytology); Evans & al. (2015: phylogeny Bromelioideae); Roguenant & al. (2016: ill. synopsis); Palma-Silva & al. (2016: perspectives on evolution); Schütz & al. (2016: phylogeny Pitcairnioideae); Barfuss & al. (2016: phylogeny/classification Tillandsioideae);

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Males (2016: ecophysiology); Brown (2017: history of classification); Males & Griffiths (2017: functional types); Gouda & al. (cont.updated). Distribution: Tropical and subtropical America (Florida to C Argentina), one species (Pitcairnia feliciana, not succulent) in tropical W Africa. This large family — 58 genera and 3140 species (Givnish & al. 2014) to 75 genera and 3590 species (Gouda & Butcher 2016+: accessed 19 March 2019) — is well-known in cultivation, where its species are colloquially referred to as bromeliads. Numerous species are cultivated as house and foliage plants, and a multitude of hybrids and cultivars is generally available in the horticultural trade. Species of Tillandsia (“air plants”) are esp. popular among specialized collectors, and numerous hobby clubs and societies exist worldwide. Ananas (“pineapple”) is an important fruit crop, and several species have in the past been valued for their strong fibres (e.g. species of Aechmea, Bromelia, Neoglaziovia) (see Bennett (2000) for a complete list of known uses). Classification and History: The circumscription of the family is unambiguous, and all recent studies have confirmed its monophyly. Bromeliaceae are part of a basal grade within Poales, and its closest relationship is with the small South American family Rapataceae (16 genera, 90 species; no succulents). In traditional classifications (e.g. Smith & Downs (1974), Smith & Downs (1977) and Smith & Downs (1979)), three subfamilies (Bromelioideae, Pitcairnioideae, Tillandsioideae) were recognized. Recent molecular investigations do not support such a classification, and esp. the Pitcairnioideae in their traditional circumscription are widely paraphyletic (Givnish & al. 2007). Today, eight named clades are recognized at the rank of subfamily (Givnish & al. 2011) (Fig. 1). Palma-Silva & al. (2016) provide a summary of recent evolutionary and phylogenetic studies at different levels of the family, and also discuss biogeography and speciation in the light of current knowledge.

U. Eggli and E. J. Gouda

According to the analysis of Givnish & al. (2011), the origin of the family is on the Guayana Shield and can be dated to 100 mybp, while Bouchenack-Khelladi & al. (2014) place the origin of the family around 80 mybp. The modern lineages only started to differentiate much later beginning around 19 mybp, and the diversification intensified considerably after about 15.4 mybp, concurrent with major geological events in South America, esp. the rising of the Andes. Many genera have a comparatively recent origin, as witnessed by a calculated age of only 2 mybp for the speciose genus Dyckia. Diversification rates can be linked to the evolution of several characteristics (e.g. epiphytism, impounding tank rosettes, ornithophily) and are strongly coupled to life in fertile, moist, montane habitats (Givnish & al. 2014). Silvestro & al. (2014) showed that diversification rates in Bromelioideae are correlated both with the evolution of CAM photosynthesis (enhancing speciation rate) and the formation of tank rosettes (diminishing extinction rate). Terminology: Scharf & Gouda (2008) present a detailed discussion of the erroneous terminology frequently used in the past for describing features of the inflorescence, and esp. the different ways of branching. Inflorescences of bromeliads can be unbranched spikes (flowers sessile) or racemes (flowers pedicellate), or are branched and are then correctly termed panicles. In traditional literature, such as the Smith & Down volumes, the terms ‘pinnate’, ‘bipinnate’, ‘tripinnate’, etc. are incorrectly used to describe inflorescence branching, but these terms refer to compound leaves, rather than to different branching orders of branched inflorescences. In addition, the peduncle of the inflorescence (i.e. the basal, usually bract-bearing part of several internodes without flowers) is often incorrectly referred to as ‘scape’ (whose correct application describes the base of an inflorescence that consists of a single internode and is thus without bracts, as e.g. in many Amaryllidaceae or Hyacinthaceae). In addition to the problematical terminology used for inflorescence descriptions, that used for the parts of the perianth is also to some degree in dispute: Bromeliads, like other monocotyledons,

Bromeliaceae BL

837 Tillandsioideae

H N

Pitcairnioideae

Puy.

Bromelioideae

400 species

Fig. 1 Summary phylogeny of Bromeliaceae (3140 species total, data from Givnish & al. (2014) and Stevens (2001+), accessed Aug 2016). B, Brocchinioideae; H,

Hechtioideae; L, Lindmanioideae; N, Navioideae; Puy, Puyoideae. (Copyright: U. Eggli)

have a perigone, and its elements would, according to a strict terminology, have to be referred to as tepals. Because frequently only the inner whorl is petaloid, the use of the terms sepals and petals has become established by tradition throughout bromeliad literature, and this is also the usage followed here. A further complication is caused by the position of the ovary, which is inferior in Bromelioideae but partly half-inferior in the traditional Pitcairnioideae and Tillandsioideae (Böhme 1988, Bernardello & al. 1991), or superior in Dyckia (Fagundes & Mariath 2010, Santos-Silva & al. 2015). In the case of Tillandsioideae, the ovary appears to be superior macroscopically, but both Böhme (1988) and Bernardello & al. (1991) showed that the ovary is partially sunken into a receptacle. Accordingly, the often used term ‘pedicel’ cannot be used in descriptions of Tillandsia.

Brocchinia growing on nutrient-poor soils, have for long been suspected to be partly carnivorous (Benzing 2000: 217–223). Benzing (2000: 112) refined earlier concepts first formulated by Pittendrigh (1948), and differentiates between five different ecophysiological types, based on the presence/absence of tanks (= impounding rosettes = phytothelmata), the role of roots, the place of water and nutrient uptake, and photosynthetic pathways:

Ecology: Ecological specializations are numerous. Especially epiphytic taxa are often densely covered in greyish-white scales which absorb water from the atmosphere (“atmospheric epiphytes”). The large tank bromeliads from humid climates harbour an often unique microfauna (frogs, insects etc.) and -flora (specialized Utricularia species) in their water-filled rosettes. A limited number of bromeliads, such as

Type I: Terrestrial, without tanks, water and nutrient uptake via roots, with C3 or CAM Type II: Terrestrial, with weakly developed tanks, water and nutrient uptake via roots and absorptive trichomes on the leaf bases, with CAM Type III: Terrestrial (usually saxicolous) or epiphytic, with well-developed tanks, roots for mechanical support, water and nutrient uptake through absorptive scales on the leaf bases, mostly with CAM Type IV: Usually epiphytic, with well-developed tanks, roots for mechanical support, water and nutrient uptake through absorptive scales on the leaf bases, mostly with C3 Type V: Epiphytic or more rarely terrestrial and then mostly saxicolous, roots for mechanical support or absent, water and nutrient uptake through absorptive scales over the entire shoots

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and leaves, with CAM — this group conforms to the so-called “atmospheric bromeliads”

According to Males & Griffiths (2017) these types conform to different functional types, with close congruence of physiological parameters (such as degree of succulence, or photosynthetic pathway) with the climatic niches they inhabit. These authors also stress that different combinations of drought avoidance (by developing water storage tissues) and drought tolerance (by allowing lower tissue water potentials) are found in species of the family. Heteroblasty, i.e. an abrupt change in leaf shape during the development of young plants into adult plants, is known from several species (Meisner & Zotz 2012, Meisner & al. 2013), and is probably associated with the transition to the tank-forming habit of adult plants. Reyes-García & Griffiths (2009) stress that there are gradual switches from roots that function as absorptive organs to roots as anchor organs only, and that this grade is inversely parallel to an increasing presence of trichomes as absorptive structures. CAM photosynthesis is common in several clades (Horres & Zizka 1995), but Loeschen & al. (1993) did not find a correlation between the presence of CAM and the degree of succulence. Overall, about 45% of all bromeliads show some degree of CAM (Silvera & al. 2010) (see Matiz & al. (2013) and Crayn & al. (2015) for lists of taxa and references). CAM evolved at least five times independently from the supposedly ancestral C3 condition (Crayn & al. 2015) and is more predominant in terrestrial taxa (e.g. Deuterocohnia, Dyckia, Encholirium, where all species exhibit CAM). According to Quezada & Gianoli (2011) and Silvestro & al. (2014), the evolution of CAM is an evolutionary key innovation, and clades where CAM has evolved are generally more speciose than sister clades without CAM. Pierce & al. (2001) hypothesize that the peculiar foliar trichomes so common in the family were of hydrophobic nature ancestrally and served to repell water esp. from the abaxial leaf faces.

U. Eggli and E. J. Gouda

Benzing (2000) ascribes a general function of deterring predators and pathogens to the bromeliad trichomes. In Tillandsioideae, the scales have developed into complicated multicellular organs that serve in water uptake, and probably also for photoprotection (Pierce 2007), although the increase in light reflection is generally small (Pierce & al. 2001). The ontogeny of the absorptive trichomes has been studied in detail by Papini & al. (2010): The scales consist of a stem or axis that is connected to the internal leaf tissue (mesophyll parenchyma), and a shield that consists of several cells. The shield cells die when the trichome is fully developed and intercellular channels appear which connect the shield cells amongst them, and to the central stem. When plants with such trichomes become wet, a water film develops under the trichome shields, and is then passively absorbed by the shield cells. Succulence: Many bromeliads are slightly to distinctly leaf-succulent, and some have somewhat succulent stems. There is a transition from mesic to coriaceous and to distinctly succulent leaves, although even relatively mesic forms such as Catopsis floribunda or Vriesea incurvata or many Pitcairnia species have minimal amounts of water storage tissue (Benzing 2000: 65). In the case of the more succulent species, the degree of succulence is comparable to that of many undisputed leaf succulents from the Crassulaceae (Horres & Zizka 1995). The succulent tissue, which can make up to 85% of the total leaf cross-sectional area, is parenchymatous mesophyll, and is usually devoid of chlorophyll; epidermis and hypodermis are not contributing significantly to succulence (Metzler 1924, Loeschen & al. 1993, Horres & Zizka 1995). Some authors interprete the succulent tissues as representing a hypodermis (e.g. Pereira & al. (2011) for four species of Bromelioideae, Gomes-da-Silva & al. (2012) for species of the Vriesea corcovadensis group), but the hypodermis proper consists of one to five layers of sclerified cells (Monteiro & al. 2011, for Bromelia spp., although these authors also explicitely refer to a “water storage hypodermis” in the abstract). There is an abrupt change between the water-storage tissue and the chlorenchyma in most taxa. The architecture, however, can be variable even within a genus,

Bromeliaceae

and Monteiro & al. (2011) report an abrupt change for most species of Bromelia, but a gradual change in B. auriculata, B. morreniana, B. scarlatina and B. tubulosa. The water-storage tissue is usually predominantly located on the adaxial face of the leaves, but conditions vary and are of limited diagnostic value (e.g. both adaxial and abaxial in Dyckia and Encholirium, but adaxial only in most species of Pitcairnia, see Santos-Silva & al. (2013)), and can also be variable within a genus (e.g. abaxial in Bromelia morreniana, and present on both faces in B. tubulosa or B. scarlatina (Monteiro & al. 2011)). Horres & Zizka (1995: 53–55) tabulate the presence of water-storage tissue on both leaf faces for species of many additional genera. When the water-storage tissue is clearly demarcated from the chlorophyllous tissue, cells are often  isodiametric, but can also be narrow and elongate resembling a palisade parenchyma. Intercellular spaces are generally absent in the water storage tissue. The leaves of Guzmania lingulata consist of a rather uniform large-celled parenchymatic tissue, of which all cells except those of the uppermost one to two layers contain about equal and low amounts of chlorophyll (Vieira da Silva & al. 2011: Fig. 10). Under drought conditions, water is preferentially drawn from the water storage tissue (e.g. Freschi & al. (2010) for Guzmania monostachia), resulting in increasingly undulate cell walls concomitant with the loss of volume (e.g. Vieira da Silva & al. (2011: Fig. 15) for Guzmania lingulata). Pereira & al. (2011) describe reversible cellshrinkage on the base of the concertina-pattern observable in anticlinal walls (their Fig. 26, with scant evidence, however). Collapsible cells are also reported from species of Bromelia by Benzing (2000: 32, Fig. 2.13b), and for Alcantarea by Versieux & al. (2010), but these plants do not appear to be truly succulent, although the water-storage tissue can be quite well-developed in species from inselberg habitats (Versieux & al. 2010, Versieux & al. 2012). Succulence is very much a matter of degree in the family, and can be quite cryptic—even Tillandsia usneoides shows considerable succulence, although no specialized succulent tissue is

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present (Horres & Zizka 1995). Interestingly, the degree of succulence is, at least in some cases, not linked to environmental conditions, and Flores (1975) found the leaves of Aechmea mexicana (a moist-forest epiphyte) to have more pronounced water storage tissue than those of Hechtia glomerata (terrestrial in arid zones). Moreover, the extent of water storage tissues is under environmental control at least in some taxa, but contrary to expectations, water storage is more pronounced (but still insignificant) in shade plants of Billbergia elegans and Neoregelia mucugensis than in plants growing in open conditions (Rodrigues Pereira & al. 2013). Alcantarea glaziouana from the steep granite inselbergs in the Rio de Janeiro area shows a remarkable shift in life strategy because young plants have normal rosettes of very succulent leaves, and the large impounding tank rosettes are only present in adult plants, when leaf succulence disappears (Reinert & Meirelles 1993: as Vriesea geniculata, Versieux & al. 2010). Some species of Puya and Tillandsia form a conspicuous bulbous base. For Tillandsia, Rauh (1990) distinguishes “true bulbs” (e.g. T. argentea), with tightly packed leaf sheaths, from “pseudobulbs” (e.g. T. caput-medusae), where the leaf sheaths are thin and not succulent, and are separated from each other by a space of free air, which is often inhabited by ants. None of these two forms represents a monophyletic clade, however (Chew & al. 2010). Succulence has evolved independently in five of the eight currently recognized subfamilies. The succulent or predominantly succulent genera Hechtia (Hechtioideae), Deuterocohnia, Dyckia, Encholirium (Pitcairnioideae) and Puya (Puyoideae) show a remarkable degree of “concerted convergence” in six anatomical and physiological leaf characters that are all associated with an occurrence at (semi-) arid sites (Givnish & al. 2007). Apart from the genera with a full or partial treatment in this handbook, some degree of succulence is also present in at least some species of the genera shortly discussed in the list below. According to present knowledge, the water storage tissues are less well-developed in these

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cases, and the claim to succulence (according to the definition of Eggli & Nyffeler (2009)) remains to be tested: Ananas (Bromelioideae): A genus of three species (Butcher & Gouda 2014), of which A. ananassoides (“pineapple”, still usually known as A. comosus) is the most wellknown. A. ananassoides has a well defined water-storage tissue with collapsible cell walls under drought conditions, while the chlorenchyma tissue does not shrink (Sideris & Krauss 1955 cited from Benzing (2000)). A distinct water-storage tissue, making up 50% of the leaf cross-section, has been illustrated by Vieira da Silva & al. (2011: Fig. 1), showing a distinctly large-celled palisade-like hydrenchyma and a small-celled chlorenchyma. About 30% water-storage volume is reported for A. lucidus (Krahl & al. 2013). Brocchinia (Brocchinioideae): A genus of 20 species. Horres & Zizka (1995) report a water-storage tissue of 48% cross-sectional leaf area for B. uaipanensis (as Ayensua uaipanensis). The plants appear quite succulent (see Oliva-Esteve & Steyermark (1987: 307) for an illustration), but in view of its habitat (rocks on tepuis in an overall humid climate), the taxon does not appear to be truly succulent (E. Gouda, pers. obs.).

U. Eggli and E. J. Gouda

al. (2004) found that adaxial water-storage tissue is present throughout the genus and is generally more developed in semi-mesophytic species compared with the mesophytic species. Disteganthus (Bromelioideae): A genus of five species (Aguirre-Santoro & Michelangeli 2015). The leaves of some species are described as succulent, but they are merely  markedly coriaceous, similar to those of many species of Bromelia. Disteganthus species usually grow in at least partial shade and thus are not specially adapted to cope with water shortage. Guzmania (Tillandsioideae): A genus of 220 species. For G. lingulata, Vieira da Silva & al. (2011) illustrate a uniform tissue throughout the leaf with large cells with primarily waterstorage function, and few chloroplasts, but this species is mesophytic and has thin leaves (E. Gouda, pers. obs.). Freschi & al. (2010) found that there is a clear partitioning of functions (water storage vs. photosynthesis) within the leaves of G. monostachia from the base to the tip. Versieux & Medeiros (2018) found small amounts of water storage parenchyma both in adaxial as well as abaxial position for several allegedly mesophytic Guzmania species.

Bromelia (Bromelioideae): A genus of 70 species. The majority have distinctly coriaceous leaves and can withstand considerable aridity. Benzing (2000: 32, Fig. 2.13b) illustrates collapsible water-storage cells for B. balansae. For other species, minimal volumes of waterstorage tissues are reported (B. antiacantha: almost nil, Pereira & al. (2011); B. pinguin: 16%, Horres & Zizka (1995)).

Navia (Navioideae): A genus of 95 species. A number of species from arid sites (dry rocks, xerophytic savanna, dry cliffs) are described as having distinctly thickly coriaceous leaves by Robinson (1969) and Horres & Zizka (1995) report 67% water-storage cross-sectional leaf area for N. splendens. Illustrations (e.g. OlivaEsteve & Steyermark (1987: 332–355)) suggest that some might be truly succulent, but more research is necessary to evaluate whether this is really the case.

Catopsis (Tillandsioideae): A genus of 18 species. Water-storage tissue makes up 50% of the cross-sectional leaf area for C. delicatula, C. juncifolia and C. morreniana (Horres & Zizka 1995), but these plants are mesophytic with overall very thin leaves. Palací &

Pitcairnia (Pitcairnioideae): A genus of 408 species (incl. Pepinia). Horres & Zizka (1995) report a water-storage cross-sectional leaf area of 50% for P. andreana and P. burle-marxii. In addition, P. balsaminea, P. cristalinensis and P. encholirioides appear

Bromeliaceae

to be somewhat succulent (Santos-Silva & al. 2013), as well as the recently described P. volker-schaedlichii. Portea (Bromelioideae): A genus of 8 species. Horres & al. (2007) report fairly well-developed water-storage for P. leptantha. Quesnelia (Bromelioideae): A genus of 22 species. The water-storage tissue makes up 25 to >33% of the cross-sectional leaf area according to Mantovani & al. (2012). For Q. strobilispica, 40% water-storage volume is reported by Pereira & al. (2011). Vriesea (Tillandsioideae): A genus of 225 species. Horres & Zizka (1995) report a water-storage tissue making up 50% of the cross-sectional leaf area for V. espinosae (later transferred to Tillandsia), but the species appears to be at most xerophytic, rather than truly succulent (E. Gouda, pers. obs.).

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Bromeliads and malaria: Species with impounding rosettes are a malaria (and probably also dengue, zika, Mocellin & al. (2009)) vector, esp. in urban areas close to forests or rock outcrops that are inhabited by such species (e.g. the large-growing Alcantarea species in SE Brazil), and that are also breeding places for mosquitoes. This continuously encourages tree cutting and heavy herbicide use to eliminate the offending bromeliads in parts of Amazonian Brazil (Reitz 1983), but also in SE Brazil and Colombia. Bromeliads and their relationship with malaria were first intensely studied during World War II in Trinidad, where native species like Aechmea aquilega support Anopheles bellator that transmit malaria (Downs & Pittendrigh 1946, Pittendrigh 1948). Anopheles bellator is a small forest mosquito that breeds in water-filled bromeliad tanks. It has a disjunctive distribution and is a primary human malaria vector species (“bromeliad malaria”) in Trinidad and the coastal Atlantic forest of S Brazil (Gadelha 1994, Carvalho-Pinto

Key to subfamilies and genera with succulents. NB: A complete key to all genera of the family, updated from the keys in the three Smith & Downs-volumes, has been published by Butcher (2000) 1a

Fr indehiscent berries dispersed by animals (usually birds)

1b 2a

Fr usually capsular and dehiscent, or rarely indehiscent (Pitcairnia p.p.) but then not berries, Se dispersed by wind Se with a plumose appendage

2b 3a

Se winged, with a single hair-like appendage at both ends, or naked Fl unisexual and plants dioecious (only N to C America)

3b

Fl hermaphrodite or rarely unisexual and plants then monoecious or polygamodioecious, or dioecious and then from the Brazilian Shield Pet blades showy, tightly spiralled after anthesis, broad and distinct from the claw Pet blades showy or not, remaining free after anthesis or if slightly coiled then not clawed Pet large and conspicuous, or if minute then Sep imbricate and Anth basifixed, linear Pet minute and Sep cochlear (= abaxial sepal margins overlapping both axial ones), or Pet and Bra various and Sep twisted-convolute (= each sepal overlapping another with the margin at one side) or cochlear Sep twisted-convolute, Pet mostly many times longer than the sepals Sep cochlear and Pet minute L entire, star-like cells in the chlorenchyma abundant L margins toothed, star-like cells in the chlorenchyma absent

4a 4b 5a 5b

6a 6b 7a 7b

Bromelioideae (see separate key below) 2 Tillandsioideae (only Tillandsia p.p. with succulents) 3 Hechtioideae (only genus: Hechtia) 4 Puyoideae (only genus: Puya) 5 Pitcairnioideae (see separate key below) 6

Lindmanioideae (no succulents) 7 Brocchinioideae (no succulents) Navioideae (no succulents)

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Key to genera of subfamily Bromelioideae with succulents 1a 1b 2a 2b 3a 3b 4a 4b 5a 5b 6a 6b 7a 7b

Fl laxly arranged in simple or compound Inf, axes wholly visible Fl in dense spikes, racemes, or in dense compound and often prominently bracteate Inf Inf simple Inf compound Fl pedicellate; Sep with soft and usually obtuse apices Fl sessile Per slightly zygomorphic; Sep 15–35 mm; Pet 33–65 mm; pollen mostly sulcate (C America to Argentina and Uruguay) Per actinomorphic; Sep 6–9 mm; Pet 13–20 mm Ovules obtuse; pollen sulcate; L attenuate,  as thick as wide (NE Brazil) Ovules long-caudate; pollen porate; L flat and apiculate (C-E Brazil) Sep mucronate or pungent (N & S America) Sep unarmed Ovules long-caudate; Sep mostly 40 cm, sheaths broadly ovate, thick, dark, L lamina very narrowly triangular, becoming involute, with pungent tip, sharply carinate and appressedly palelepidote abaxially, entire (inner) or basally minutely spinose (outer), Sp brown, hooked; Inf to 50 cm, simple; peduncle 16 cm, slender, lepidote or glabrous; peduncular Bra suborbicular with long linear lamina, slightly shorter than the internodes, entire; fertile Inf part lax, 2- to 6-flowered, axis very slender,  glabrous; floral Bra spreading, broadly ovate with 10–13 mm long acuminate tips (lower), exceeding the flowers; Fl suberect, subsecund, 9 mm or longer, glabrous, subsessile; Sep subelliptic with a minute recurved mucro at the apex; Pet erect, lamina large, suborbicular, obtuse, colour not described; St included; Fil free above the common tube; Anth elliptic, acute, straight or slightly recurved; Sty very short; Fr and Se not described. Insufficiently known and not recently collected. D. burle-marxii L. B. Smith & Read (Phytologia 38: 137–138, t. 5, 1977). Type: Brazil, Bahia (Burle Marx s.n. [HB, US [photo]]). — Distr: Brazil (Bahia: Chapada Diamantina); ecology not recorded. [1?] Ros not described; L >30  1.5 cm, sheath not described, L lamina very narrowly triangular, between the veins finely pale-lepidote abaxially, soon glabrous and lustrous adaxially except on the spines, laxly spinose-serrate, Sp curved, 9 mm; Inf 1.7 m or more, compound; peduncle glabrous at least with age; peduncular Bra ovate, lamina narrowly triangular (lower) to attenuate (upper), exceeding the internodes but not imbricate, serrulate (lower) or entire (upper); fertile Inf part finely cinereous-tomentose; Br suberect, slender, 30 cm; primary Bra lanceolate-ovate, attenuate, to 5 cm, serrulate or entire; floral Bra broadly ovate, acuminate, nearly equalling the sepals; Ped 3 mm, obconical; Sep broadly subelliptic, 6 mm, broadly rounded at the apex, ecarinate, cucullate at anthesis; Pet lamina spreading, elliptic, 8 mm, broadly rounded, ecarinate, orange; St  equalling the petals; Fil high-connate above the common tube; Ov broadly ovoid; Sty very short.

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Probably related to D. encholirioides. — [F. Krapp] D. cabrerae L. B. Smith & Reitz (Sellowia 14: 101, fig. 2, 1962). Type: Brazil, Santa Catarina (Smith & al. 9262 [US, HBR]). — Distr: Brazil (Santa Catarina); dry open ground and rocks. I: Smith & Downs (1974: 523). [1/2?] Ros dense, many-leaved; L 31–40  1.5 cm, rigid, slightly curved, sheath ovate, 3  2–4 cm, glabrous and sublustrous adaxially, densely cinereous-lepidote towards the apex abaxially, L lamina linear, canaliculate, acute, abruptly acute and mucronulate, appressedly cinereous-lepidote, soon glabrous adaxially, laxly serrate, Sp slender, retrorsely curved, 3 mm, brown; Inf to 1.2 m, branched; peduncle  flattened, 70 cm, glabrous; peduncular Bra broadly ovate, acuminate or the lowest leaf-like, the uppermost remote, cinereous-lepidote towards the apex, glabrous elsewhere; fertile Inf part with 1–2 longstipitate racemes at the base, lax, 38–54 cm, diffusely cinereous-lepidote, soon glabrous; Br few-flowered; primary Bra like the upper peduncular bracts, to 2 cm, much exceeded by the sterile branch bases; floral Bra acuminate from a broadly ovate base, ascending, reaching  the middle of the sepals, minutely serrulate; Fl suberect, to 17 mm, orange; Ped 5 mm, slender; Sep broadly ovate, 8–13 mm, acute, strongly incurved, subeven, the posterior ones strongly carinate; Pet distinctly unguiculate, curved-spreading towards the apex, lamina rhombic, obtuse; St slightly shorter than the petals; Fil short-connate above the common tube; Anth narrowly triangular, 3 mm, recurved towards the apex; Ov ovoid; Sty short; Fr stoutovoid, acute, 18 mm, dark castaneous, lustrous; Se subovate, 4 mm, strongly complanate, wing asymmetrical and circumferential. D. cangaphila P. J. Braun & al. (Bromelie 2010(3): 107, ills. (pp. 105–110), 2010). Type: Brazil, Goiás (Esteves Pereira 392 [UFG]). — Distr: Brazil (SW Goiás); iron-rich gravel deposits (“pedra canga”) on and between sandstone. I: Esteves Pereira & Gouda (2017: 180). [2/3a] Ros with up to 70 living leaves, to 35 cm ∅, with a stem to 11  4.7 cm; L to 19  2.6 cm,

F. Krapp and U. Eggli

sheaths broadly ovate, to 1.8  2.8 cm, whitish, glabrous adaxially, L lamina arching, very rigid, concave, triangular-lanceolate, brownish-red to purple, sometimes green basally, veined, adaxially glossy and sparsely lepidote basally, abaxially densely light grey-lepidote, serrate, Sp strong and claw-like, pungent, brownish; Inf to 1.3 m, simple or with up to 3 branches; peduncle  erect, 85–107 cm, pale green to pale brownish, densely grey-lepidote; peduncular Bra leaf-like, triangular-lanceolate, to 65  5.5 mm (uppermost only to 17 mm), green to brownish-green (lower) or dull pale green (upper), later stramineous; fertile Inf part to 41 cm, 37- to 52-flowered, basally lax, above much denser, rachis initially pale green, towards the apex changing to greenishorange or orange, grey- or orange-lepidote; Br slender, 15 cm; floral Bra 19 mm, sometimes equalling the petals, smaller higher up, brownishgreen, soon stramineous, strongly veined; Fl slightly ascending, 16  8 mm; Ped absent in bud, to 7 mm at anthesis; Sep 7.7  6.5 mm, orange-yellowish, creamish-yellow-lepidote, base succulent; Pet obtuse, 11  7.6 mm, emarginate (V-like), orange-yellowish; St included at anthesis; Fil 7.6 mm, apical 1.4 mm free, yellow; Anth yellow, 3.5 mm, apex curved; Ov narrowly pyramidal, 5.5  2.8 mm, yellow to light yellow; Sty 4 mm; Sti lobes linear, yellow; Fr conically ovoid, to 15  12 mm, chestnut-brown to blackish-wine-coloured. Closely related to D. pumila. D. choristaminea Mez (Repert. Spec. Nov. Regni Veg. 16: 71, 1919). Type: Brazil, Rio Grande do Sul (Malme s.n. [B]). — Distr: S Brazil (Rio Grande do Sul); open rocky places. I: Smith & Downs (1974: 528). – Fig. 2. [3a] Ros solitary or with some congested offsets, to 15 cm ∅; L 7–12  0.5 cm, sheath suborbicular, 2 cm, pale, glabrous, L lamina linear, deeply canaliculate, both faces cinereouslepidote, margins laxly serrate, Sp slender, curved, 2.5 mm; Inf 15–25 cm, simple; peduncle slender, glabrous; peduncular Bra densely imbricate, broadly ovate, abruptly acute, somewhat inflated, minutely lepidote; fertile Inf part few-flowered, 3–5 cm, rachis densely white-

Dyckia BROMELIACEAE

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Fig. 2 Dyckia choristaminea (Kirschnek s.n.: Brazil; Rio Grande do Sul, sine loco). (Copyright: U. Eggli)

villous, subterete; floral Bra like the peduncular bracts, carinate towards the apex, 15 mm, equalling the sepals, entire; Ped short, stout; Fl 18–24 mm,  funnel-shaped; Sep very broadly ovate, acute, 10–11 mm; Pet lobes spreading to recurved at anthesis, yellow; St included to slightly exserted; Fil free above the very short common tube; Ov 8 mm; Sty very stout, 8 mm. D. cinerea Mez (in Martius, Fl. Bras. 3(3): 469, 1894). Type: Brazil, sine loco (Glaziou 18570 [B, K, F [photo]]). — Distr: Brazil (Minas Gerais); high-altitude grasslands, rocky fields, on rocks. [2] Ros acaulescent; L very numerous, 50  >3 cm, very rigid, sheath not described, L lamina narrowly triangular, densely cinereouslepidote, laxly serrate, Sp stout, curved, 2.5 mm; Inf 1 m or more, compound or pseudo-racemose with reduced branches or buds only in the axils of the lower bracts; peduncle stout, cinereoustomentulose or glabrescent; peduncular Bra broadly ovate with narrowly triangular lamina, exceeding the internodes, serrulate; fertile Inf part cinereous-tomentulose; floral Bra broadly ovate, apiculate, reflexed (upper), equalling the flowers or shorter (lower), serrulate; Ped short, stout; Fl erect to spreading, 17–18 mm, orange; Sep 11 mm, abruptly obtuse, cucullate, obtusely carinate towards the apex, subglabrous, minutely fimbriate; Pet erect, elliptical-oblong, 16 mm,

subacute or obtuse, orange; St much shorter than the petals; Fil 2 mm connate above the common tube; Anth sublinear, stout,  recurved; Sty  none. D. commixta Hassler (Annuaire Conserv. Jard. Bot. Genève 20: 305, 1919). Type: Paraguay, Cordillera (Chodat & Vischer 97 p.p. [G, F [photo]]). — Distr: Brazil (Paraná), Paraguay; ecology not reported. [3a] Known only from fragments; inner L 6  0.5 cm, sheaths triangular-elliptic, 3  1.5 cm, membranous, glabrous adaxially, striate abaxially with whitish scales between the veins, margins entire and hyaline, L lamina linear-triangular, attenuate, pungent,  glabrate adaxially, like the sheaths abaxially, serrate, Sp antroretrorsely curved or straight, 1–1.5 mm; Inf size unknown, pseudosimple; peduncle flexuous, long, glabrous; peduncular Bra broadly ovate, acuminate (lower), much shorter than the internodes, 6–20 mm (lowest), striately immersed-lepidote (lowest), with hyaline margin and  finely fimbriate (uppermost); fertile Inf part with undeveloped buds in the axils of the lowest bracts, laxly flowered, rachis flexuous, subquadrangular, glabrous; floral Bra erect to spreading, broadly ovate, acute and mucronulate, convex, 5–6 mm,  as long as the sepals, glabrous; Fl 14–15 mm; Ped 1–2 mm, rather stout; Sep ovate-elliptic, 10  7 mm, subacute, thin, with hyaline margin, glabrous,

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rugulose when dry; Pet erect, ovate-elliptic, 14 mm, ecarinate, inflexed at the apex, colour not described; St included; Fil free above the 2 mm high common tube; Anth linear-triangular, sagittate at the base, 6 mm; Ov trigonous, 10 mm; Sty 1 mm. D. conceicionensis O. B. C. Ribeiro & Leme (J. Bromeliad Soc. 65(1): 15–19, ills., 2015). Type: Brazil, Minas Gerais (Ribeiro 267 [BHCB]). — Distr: Brazil (Minas Gerais: Conceição do Mato Dentro); on iron-rich rock (“pedra canga”) outcrops, 1000 m. [3a] Ros 20 cm ∅; L 20, erect to spreading, straight to arcuate, coriacous, sheath 1.7–1.9  2–2.2 cm, whitish, glabrous and glossy on both faces, L lamina narrowly triangular, 9–10  1.5 cm, canaliculate, dark red to bronze-coloured towards the tip, adaxially glabrous and glossy, abaxially distinctly veined and between the veins rather densely white-lepidote, tip acuminately spine-tipped, pungent, margins laxly spinose, Sp retrorse-uncinate, 2.5–3 mm, subtriangular, 15 mm apart; Inf 50 cm, simple, erect; peduncle 30–35 cm, glabrescent, reddish-green; peduncular Bra ovate-acuminate, 9–11  5–6 mm, much shorter than the internodes, greenish near the base to ochraceous near the tip, veined and with prominent midvein near the tip, sparsely white-lepidote, margins minutely denticulate to subentire; fertile Inf part 18–20 cm, laxly 6- to 20-flowered, rachis reddish, sparsely lepidote; floral Bra broadly ovate, 9–10  5–6 mm, acute, remotely denticulate, somewhat spreading; Fl 14–15 mm, porrectspreading, tubular, sessile; Sep broadly ovate, 6–7  5–6 mm, orange-red, obtuse-emarginate, convex, sparsely white-lepidote; Pet broadly spatulate, 10–11  9 mm, erect, orange, ecarinate, broadly obtuse-emarginate; St included; Fil yellow, free above the common tube; Anth suboblong-lanceolate, straight, base bilobed; Ov 6 mm, suboblong, yellow; Sty 1 mm, distinct, orange; Sti conduplicate-spiral; Fr and Se unknown. A member of the D. saxatilis complex, and compared with the similar D. brachyphylla in the protologue. — [U. Eggli]

F. Krapp and U. Eggli

D. consimilis Mez (in Martius, Fl. Bras. 3(3): 479, t. 90, 1894). Type: Brazil, Minas Gerais (Weddell 1407 [P, GH [photo]]). — Distr: Brazil (Minas Gerais); rocky fields, on rocks. I: Smith & Downs (1974: 575). [3b] Ros acaulescent, with a basal bulbous part and a short thick rhizome; L numerous, densely arranged, 5  0.7 cm, sheath broad and conspicuous, depressedly semicircular, glabrous and glossy, abaxially brownish, L lamina narrowly triangular, glabrous adaxially, appressedly ferrugineous- or canescent-lepidote abaxially, laxly serrate, Sp stout, 3 mm; Inf 10  1.5 cm, sheath suborbicular, 1.5 cm, brown, L lamina narrowly triangular, pungent, appressedly cinereous-lepidote on both faces, margins very laxly serrate, Sp flat, mostly retrorse, 1 mm; Inf 30 cm, simple; peduncle erect, very sparsely and minutely white-lepidote; peduncular Bra ovate, acuminate, pungent, small, remote; fertile Inf part laxly 7-flowered, rachis slender, 4 cm, very sparsely and minutely lepidote; floral Bra broadly ovate, acuminate, the lower  equalling the sepals; Fl spreading; Ped 1 mm, obconical;

Dyckia BROMELIACEAE

Sep triangular-ovate, 5 mm, obtuse, the posterior carinate; Pet suborbicular, 10 mm, slightly carinate, orange; St slightly exserted; Fil free above the short common tube; Sty very short. Similar to D. pumila. The recently described D. secundifolia is also compared with D. coximensis in its protologue. D. crassifolia Rauh (Trop. subtrop. Pfl.-welt 79: 10–11, fig. 4, 1991). Type: Bolivia, Santa Cruz (Rauh 40670 [HEID]). — Distr: Bolivia (Santa Cruz); dry forest, 1200 m. [3a] Ros few-leaved, to 16 cm ∅, 8 cm high, acaulescent; L 10–16  1.2 cm, strongly succulent, sheath 1.5  1.3 cm, white, glabrous adaxially, lepidote abaxially, L lamina narrowly triangular, with a long pungent tip, pale green to reddish adaxially, densely white-lepidote abaxially, margins spinose, Sp strong, retrorse, to 5 mm, dark brown, lepidote, 10 mm apart; Inf to 45 cm, simple; peduncle slender, 30–40 cm, green to reddish, sparsely white-lepidote; peduncular Bra oblong-ovate, acuminate, 5 mm, shorter than the internodes, veined, indistinctly serrulate; fertile Inf part lax, 10 cm, 10-flowered, rachis green to reddish, sparsely white-lepidote, slender; floral Bra broadly ovate, short-acuminate, very small, 2 mm, brownish, membranous, appressed; Fl spreading, 17 mm, narrowly tubular and narrowed towards the throat; Ped very short, stout; Sep oblong, 7 mm, obtuse, succulent, slightly carinate, orange-red, darker at the tip, sparsely whitelepidote, margins membranous, indistinctly serrulate; Pet 15 mm, obtuse, erect, orange-red, margins smooth; St slightly exserted; Fil free above the very short common tube; Anth curved outwards; Sty much shorter than the conspicuous Sti branches; Fr and Se not described. Similar to D. leptostachya. Somtimes treated as synonym of D. pulquinensis. — [F. Krapp] D. crocea L. B. Smith (Phytologia 19: 282, t. 1, figs. 7 and 8, 1970). Type: Brazil, Paraná (Hatschbach 17391 [US]). — Distr: Brazil (Paraná); open fields; I: Smith & Downs (1974: 575). [3b] Ros many-leaved; L to 25  0.8–1.3 cm, sheath suborbicular, 3 cm, yellow, glabrous and

905

lustrous adaxially, L lamina very narrowly triangular, soon glabrous adaxially, appressedly palelepidote abaxially, laxly serrate, Sp recurved or spreading, 1–2.5 mm; Inf 60–120 cm, usually simple; peduncle straight or nearly so, glabrous; peduncular Bra ovate, acuminate, shorter than most of the internodes, entire or inconspicuously serrulate; fertile Inf part laxly many-flowered, 17–32 cm, soon glabrous; floral Bra spreading, ovate, acuminate, shorter than the sepals, entire; Sep ovate, acute,  carinate, 8–9 mm; Pet lamina elliptic, cucullate, 15 mm, orange; St included; Fil connate above the common tube; Sty  none. Similar to D. aurea. — [F. Krapp] D. dawsonii L. B. Smith (Contr. Sci. Nat. Hist. Mus. Los Angeles County 17: 2, figs. 2–4, 1957). Type: Brazil, Goiás (Dawson 15236 [R, LAM, US]). — Distr: Brazil (Goiás); rock outcrops along a canyon, Cerrado vegetation. – Fig. 3. [3b] Ros acaulescent, 30–40 cm ∅, slowly offsetting and forming a dense clump; L to 22  0.7 cm, mostly spreading, sheath suborbicular, >2 cm wide, stramineous, lustrous, glabrous, L lamina linear, appressedly cinereouslepidote on both faces, laxly serrate, Sp slender, curved, 5–7 mm, mostly subopposite; Inf 70 cm, simple; peduncle glabrous; peduncular Bra broadly ovate with a linear lamina, thin, all but the lowest much shorter than the internodes, entire; fertile Inf part lax, 25–30 cm, glabrous; floral Bra suborbicular, apiculate, thin, 4 mm, erose; Ped 3 mm, stoutly obconical; Sep broadly elliptic, obtuse, thin, ecarinate, 5–6 mm; Pet ecarinate, lamina broadly obovate, 9 mm, yellow; St exserted; Fil 2 mm connate above the 1 mm high common tube; Sty  none. The protologue description is incomplete; rosette characters have been added from offspring of the type collection cultivated at the Sukkulenten-Sammlung Zürich. The species has recently used for hybridization, with Dyckia ‘Brittle Star’ (likely = D. fosteriana  D. platyphylla) as second parent, yielding very attractive plants (Saul 2018). D. delicata Larocca & Sobral (Novon 12(2): 234–236, ills., 2002). Type: Brazil, Rio Grande

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F. Krapp and U. Eggli

Fig. 3 Dyckia dawsonii (Dawson 15236: Brazil; Goiás, E of Formoso, type collection.). (Copyright: U. Eggli)

do Sul (Larocca & al. 96/001 [ICN, MBM, US, ZSS]). — Lit: Sachs (2011: with ills.). Distr: Brazil (Rio Grande do Sul); basalt outcrops. [1?] Ros 20–60 cm ∅; L lamina 5–20  0.7–2 cm, succulent, eventually recurved in solitary individuals, cinereous to reddish-cinereous, densely pale-lepidote on both faces, sheath broadly triangular, L lamina linear-triangular, apex forming a rigid spine, margins undulate, Sp soft, straight, antroretrorse, flexible, 5–7 mm, 3–5 mm apart near the leaf tip; Inf 120 cm, simple or with up to 9 branches to 35 cm long; peduncular Bra linear-lanceolate, papyraceous, 30–55  6–16 mm, exceeding the internodes (lower) or shorter (upper), with 10–20 parallel veins, glabrous, with teeth to 3 mm (upper); fertile Inf part 80- to 150-flowered, erect, rachis softly ridged, glabrous; floral Bra elliptic-ovate, rounded or acute, 5.5–8  5–6 mm, glabrous; Sep ovate-oblong, rounded, 6–7  3.5–4 mm, green, margins hyaline for 1 mm; Pet lanceolatespatulate to oblanceolate, 10–13  3–4 mm, yellow; St 9–10 mm; Fil somewhat unequally fused with the petals, free above the common tube; Anth lanceolate-ovate, 2  1 mm; Ov triquetrous, oblong-acute, 7  2.5–3 mm; Sty 7–10 mm, Sti lobes papillose; Fr ellipsoid, 9–12  3–5 mm; Se triangular, with a reduced wing along the longitudinal axis, 1.2–1.7  0.5–1 mm.

Similar to D. hebdingii. — [F. Krapp] D. deltoidea (L. B. Smith) L. B. Smith (Phytologia 14: 485, 1967). Type: Brazil, Paraná (Dusén 17357 [GH, S]). — Distr: Brazil (Paraná); campo, 740 m.  Dyckia coccinea var. deltoidea L. B. Smith (1943)  Dyckia tuberosa var. deltoidea (L. B. Smith) L. B. Smith (1950). [3a] Ros not described; L >40  1.4 cm, sheath suborbicular, 4 cm, brown, L lamina linear, attenuate to a finely subulate rigid apex, flat, glabrous adaxially, closely appressedly brownish-lepidote abaxially, very laxly serrate, Sp slender, curved, 2 mm; Inf to 80 cm, simple; peduncle soon glabrous; peduncular Bra erect, subfoliaceous and imbricate (lowest) to ovate and acuminate, shorter than the internodes, densely lepidote abaxially; fertile Inf part lax, 23–28 cm, rachis soon glabrous; floral Bra narrowly triangular, to 22 mm, exceeding the lowest flowers, densely appressedlepidote abaxially, minutely serrulate; Fl spreading; Ped 5 mm, subcylindrical, stout; Sep broadly ovate or elliptic, 10 mm, acuminate, mucronate, smooth except for the strong midvein, slightly if at all carinate, minutely serrulate, white-lepidote, glabrescent; Pet lamina suberect to spreading, subrhombic, 17 mm, carinate-complicate, orange-yellow; St included; Fil free above the common tube; Sty  none. — [F. Krapp]

Dyckia BROMELIACEAE

D. densiflora Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): 1194, 1830). Type: Brazil, Minas Gerais (Martius s.n. [M, US [photo]]). — Distr: Brazil (Minas Gerais: Diamantina region: Morro de Vila Rica); rocky slopes, 1070–1200 m; only known from the region of the type locality. I: Smith & Downs (1974: 535). [3b] Ros not described; L to 5  0.7 cm, sheath 2 cm wide, dark brown, glabrous, L lamina narrowly triangular, soon glabrous adaxially, cinereous-lepidote abaxially, laxly serrate, Sp 2 mm; Inf to 40 cm, simple; peduncle densely ferrugineous-lepidote; peduncular Bra ovate, acuminate, equalling the internodes, densely serrulate; fertile Inf part subdensely few-flowered, densely tomentose-lepidote; floral Bra spreading or reflexed, ovate, acuminate, equalling the sepals (lower) or shorter (upper), serrulate; Fl spreading at anthesis and then erect, 10 mm; Ped 2 mm; Sep ovate, 8 mm, apiculate, fimbriate-serrulate; Pet lamina suberect, broadly elliptic, obtusely undulate, barely carinate, orange; St slightly shorter than the petals; Fil high-connate (to completely connate) above the common tube; Anth stout, subacute,  recurved; Sty very short. — [F. Krapp] D. dissitiflora Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): 1194, 1830). Type: Brazil, Bahia (Martius s.n. [M, F [photo]]). — Distr: Brazil (Bahia, Minas Gerais, Piauí); on rocks in dense Caatinga vegetation or savanna. [3b] Ros not described; L 20  1 cm, sheath not described, L lamina narrowly triangular, densely appressed-lepidote abaxially, serrate, Sp slightly >1 mm; Inf 50–100 cm, simple; peduncle glabrous; peduncular Bra ovate with long-attenuate pungent lamina, much shorter than the internodes, entire; fertile Inf part lax, elongate, lepidote; floral Bra spreading, lanceolate and acuminate (lower) or broadly ovate and apiculate (upper), equalling or exceeding the sepals (lower) or much shorter (upper); Fl 12–13 mm; Ped 3–4 mm; Sep elliptic, broadly acute and apiculate, straight or nearly so, 7 (–9) mm, furfuraceous esp. towards the base; Pet suberect, lamina suborbicular, carinate, when dry brownish-yellow; St included; Fil completely

907

connate above the common tube; Anth narrowly triangular, acute, nearly or quite straight; Sty as long as the ovary. — [F. Krapp] D. distachya Hassler (Annuaire Conserv. Jard. Bot. Genève 20: 308, 1919). Type: Paraguay (Fiebrig 5648 [G, F [photo]]). — Lit: Pompelli & Guerra (2004: conservation); Wiesbauer (2008: biology, conservation); Voltolini & al. (2009: leaf anatomy). Distr: Brazil (Santa Catarina), Paraguay; ledges by streams and rock islands, seasonally inundated. I: Smith & Downs (1974: 542); Reitz (1983: t. 45).  Dyckia interrupta Mez (1919) (nom. illeg., Art. 52.1); incl. Dyckia distachya fa. induta Hassler (1919). [2] Ros acaulescent, 50-leaved; L 14–20  2–3.5 cm, thick, rigid, sheath suborbicular, L lamina narrowly triangular, pungent, glabrous adaxially, minutely white-lepidote between the prominent veins abaxially, very laxly serrate, Sp minute, curved; Inf to 1.3 m, simple or few-branched; peduncle soon glabrous; lowest peduncular Bra leaf-like, all others much smaller, to 10 mm, shorter than the internodes, entire; fertile Inf part to 50 cm, white-furfuraceous, Br laxly flowered, divergent; primary Bra shorter than the sterile branch base; floral Bra broadly ovate, apiculate, shorter than or  equalling the sepals (lowest), entire; Ped 1–1.5 mm; Sep elliptic, 6–8 mm, obtuse, ecarinate; Pet suborbicular, 12–15 mm, orange or yellow; St included; Fil free above the common tube; Sty  equalling the ovary; Fr trigonous, dark brown, glossy, 17  8 mm; Se 3 mm. The disjunct distribution is notable. Reitz (1983) compares the species with D. brevifolia. The species appears severely threatened by dam construction, and micropropagation possibilities have been developed (Pompelli & Guerra 2004, Pompelli & Guerra 2005). D. divaricata Leme & H. Büneker (J. Bromeliad Soc. 65(2): 129–134, ills., 2015). Type: Brazil, Mato Grosso do Sul (Kranz & al. 426 [RB, HB]). — Distr: Brazil (Mato Grosso do Sul: region of Antônio João); on horizontal rock outcrop with disturbed semi-deciduous tropical forest, 480 m.

908

[1] Ros offsetting and forming large colonies; L 20–25, densely rosulate, coriaceous-succulent, suberect to arching, sheath suborbicular, 5  6 cm, densely white-lepidote at the apex, glabrous below, at least near the apex castaneous and glossy, L lamina narrowly triangular, attenuate, 40–45  2–2.3 cm, vividly green, shallowly canaliculate towards the tip, adaxially densely white-lepidote towards the base but glabrous above, adaxially subdensely white-lepidote along the veins, tip a pungent spine, margins spinose, Sp narrowly triangular, 3–6 mm, antrorse-uncinate, 5–20 mm apart; Inf 1.7 m, ascending-spreading, broadly paniculate; peduncle to 75 cm, erect, glabrescent, densely covered with white wax, pale greenish-bronze-coloured; peduncular Bra narrowly triangular, 60–80  15–20 mm, acuminate, spinescent, subdensely white-lepidote; Br 11, spreading, 35–47 cm, densely 50- to 85-flowered, rachis straight, densely whitefloccose; primary Bra similar to the upper peduncular bracts, 40–45  10 mm, as long as or shorter than the sterile branch base; floral Bra triangularovate to broadly ovate, 3–10  2–4 mm, stramineous, densely white-floccose, irregularly dentate to entire; Fl 15–17 mm, 16 mm ∅, subverticillately arranged, broadly campanulate to substellate, spreading; Ped 3 mm, stout but inconspicuous, densely white-floccose; Sep Fig. 4 Dyckia domfelicianensis (Sachs s.n.: Brazil; Rio Grande do Sul; Arroio dos Ratos). (Copyright: U. Eggli)

F. Krapp and U. Eggli

broadly elliptic to orbicular, 5–6  4.5–5 mm, green with yellowish apex, strongly convex, densely white-floccose; Pet elliptic to obovate, spreading, obtuse-emarginate, 11–12  7–7.5 mm, yellow, ecarinate, basally connate for 2 mm, glabrous; St  as long as the petals, exposed, erect; Fil free above the common tube, yellow; Anth sublinear, 4 mm, straight, base deeply bilobed; Ov slenderly ovoid, 6  2.5 mm, yellow; Sty 4 mm, yellow; Sti conduplicate-spiral, yellow, 2 mm; Fr subglobose, dark brown, glossy, size not described; Se unknown. Compared with D. walteriana, D. microcalyx and D. exserta in the protologue. The species occurs within 10 km of the border with Paraguay and probably occurs in that country as well. — [U. Eggli] D. domfelicianensis Strehl (Vidalia 2(2): 30–32, figs. 5–8, 2004). Type: Brazil, Rio Grande do Sul (Strehl & Silva 1283 [HAS]). — Distr: S Brazil (Rio Grande do Sul); rocky outcrops between grasses. – Fig. 4. [1] Ros very many-leaved (100–300 leaves), 50 cm ∅, with a short stem; L sheath 3.5  5.5 cm, dark brown, glossy on both faces, L lamina 23–25  3.5 cm, tough and succulent, acuminate, acute, clear green, cinereously whitelepidote, with pungent tip, margin spinose, Sp hooked, antrorse, brown; Inf to 120 cm, 1–2

Dyckia BROMELIACEAE

branched, paniculate; peduncle 50 cm, green to brownish, with fine white tomentum; peduncular Bra leaf-like (lowest), long-triangular, acuminate, to 50 mm, sparsely serrate, laxly arranged; fertile Inf part 60–80 cm, lax, white-lepidote; Br to 45 cm, rachis slightly angular, white-lepidote; floral Bra broadly triangular, acute, 2  3 mm, green to brown, abaxially puberulous; Fl sessile, narrowly tubular; Sep ovate-acuminate, 5–6  3 mm, carinate, connate for of their length, abaxially whitelepidote, tip  bifid; Pet ovate, 9  3–4 mm, clear yellow, abaxially white-lepidote; St exserted; Fil free, 6 mm; Ov green, 3–4 mm; Sty 6 mm, yellow; Fr and Se not described. Similar to D. hebdingii. D. duckei L. B. Smith (Bol. Mus. Paraense “Emilio Goeldi”, n.s., Bot. 1: 3, fig. k–o, 1958). Type: Brazil, Pará (Ducke s.n. [US, MG 14856]). — Distr: Brazil (Maranhão, Pará); iron-rich rocky (“pedra canga”) fields. I: Smith & Downs (1974: 558). [3a] Ros acaulescent, very dense, very manyleaved; L to 26–40  1.5–2 cm, arching-recurved, sheath broadly elliptic, 2–3 cm, L lamina very narrowly triangular, with pungent tip, soon glabrous adaxially, appressedly cinereous-lepidote abaxially, margins laxly serrate, Sp slender, spreading, 2 mm; Inf 40–130 cm, simple or subsimple; peduncle very slender, glabrous; peduncular Bra ovate, acuminate, very small, entire (except the lowest), remote; fertile Inf part lax, 5–26 cm, soon glabrous; floral Bra broadly ovate, apiculate, shorter than the pedicels, entire; Fl divergent or spreading; Ped to 3 mm, slender; Sep elliptic, 5.5 mm, obtuse, ecarinate, erose; Pet lamina rhomboid, 10 mm, carinate, orange; St slightly but distinctly exserted; Fil free above the 1 mm high common tube, 10 mm, orange; Sty 1 mm; Fr subglobose, 10–12 mm, dark castaneous. Monteiro & Forzza (2016, 1260) describe and illustrate material under this name with rosettes with few erect to spreading leaves only 10–13 cm long, and much smaller inflorescences, but with similar flowers. D. dusenii L. B. Smith (Contr. Gray Herb. 98: 6, t. 2, 1932). Type: Brazil, Paraná (Dusén 18081

909

[S, GH [photo]]). — Distr: Brazil (Paraná, Santa Catarina, Rio Grande do Sul); cliffs and dry fields. I: Smith & Downs (1974: 539). [3b] Plants stout; L 30–40  1.5–2 cm, sheath broad, suborbicular, dark castaneous, L lamina narrowly triangular, attenuate, yellow-green, glabrous adaxially, densely pale-lepidote abaxially, laxly serrate, Sp curved, 2–3 mm; Inf to 70 cm, simple; peduncle stout, soon glabrous; peduncular Bra broadly ovate, acuminate, large, much exceeding the internodes, densely pale-lepidote, entire or minutely denticulate; fertile Inf part subdense, many-flowered, 20–30 cm; floral Bra broadly ovate, acuminate, exceeding the sepals (all) and the petals (lowest); Fl subspreading, soon glabrous; Ped 5 mm, stout; Sep ovate, 16 mm, acute, strongly alate-carinate, frequently denticulate; Pet lamina broadly elliptic, 15–20 mm, obtuse, ecarinate, suberect, yellow; St much shorter than the petals; Fil shortly connate above the common tube; Anth sublinear, acuminate; Sty very short. D. edwardii P. J. Braun & al. (Bromelie 2008 (3): 116–123, ills., 2008). Type: Brazil, Goiás (Esteves Pereira 467 [UFG]). — Lit: Braun & Esteves Pereira (2009a). Distr: Brazil (Goiás); granitic rocks and granite-derived soils in very dry Cerrado vegetation. I: Braun (2018: 75). [2] Ros with >50 living leaves, to 1 m ∅, usually solitary, sometimes forming small groups, with a stem to 50  17 cm in very old plants, compactly covered by old leaves; L 56  to 4.3 cm, sheath 3.2  4.3 cm, white, apically densely yellowish-lepidote, L lamina erect to slightly arching, narrowly triangular to lanceolate, apex a flat brownish pungent tip to 35 mm long, light-green to chocolate-brown, adaxially lustrous, abaxially white-lepidote, strongly veined, sparsely serrate, Sp pungent, uncinate, usually antrorse, retrorse towards the leaf tip, to 5.5 mm, 6–18 mm apart; Inf to 1.8 m, compound; peduncle hard and strong, terete, slightly sinuous and sulcate, 50 cm, green to reddish-green; peduncular Bra linear to lanceolate, acuminate, pungent, exceeding or equalling the internodes, 3.1–4.4  1 cm, pale green, soon stramineous, serrate; fertile Inf part lax, 1.4 m, ascending, below the first

910

branch with 1–2 dormant buds; Br 7–15, ascending, spreading, flexible, 26–53 cm, dark rose, sparsely white-lepidote; primary Bra like the upper peduncular bracts but shorter; floral Bra narrowly triangular to lanceolate, 22  7 mm (lower) to 6 mm (upper), brownish-green, soon stramineous, densely white-lepidote, serrate; Fl ascending, 18–20  7–9 mm; Ped 0–2 mm; Sep lanceolate, 11  5.9 mm, yellowish to reddishorange, scatteredly white-lepidote; Pet obtuse, 11  8 mm, orange but more yellowish than the sepals, basally light orange to yellow; Fil 9 mm, apical 1.8 mm free above the 7 mm high common tube, pale yellow; Anth to 2.6  1.2 mm, yellow, recurved at anthesis; Ov narrowly subpyramidal, 5 mm; Sty 1.3 mm, yellow, Sti 2.5 mm, yellowish-orange, hidden at anthesis; Fr ellipsoid-ovate, apex acuminate, 15–16  8–10 mm, dark brown, lustrous. Similar to D. goiana and D. uleana, but forming extraordinary stems to 50 cm long in old plants. D. elata Mez (in A. & C. de Candolle, Monogr. Phan. 9: 508, 1896). Type: Brazil, Minas Gerais (Schwacke 8739 [B, F [photo]]). — Distr: Brazil (Minas Gerais); rocky fields. [2?] Insufficiently described; L to 40  1.7 cm, sheath suborbicular, large, L lamina narrowly triangular, green, subglabrous adaxially, laxly serrate, Sp 4 mm; Inf probably >1 m, laxly paniculate; peduncle stout, 50 cm, soon glabrous; peduncular Bra subfoliaceous, equalling or exceeding the internodes, sparsely serrulate; fertile Inf part many-flowered, narrow, rachis tomentulose but soon glabrous; Br elongate, laxly flowered; primary Bra much shorter than the sterile branch bases; floral Bra reflexed, broadly ovate, acute, shorter than the flowers or equalling the sepals (lowest), finely serrate; Fl spreading or reflexed, 12 mm; Ped short; Sep elliptic, 7 mm, broadly rounded, glabrous; Pet erect, lamina large, suborbicular, neither carinate nor undulate when dry, orange; St included; Fil free above the common tube; Anth short, subtriangular, apiculate, scarcely recurved; Sty to as long as the ovary. — [F. Krapp]

F. Krapp and U. Eggli

D. elisabethae S. Winkler (Doc. Nat. 3: 36–37, fig. 3, 1982). Type: Brazil, Rio Grande do Sul (Winkler 741 [ICN]). — Distr: Brazil (E Rio Grande do Sul); barren granite rocks in moist ground. [1?] Ros to 30 cm ∅; L 35  3 cm, sheath dark, L lamina adaxially light green, abaxially white-lepidote, margins conspicuously serrate, Sp curved, to 5 mm, irregularly arranged; Inf to 1.7 m, usually branched or rarely simple, branches not further described; peduncle reddish to red (younger parts), white-pubescent; peduncular Bra serrate; floral Bra elliptic, acuminate, pubescent, to 8 mm; Fl sessile; Sep lanceolate-ovate, to 12 mm, yellow, white-pubescent, tip rounded; Pet orbicular, to 18 mm, light yellow, finely lepidote, ecarinate, margins shortly ciliate; St included or equalling the petals; Fil free above the common tube; Ov 3 mm; Sty 3 mm. The incomplete description from the protologue was expanded on the base of the comparison with D. pontesii in the protologue of the latter. The flowers are described as sessile in the protologue, but as pedicellate in the comparison with D. pontesii. D. elongata Mez (in A. & C. de Candolle, Monogr. Phan. 9: 529, 1896). Type: Brazil, sine loco (Sellow Brom. Paris 58 [P, GH [photo]]). — Distr: Brazil (Bahia); on rocks. I: Smith & Downs (1974: 568). [3a] Ros not described; L to 40  1.5 cm, sheath broadly ovate, L lamina narrowly triangular, appressedly pale-lepidote abaxially, serrate, Sp stout, straight or curved, 6 mm; Inf almost 1 m, simple; peduncle slender, soon glabrous; peduncular Bra ovate-triangular (upper), much shorter than the internodes, 10 mm; fertile Inf part very laxly few-flowered, soon glabrous; floral Bra spreading or subreflexed, ovate-triangular, shorter than the sepals; Fl suberect, 11 mm, subsessile; Sep broadly ovate, 7–8 mm, obtuse; Pet lamina suberect, broadly obovate, obscurely carinate, colour not described; St included; Fil free above the common tube; Anth triangular, acute, straight or slightly recurved; Sty very short. — [F. Krapp]

Dyckia BROMELIACEAE

911

D. eminens Mez (Bot. Jahrb. Syst. 30 (Beiblatt 67): 5, 1901). Type: Brazil, Goiás (Glaziou 22192-A [B, K, P]). — Distr: Brazil (Goiás); ecology not reported. [3a?] Ros not described; L 30  >2 cm, sheath not described, L lamina narrowly triangular, with a strongly pointed hard tip, sparsely appressedlepidote abaxially, laxly serrate, Sp 1 mm, far apart; Inf 1 m or more, simple; peduncle stout, soon glabrous; peduncular Bra broad with narrowly triangular pungent lamina, exceeding the internodes, serrulate; fertile Inf part lax, densely furfuraceous; floral Bra reflexed, broadly ovate, acuminate (lower), 20 mm; Fl spreading or subreflexed, 14 mm; Ped 70  4.5 cm, lamina narrowly triangular, punctulate and sublustrous adaxially, finely appressedly white-lepidote and striate abaxially, Sp slightly curved, to 9 mm, 20–30 mm apart; Inf probably 1 m or more, few-branched; peduncle stout, glabrous; peduncular Bra broadly ovate, acute, 15–20 mm, upper much shorter than the internodes; fertile Inf part lax, glabrous, terminal part to 60 cm; Br densely flowered, subspreading, to 16 cm; primary Bra like the peduncular bracts, distinctly shorter than the sterile branch bases; floral Bra broadly ovate, acute, except for the lowest shorter than the sepals; Fl yellowish, subsessile; Sep suborbicular or broadly ovate, 6 mm, apiculate; Pet elliptic, 11–13 mm, obtuse, not unguiculate, pale-margined when dry, yellowish; St long-exserted, slightly exceeding the style; Fil free above the 1 mm high common tube; Anth straight, 3.5 mm; Sty slender, elongate, longer than the ovary; Se narrowly winged. — [F. Krapp]

914

D. ferox Mez (in A. & C. de Candolle, Monogr. Phan. 9: 511, 1896). Type: Paraguay (Kuntze s.n. [NY]). — Distr: Brazil (Mato Grosso), Paraguay, Bolivia (Santa Cruz), Argentina (Chaco, Corrientes, Entre Ríos, Formosa); dry fields and woods. I: Smith & Downs (1974: 544). Incl. Dyckia meziana Kuntze (1898); incl. Dyckia hamosa Mez (1906)  Dyckia ferox ssp. hamosa (Mez) Hassler (1919)  Dyckia ferox fa. hamosa (Mez) Castellanos (1933); incl. Dyckia ferox fa. vulgaris Hassler (1919); incl. Dyckia ferox fa. australis Hassler (1919) (nom. inval., Art. 26.2). [1] Ros slowly offsetting and clump-forming; L 20–50  0.8–2 cm or longer, sheath suborbicular, 3 cm, L lamina narrowly triangular, closely appressedly pale-lepidote abaxially, Sp acicular, antroretrorsely curved, 4–6 mm; Inf 60 cm, simple or few-branched with short branches; peduncle slender, cinereous-tomentose, soon glabrous; peduncular Bra very broadly ovate, abruptly acute, shorter than the internodes, entire; fertile Inf part lax, cinereous-tomentose to glabrous; primary Bra scarcely larger than the floral bracts; floral Bra spreading or reflexed, suborbicular-ovate, obtuse or apiculate, much shorter than the sepals, subglabrous; Fl subsessile, narrowly funnel-shaped; Sep very broadly elliptic, 5–7 mm, very broadly rounded at the apex, subglabrous; Pet lamina suberect, subrhombic, to 12 mm, rounded, scarcely undulate, little if at all carinate, yellow; St shorter than the petals; Fil free above the common tube; Anth triangular, subacute, strongly recurved at the apex; Sty very short. — [F. Krapp] D. ferrisincola O. B. C. Ribeiro & Leme (J. Bromeliad Soc. 65(1): 19–22, ills. (incl. p. 18), 2015). Type: Brazil, Minas Gerais (Ribeiro 249 [BHCB]). — Distr: Brazil (C Minas Gerais: Sarzedo); white iron-rich rock outcrop (“canga couraçada”), in full sun, 1200 m; known from the type area only. [3b] Ros 40 cm ∅, offsetting; L 11–14, suberect to arching-spreading, coriaceous, sheath 3.5–4  4.5–5 cm, distinct from the lamina, whitish, both faces glabrous and glossy, L lamina

F. Krapp and U. Eggli

narrowly triangular, 20–22  2.5–3 cm, strongly canaliculate, mostly wine-red or sometimes green, adaxially glabrous and glossy, abaxially whitelepidote at the base but glabrous further up, distinctly veined, tip acuminate, pungent, margins laxly spinose, Sp narrowly subtriangular, spreading to retrorse, 3.5–4 mm, 20 mm apart; Inf 60–80 cm, erect, simple; peduncle 35–40 cm, reddish-green, densely white-hairy; peduncular Bra broadly triangular with a long lanceolate acuminate-caudate lamina, 15–17  6–8 mm, erect, with prominent midvein and appearing carinate, densely white-lanate; fertile Inf part 30–40 cm, laxly 15- to 40-flowered, rachis red, densely white-lanate; floral Bra subtriangular, acuminate, 10–11  5–6 mm, with prominent midvein and appearing carinate, stramineous towards the tip, minutely denticulate; Fl 15–16 mm, porrect-spreading, tubular, unscented; Ped 4–5 mm, white-lanate; Sep broadly ovate to suborbicular, 6–7  5–6 mm, convex, ecarinate, reddish, sparsely white-lanate, margins with recurved fimbriate trichomes; Pet broadly spatulate, emarginate, 11–12  9–10 mm, orange, basally connate for 2.5 mm, ecarinate; St included and distinctly shorter than the petals; Fil yellow, connate for 2 mm above the common tube; Anth oblong-lanceolate, 2 mm, straight, base bilobed; Ov suboblong, 6  2 mm, yellow; Sty 2 mm, distinct, orange; Sti conduplicatespiral, yellow, lobes shortly lacerate; Fr and Se unknown. A member of the D. saxatilis complex, and compared with D. saxatilis in the protologue. — [U. Eggli] D. ferruginea Mez (in A. & C. de Candolle, Monogr. Phan. 9: 533, 1896). Type: Brazil, Mato Grosso (Kuntze s.n. [NY]). — Distr: Brazil (Mato Grosso); vertical walls of red sandstone. I: Smith & Downs (1974: 535). [3a] Ros not described; L to 80  2.5 cm, sheath suborbicular, 5 cm, castaneous, glabrous, L lamina narrowly triangular, soon glabrous adaxially, subappressedly white-lepidote abaxially, margins densely and coarsely serrate, Sp curved, 10 mm; Inf 1 m or more, simple; peduncle stout, ferrugineous-tomentulose or glabrous with age;

Dyckia BROMELIACEAE

peduncular Bra leaf-like, imbricate, the upper ones linear-lanceolate, entire; fertile Inf part dense or  interrupted, many-flowered, rachis stout, densely ferrugineous-tomentulose; floral Bra divergent to spreading, narrowly triangular, to 7 mm, shorter than the sepals; Fl suberect to spreading, to 15 mm; Ped very short; Sep ovate, 7–8.5 mm, broadly rounded; Pet lamina broadly elliptic, 10–14 mm, obtuse, ecarinate, suberect, yellowish-green, ferrugineous-tomentulose outside; St exserted ( equalling the petals according to the protologue); Fil free above the common tube; Anth linear, obtuse, scarcely recurved; Ov sparsely lepidote; Sty as long as the ovary. The protologue is based on immature flowers according to Smith & Downs (1974), and in fresh material, the stamens are exserted. D. floribunda Grisebach (Gött. Abh. 24: 331, 1879). Type: Argentina, Córdoba (Hieronymus 566 p.p. [GOET]). — Lit: Vesprini & al. (2003: ecology). Distr: Argentina (Catamarca, Córdoba, La Rioja, San Luís, San Juan, Jujuy, Salta); dry rocky slopes. I: Smith & Downs (1974: 549); Subils (2009: 349, 423). Incl. Dyckia gilliesii Baker (1889); incl. Dyckia chaguar A. Castellanos (1931). [1] Ros densely many-leaved, acaulescent, with  sessile offsets; L 20–100  2.5 cm, sheath suborbicular, 3–6 cm, pale, glabrous, L lamina attenuate to a pungent tip, white- or cinereouslepidote esp. abaxially, laxly serrate, Sp curved, 4–6 mm; Inf 60–100 cm, paniculate; peduncle elongate, glabrescent; peduncular Bra broadly triangular-ovate, acuminate, thin, shorter than the internodes; fertile Inf part laxly branched, lepidote at least at anthesis, Br subspreading, slightly flexuous, laxly to densely flowered; primary Bra ovate, apiculate, small; floral Bra ovate-triangular, acute or apiculate, much exceeded by the sepals; Fl divergent to spreading, orange; Ped very short at anthesis, to 4 mm in fruit; Sep broad, 7 mm, rounded, obtuse or apiculate, strongly convex; Pet obovate-elliptic, 10–14 mm, broadly rounded and emarginate, undulate, carinate; St slightly exserted; Fil free above the very short common tube; Anth sublinear, acute,  recurved; Sty free, very short.

915

D. formosensis Leme & Z. J. G. Miranda (Phytotaxa 67: 15–17, ills., 2012). Type: Brazil, Goiás (Leme & Miranda 6451 [RB, HB]). — Distr: Brazil (Goiás); open Cerrado vegetation. [3b] Ros solitary, subdense, 18-leaved; L 60–65  3.5–4 cm, rigid, thickly coriaceous, sheath wider than the lamina, L lamina narrowly triangular, canaliculate, suberect, straight, acuminate and with spinescent tip, green to reddishferrugineous, veined, adaxially inconspicuously white-lepidote, abaxially densely non-obscuringly white-lepidote, margins glabrescent and laxly spinose, Sp shortly acicular, retrorse-uncinate, 1.5–5 mm, pale castaneous, glabrous, 10–20 mm apart; Inf 1.9–2 m, simple, erect; peduncle 95–100 cm, reddish-ferrugineous, densely whitesublanate; peduncular Bra subfoliaceous (lowest) to narrowly triangular-lanceolate (upper), acuminate, with spinescent tip, erect, distinctly shorter than the internodes, stramineous, inconspicuously white-sublanate, veined, densely spinulose, uppermost like the floral bracts; fertile Inf part 70- to 75-flowered, lax basally, subdense apically, 90–100 cm,  straight, reddish-ferrugineous towards the base and orange towards the apex, densely white-sublanate; floral Bra suberect to slightly reflexed, ovate-triangular to lanceolate, acuminate-caudate, 15–35  6–15 mm, stramineous, inconspicuously white-sublanate abaxially, veined, densely spinulose; Fl spreading to slightly reflexed at anthesis,  erect afterwards, 18–22 mm, campanulate, slightly fragrant; Ped stout, 3–5 mm; Sep broadly ovate, 8–10  7–8 mm, apex obtuse, ecarinate, convex, orange, densely and coarsely white-lepidote, margins with fimbriate trichomes, remotely and densely spinulose-crenulate; Pet broadly spatulate, 13–14  8–12 mm, obtuse and emarginate, the base narrower, ecarinate, orange, glabrous; St included; Fil 5–6 mm connate above the 2–4 mm high common tube, complanate, 9–10 mm, orange; Anth subtriangular, 3–5 mm, strongly recurved at anthesis, acute and apiculate, base bilobed; Ov narrowly oblong-subpyramidal, yellowish; Sty slender, 3–3.5 mm, orange; Sti slightly exceeding the anthers after anthesis, 1.5 mm, orange, margins crenulate-lacerate. Closely related to D. brasiliana, and also similar to D. goiana. — [F. Krapp]

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D. fosteriana L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 107, t. 106, 1943). Type: Brazil, Paraná (Foster 1154 [GH]). — Distr: Brazil (Paraná). [3b] Ros dense, flat, clustering; L numerous, 9–17  0.8–1.2 cm, arching-recurved, sheath suborbicular, L lamina narrowly triangular, covered with a cinereous membrane of fused scales, Sp antroretrorsely curved, 2–4 mm; Inf 10–45 cm, simple; peduncle sparsely flocculose; peduncular Bra lanceolate, acuminate, exceeding the internodes or the upper slightly shorter, densely lepidote, serrulate; fertile Inf part 3.5–13 cm, densely lepidote; floral Bra like the peduncular bracts, carinate towards the apex, lowest  equalling the flowers, serrulate; Fl suberect or spreading; Ped 2 mm; Sep ovate, 6–9 mm, acute, carinate, ferrugineous-lepidote; Pet lamina spreading, rhombic, 8–10 mm, orange; St included; Fil short-connate above the common tube; Sty very short. — [F. Krapp]

pale or glaucescent, densely appressed-lepidote abaxially, Sp 2–5 mm; Inf 1 m, amply paniculate; peduncle ascending, stout, lepidote when young; peduncular Bra sparse, acute from a broadly ovate base, exceeding the internodes or the upper shorter, minutely serrulate; fertile Inf part many-flowered, subferrugineous-tomentellous, Br spreading, subdense towards the apex, to 30 cm; primary Bra like the upper peduncular bracts, usually shorter than the sterile branch bases; floral Bra broadly ovate, abruptly acute, shorter than the flowers, the lower serrulate; Fl spreading-ascending, 11–18 mm, narrowly funnel-shaped; Ped to 3 mm; Sep ovate, 5–8 mm, acute, carinate, fimbriate; Pet subovate, rounded, undulate, strongly carinate, spreadingrecurved, yellow or pale yellow; St much shorter than the petals but well visible in the throat; Fil free above the common tube; Anth broadly truncate at the base, attenuate, strongly recurved; Ov narrowly ovoid; Sty very short.

D. fosteriana var. fosteriana — Distr: Brazil (Paraná); sandstone outcrops. I: Smith & Downs (1974: 531). [3b] L lamina 0.8–1 cm wide, repand-serrate with recurved Sp 2–3 mm long; Inf peduncle slender; Fl dense to lax; Sep 6 mm. — [F. Krapp]

D. glabrifolia Leme & O. B. C. Ribeiro (Phytotaxa 67: 18–20, ills., 2012). Type: Brazil, Minas Gerais (Leme & al. 7273 [RB, HB]). — Distr: Brazil (Minas Gerais); on rocky quartzite outcrops, Campo Rupestre vegetation. [2] Ros solitary or in small groups, 17- to 25-leaved; L 25–42  2–4 cm, distinctly succulent mainly towards the base, sheath subreniform, 3.5  7.5 cm, whitish towards the base, green to reddish at the apex, lustrous, glabrous, L lamina narrowly triangular, attenuate, flat, suberectarcuate, acuminate-caudate, with pungent tip, green to reddish, lustrous and veined adaxially, opaque abaxially, glabrous on both faces except adaxially at the base, margins laxly spinose, Sp straight to slightly antrorse, narrowly triangular, 1–2 mm, tip castaneous, 7–17 mm apart; Inf 90–150 cm, erect, usually simple or sometimes with a short branch near the base; peduncle 36–81 cm, greenish, sparsely white-sublanate, soon glabrous; peduncular Bra subfoliaceous (lowest), erect, narrowly ovate-triangular, acuminate, 15–30  6–10 mm, from equalling to distinctly shorter than the internodes, green or reddish, stramineous at anthesis, subdensely white-sublanate basally, glabrous apically, veined,

D. fosteriana var. robustior L. B. Smith (Phytologia 19: 283, 1970). Type: Brazil, Paraná (Hatschbach 17310 [US]). — Distr: Brazil (Paraná); in soil. [3b] L lamina 1.2 cm wide, laxly serrate but scarcely repand, the mostly antrorse Sp to 4 mm; Inf peduncle stout, 6 mm ∅; Fl dense; Sep to 9 mm. — [F. Krapp] D. frigida Hooker fil. (Curtis’s Bot. Mag. 103: t. 6294 + text, 1877). Type: Brazil (Anonymus ex Hort. Linden s.n. [K, LG [status?]]). — Distr: Brazil (Paraná); on rocks. I: Smith & Downs (1974: 519). Incl. Dyckia regalis Linden & Morren ex Baker (1889) (nom. inval., Art. 34.1a). [1] Ros robust, acaulescent; L 100  3.5 cm, sheath reniform, 9 cm wide, dark castaneous, L lamina narrowly triangular, with pungent tip,

Dyckia BROMELIACEAE

densely and minutely denticulate; fertile Inf part 50- to 110-flowered, 33–77 cm, rachis straight, greenish to pale orange, subdensely to densely pale-lanate; floral Bra spreading to suberect, narrowly subovate-triangular (lowest), acuminatecaudate, 10–25  4–8 mm, stramineous towards the apex at anthesis, pale-lanate, veined, denticulate to subentire; Fl spreading at anthesis, erect afterwards, 16–19 mm, broadly tubular with narrowed throat, unscented; Ped 2–4 mm, orange, densely white-sublanate; Sep ovate to broadly ovate, 6–8  6 mm, convex, apex subacute and apiculate, ecarinate, orange, subdensely whitelanate, fimbriate; Pet broadly obovate-spatulate, 12–13  10–11 mm, apex obtuse to subacute and inconspicuously apiculate, ecarinate, orange, glabrous, entire or sometimes irregularly crenulate near the apex; St included; Fil 9–10 mm, 3–4 mm connate above the 1 mm high common tube (according to the protologue, completely connate according to Fig. 3 in Pinangé & al. (2017)), yellowish towards the apex; Anth narrowly subtriangular, 4 mm and straight before anthesis, after anthesis distinctly smaller and strongly recurved, base bilobed, acute; Ov narrowly suboblong, 6.5 mm, pale yellow; Sty 1 mm, Sti 1.5 mm, yellow, margins minutely crenulate-lacerate; Fr broadly ovoid, beaked, 15  11 mm, dark castaneous, lustrous; Se flat, asymmetrical, subcuneate, 3–4  2–2.5 mm. Closely related to D. trichostachya, and also similar to D. weddelliana. D. glandulosa L. B. Smith & Reitz (Phytologia 14: 484, t. 1, figs. 20–24, 1967). Type: Brazil, Minas Gerais (Magalhães 18959 [HB, NY, US [photo]]). — Distr: Brazil (Minas Gerais: Diamantina Plateau); among rocks in Campo Rupestre vegetation, 1400–1800 m. I: Smith & Downs (1974: 565). [3a] Ros not described; L to 23  2.3 cm, sheath suborbicular, 2 cm, L lamina narrowly triangular, attenuate to an abruptly acute pungent apex, smooth and glabrous adaxially, appressedly cinereous-lepidote abaxially, laxly serrate, Sp antrorsely uncinate, 2 mm; Inf 70 cm, simple; peduncle laxly glandular-lepidote with yellow trichomes; peduncular Bra triangular to linear-

917

laminate (lowest), small, shorter than the internodes (uppermost), peltate-lepidote (lower) to vestite with glandular trichomes alone, entire; fertile Inf part laxly many-flowered, to 27 cm, all parts incl. pedicels and sepals laxly glandulartomentose; floral Bra triangular-ovate, acuminate, the lowest exceeding the sepals, entire; Fl spreading; Ped 5 mm, subcylindrical; Sep ovate, 7 mm, obtuse, cucullate, ecarinate, rugose when dry as if initially fleshy; Pet lamina rhombic, 12 mm, obtuse, carinate, orange; St included; Fil free above the common tube; Sty 1 mm. — [F. Krapp] D. goehringii E. Gross & Rauh (Trop. subtrop. Pfl.-welt 79: 12–14, ills., 1991). Type: Brazil, Goiás (Esteves Pereira s.n. in Rauh 67622 [HB, HEID]). — Distr: Brazil (Minas Gerais); rocky fields, on rocks or quartz sand. I: Braun & Esteves Pereira (2006b). – Fig. 6. [3b] Ros many-leaved, 50 cm ∅, 30 cm high, with a short stem; L to 23  2 cm, reflexed, succulent, sheath broadly ovate, appressed to the stem, 3  3.5 cm, whitish adaxially, castaneous and lustrous abaxially, glabrous, L lamina triangular, with pungent tip, both faces appressedly white-lepidote, adaxially with conspicuous imprint from neighbouring leaves, abaxially veined, trichomes arranged in distinct rows, margins spinose, Sp strong, antroretrorse, white-lepidote, 10 mm apart; Inf 80 cm, usually simple; peduncle 50 cm, green, glabrous at the base, upper part white-lepidote; peduncular Bra broadly lanceolate, acuminate, 10 mm and much shorter than the internodes (upper), the lowest longer than the internodes, sparsely lepidote, veined, indistinctly serrulate; fertile Inf part 25-flowered, lax, 35 cm, rachis green, densely white-lanate, terete; floral Bra broadly ovate, acuminate, much shorter than the sepals, 5  6 mm, densely white-lanate, indistinctly serrulate; Fl spreading, 15 mm, subsessile; Sep oblong-ovate, 7 mm, obtuse, ecarinate, slightly succulent, greenish-orangered, white-floccose, margins hyaline, indistinctly serrulate; Pet erect, rhombic, 10  10 mm, obtuse, ecarinate, forming a narrow tube, orange-red, margins indistinctly serrulate; St slightly exserted; Fil short-connate above the common tube

918

F. Krapp and U. Eggli

Fig. 6 Dyckia goehringii (Esteves Pereira s.n. in Rauh 67622: Brazil; Goiás, near Portel^andia; type collection). (Copyright: U. Eggli)

(or completely free according to Fig. 3 in Pinangé & al. (2017)), basally broadly complanate, attenuate; Anth golden-yellow, sagittate; Sty very short, Sti papillate. Related to D. crassifolia and D. leptostachya. — [F. Krapp]

persistently brown-black-lepidote; Pet spreading, elliptic, 10–12 mm, rounded, carinate, reddishorange; St  equalling the petals; Fil shortly connate above the common tube; Sty 1 mm. Similar to D. elata and compared with D. edwardii in the protologue of the latter.

D. goiana L. B. Smith (Phytologia 14: 479, t. 1, figs. 1–5, 1967). Type: Brazil, Goiás (Irwin & al. 15163 [US, NY]). — Distr: Brazil (Goiás); Cerrado vegetation, 800 m. I: Smith & Downs (1974: 528); Esteves Pereira & Gouda (2017: 181). [1] Ros 1.5–1.8 m ∅; L 80–100 cm long, adaxially lepidote, margins serrate but apical 1=3 entire; Inf to 2.5 m, laxly compound with up to 8 suberect branches; peduncle stout, soon glabrous; peduncular Bra presumably foliaceous (lower) to ovate with a long narrowly triangular apex (upper), exceeding the internodes, serrulate; fertile Inf part 100 cm, initially finely cinereouslepidote, sublax; Br slender, slightly flexuous, to 50 cm; primary Bra like the upper peduncular bracts, to 70 mm; floral Bra broadly ovate, longacuminate (lower) to apiculate (upper), exceeding the flowers (lower) or shorter than the sepals (upper), serrulate; Fl spreading and slightly descending; Ped to 4 mm in fruit, stout; Sep broadly deltoid-ovate, to 6 mm, ecarinate,

D. gouveiana Leme & O. B. C. Ribeiro (Phytotaxa 67: 21–23, ills., 2012). Type: Brazil, Minas Gerais (Leme & al. 8485 [RB]). — Distr: Brazil (Minas Gerais: Diamantina Plateau); rock outcrops in Campo Rupestre vegetation, in shallow humus on rocks; only known from the type locality. [3b] Ros dense, 12- to 18-leaved; L 12–18  2–2.6 cm, strongly coriaceous, sheath inconspicuous, L lamina narrowly triangular, strongly canaliculate, suberect to nearly erect and unilaterally curved, acuminate and spinetipped, light green, obscurely white-lepidote on both faces, veined mainly adaxially, subdensely to densely spinose, Sp narrowly triangular, straight to prevailingly antrorse-uncinate, 1–2.5 mm, with pale castaneous tip, 2–7 mm apart; Inf 35–60 cm, simple, erect; peduncle 18–30 cm, dark reddishgreen, sparsely to subdensely white-lepidote, glabrescent; peduncular Bra base suborbicular to subreniform and slightly gibbose, lamina long narrowly triangular and acuminate-spinescent,

Dyckia BROMELIACEAE

ecarinate, to 18–45  3–6 mm, longer (lowest) to shorter (uppermost) than the internodes, stramineous towards the apex, veined, subdensely white-lepidote, densely spinulose; fertile Inf part 8- to 26-flowered, lax, 13–24 cm, straight, dark reddish, (sub-) densely non-obscuringly whitesublanate; floral Bra ovate-triangular, acuminate-caudate, spreading at anthesis, ecarinate with raised midvein, the upper ones convex at the base, 13–19  7–10 mm, soon becoming stramineous towards the apex, (sub-) densely white-lepidote, finely veined, densely spinulose to inconspicuously denticulate; Fl spreading, 16–17 mm, subtubular, unscented; Ped 3–4 mm, stout, red, densely white-lanate; Sep broadly ovate, 7–8  5–6 mm, acute and apiculate, ecarinate, convex, reddish-orange, densely white-lanate, margins inconspicuously denticulate near the apex; Pet broadly spatulate, 11–12  9 mm, apex broadly acute, ecarinate, entire, orange, glabrous; St included; Fil 1 mm connate above the 3 mm high common tube, 8 mm, pale orange towards the apex; Anth narrowly subtriangular, 3.5–4 mm, strongly recurved at anthesis; Ov narrowly suboblong, 5 mm, pale yellow; Sty 2.5 mm; Sti 1.5 mm, orange, margins minutely crenulate. Compared with D. mezii in the protologue (as D. argentea). — [F. Krapp] D. gracilis Mez (in A. & C. de Candolle, Monogr. Phan. 9: 516, 1896). Type: Bolivia, Santa Cruz (d’Orbigny 1006 [P, F [photo], G, GH [photo]]). — Distr: Brazil (Mato Grosso do Sul), Bolivia (Santa Cruz), Argentina (Chaco); seasonally deciduous lowland forests and savannas, dry rocky soil, 230–500 m. I: Smith & Downs (1974: 576); Vásquez & Ibisch (2012: 127). [3b] Ros not described; L to 30  2–2.5 cm, suberect, L lamina narrowly triangular, quite green adaxially, closely appressedly pale cinereous-lepidote abaxially, margins serrate, Sp 4 mm; Inf 50 cm, simple; peduncle very slender, glabrous; peduncular Bra remote, very broadly ovate with a short acuminate apex, entire; fertile Inf part lax, few-flowered, minutely furfuraceous; floral Bra reflexed, suborbicular-ovate, apiculate,

919

4 mm, much shorter than the sepals; Fl suberect, slender, subsessile; Sep very broadly elliptic, 7–9 mm, emarginate, nearly glabrous; Pet erect, obovate, 14–18 mm, rounded, scarcely if at all carinate, bright yellow; St equalling or slightly exceeding the petals; Fil connate above the common tube; Anth linear, acute, strongly recurved; Sty 2–3 shorter than the ovary. Lost for some 180 years before found again in Bolivia (Vásquez & Ibisch 2012). The recently described D. barthlotii is similar (see there for differences). Paggi & al. (2015) record the taxon also from the Brazilian state Mato Grosso do Sul. D. grandidentata P. J. Braun & Esteves (Cact. Succ. J. (US) 80(6): 323, ills. (pp. 322–323), 2008). Type: Brazil, Mato Grosso do Sul (Esteves Pereira & Braun 659 [UFG, HAL]). — Distr: Brazil (Mato Grosso to Sul); on barren reddishblack rocks among shrubs, trees, orchids and cacti. [2] Ros to 37 cm ∅ but usually much smaller, stem 6  5–6 cm, forming a small clump; L 23–29, to 23  3.1 cm, 5 mm thick, sheath to 4 cm wide, L lamina recurved, spreading, slightly coriaceous, succulent, glabrous, narrowly triangular to lanceolate, canaliculate, apex a hard brownish spine to 7 mm long, veined on both faces, margins with an inconspicuous pale yellow stripe, serrate, Sp pungent, uncinate, predominantly retrorse, 7–11 mm, narrowly triangular, shiny, light green to light yellow, with a brown-reddish tip, 14–23 mm apart towards the leaf tip, basally  connected; Inf to 80 cm, usually simple or rarely with 1–3 branches; peduncle erect or slightly sinuous, 40 cm, base reddish-green, red distally, shiny green to brownish-green elsewhere, glabrous or partly creamy-lepidote; peduncular Bra narrowly triangular to lanceolate, pungent, 58 mm (lowest) to 6 mm (uppermost), green to brownish, finely serrate; fertile Inf part erect, to 39 cm, rachis dark rose, distally rose to orange and red; floral Bra narrowly triangular to lanceolate, to 4.7 mm, rose, drying parchmentlike; Fl 17  8 mm, orange to yellowish, subsessile; Sep slightly triangular, 7.5  6.2 mm, succulent, abaxially carinate, reddish, margins pale; Pet 12  5–7 mm, orange; St exserted; Fil

920

apically free above the common tube, to 9 mm; Anth 5 mm, light yellow, strongly curved outwards, acuminate; Ov elongate-pyramidal, 6  2 mm, light greenish-white; Sty 0.7–0.9 mm, orange; Fr ovoid, acute, 13  10 mm, dark brownish, glossy; Se broad, drop-shaped, flattened, with a tiny acute tip towards the hilum, 3–4.5 mm ∅ — [F. Krapp]

F. Krapp and U. Eggli

(Paraná); low woods in Restinga vegetation. I: Smith & Downs (1974: 535). [3a] Only known from fragments; L to 85  4.5 cm, lamina linear-triangular, with pungent tip, initially or soon glabrous adaxially, appressedly white-lepidote abaxially, laxly serrate, Sp antrorsely uncinate, dark, 7 mm; Inf to 1.5 m, simple; peduncle slender, ferrugineoustomentose; peduncular Bra imbricate, lineartriangular from an ovate base, serrulate; fertile Inf part sublax, many-flowered, 40 cm, all parts incl. pedicels and sepals ferrugineoustomentulose; floral Bra narrowly triangular, slightly exceeding the pedicels (lower), serrulate; Fl spreading; Ped to 10 mm, straight, slender; Sep ovate, 8 mm, acute, carinate; Pet subrhombic, 14 mm, curved-spreading, colour not described; St included; Fil free above the common tube; Sty to 2 mm. Insufficiently known.

D. granmogulensis Rauh (Trop. subtrop. Pfl.welt 65: 11–12, fig. 5, 1988). Type: Brazil, Minas Gerais (Rauh 56484 [HEID]). — Distr: Brazil (Minas Gerais); rocky fields, on rocks. [3a] Ros acaulescent, in groups, spreading, many-leaved, 15 cm ∅, to 10 cm high; L to 18  1.5 cm, sheath 2.5  2.5 cm, with brownish base and green tip abaxially, glabrous, L lamina narrowly triangular, long-attenuate, slightly succulent, adaxially dark green and sparsely lepidote, abaxially veined and appressedly whitelepidote, Sp retrorse, hard, small, white; Inf to 60 cm, simple, erect; peduncle to 40 cm, green, glabrous, sparsely white-lanate-lepidote towards the apex; peduncular Bra subfoliaceous (lower) to narrowly triangular-acute (upper), as long as the internodes, sparsely lepidote, slightly serrate; fertile Inf part lax, straight, to 20 cm, green, reddish towards the apex, densely lanate; floral Bra triangular, acuminate, small, shorter than or equalling the pedicels, inconspicuously serrate (lower); Fl spreading to ascending, 8–10  4–5 mm, 3 mm at the throat, in the evening with fishy scent; Ped to 5 mm, stout, green to reddish, pubescent; Sep ovate, 5  3 mm, obtuse, ecarinate, reddish, densely lanate; Pet broadly spatulate, 6  7 mm, broadened towards the tip, obtuse, ecarinate, initially erect, later with reflexed tips, brilliant orange-red; St included; Fil free above the 2 mm high common tube, narrowing apically; Anth centripetally bent; Sty 2 mm, yellow. Compared with D. oligantha (here treated as synonym of D. saxatilis) in the protologue. — [F. Krapp]

D. hebdingii L. B. Smith (Phytologia 21: 90, t. 2, figs. 1–7, 1971). Type: Brazil, Rio Grande do Sul (Croizat 22495 [US]). — Distr: Brazil (Rio Grande do Sul); on rocks. [1] Ros dense, spreading, many-leaved; L 15  >1.5 cm, L lamina narrowly triangular, appressedly cinereous-lepidote on both faces, subdensely serrate, Sp slender, spreading; Inf >1 m, laxly 3 branched; peduncle erect, slender, 45 cm; peduncular Bra very narrowly triangular, longer than the internodes, red, serrulate, divergent, wholly exposing the axis; fertile Inf part densely cinereous-lepidote, Br manyflowered, subdense to lax, to 30 cm; floral Bra broadly ovate, apiculate, 5 mm, much exceeded by the sepals; Fl suberect to spreading and sometimes slightly secund; Ped short; Sep ovate, 4.5 mm, broadly subacute; Pet spatulate, 7 mm, obtuse, yellow; St included; Fil free above the 1 mm high common tube; Sty slender, elongate; Fr 8 mm; Se with a narrow apically pointed wing. Part of the “Prionophyllum” complex, and similar to D. maritima. — [F. Krapp]

D. hatschbachii L. B. Smith (Phytologia 14: 480, t. 1, figs. 9–12, 1967). Type: Brazil, Paraná (Hatschbach 2725 [US]). — Distr: Brazil

D. hohenbergioides Leme & Esteves (J. Bromeliad Soc. 48(3): 124–126, ills., 1998). Type: Brazil, Bahia (Esteves Pereira E-385 [HB,

Dyckia BROMELIACEAE

RB]). — Distr: Brazil (Bahia); ecology not described. [1/2] Ros 30-leaved; L 60  4–5 cm, very rigid, succulent, sheath not described, L lamina uberect, very narrowly triangular, attenuate, acuminate, with pungent tip, greenish, finely veined, densely white-lepidote on both faces, laxly spinose, Sp triangular-uncinate, 5–9 mm, densely whitelepidote; Inf 1.8–2 m, laxly 2–3 branched, erect; peduncle sulcate, 80–100 cm, green, whitesublanate, soon glabrous; peduncular Bra triangular, acuminate-caudate, ecarinate, erect, 50–100  20–30 mm, equalling to slightly exceeding (lowest) to shorter than the internodes (upper), greenish to ochre, white-lepidote to glabrescent, finely veined, laxly serrulate apically or entire; fertile Inf part dense, erect, 70 cm, white-sublanate at anthesis, glabrescent; Br 10, subsessile, densely strobilatedigitate, 7–10 cm, erect to suberect, axis whitelanate; primary Bra like the upper peduncular bracts but gradually reduced towards the apex, distinctly shorter than the branches; floral Bra suborbicular to triangular, acuminate, suberect, concave, ecarinate to slightly carinate near the apex, 10–14  13–15 mm, white-lanate at the base, glabrescent, chaffy towards the tip, minutely denticulate-crenulate to entire; Fl suberect at anthesis,  ovoid-tubular, 16–17 mm, unscented; Ped 2.5 mm, indistinct, stout; Sep suborbicular, 7–8  9 mm, subobtuse to slightly emarginate, very minutely mucronulate, ecarinate, orange-red, sparsely sublanate to glabrescent, entire; Pet subcordate, 13  11–12 mm, truncate-emarginate, ecarinate, erect at anthesis, orange-red, glabrous; St slightly exserted for  2=3 of the anther length; Fil 10 mm, sometimes connate for 1.5 mm above the 1.5 mm high common tube (according to the protologue, completely connate according to Fig. 3 in Pinangé & al. (2017)); Anth 3–4 mm, strongly recurved towards the apex, apiculate, base sagittate; Ov narrowly subpyramidal, 6 mm; Sty indistinct, Sti orange, 1.5 mm, margins irregularly scalloped. Compared with D. beateae in the protologue. — [F. Krapp] D. horridula Mez (Bot. Jahrb. Syst. 30 (Beiblatt 67): 5, 1901). Type: Brazil, Goiás

921

(Glaziou 22194 [B, F [photo], K]). — Lit: Leme & Miranda (2009: with ills.). Distr: Brazil (Goiás: Santo Antonio do Descoberto); rocky soils in Campo Rupestre vegetation, amongst herbs. [3b] Ros compact, 15 cm ∅, with a bulbous base formed by old leaf sheaths, probably solitary; L 35–40, dense, coriaceous, sheath 4.5–5  4–5 cm, L lamina sublinear, 10–12  0.8 cm, (sub-) erect, nearly flat near the base, canaliculate and obtusely carinate above and subulate towards the tip, green basally changing to reddish towards the tip, adaxially sparsely and abaxially densely white-lepidote but scales not obscuring the leaf colour, tip acuminate-caudate, pungent, margins subdensely spinose, Sp spreading to retrorse, 1–1.5 mm, reddish-brown, 2–3 mm apart; Inf 15–25 cm, simple, erect; peduncle 9–17 cm, subdensely white-furfuraceous to glabrous; peduncular Bra 10–20  5 mm, subfoliaceous (lowest) to ovate-subtriangular (upper), erect, stramineous, sparsely white-lepidote; fertile Inf part 3–4 cm, 5- to 7-flowered, subdensely to inconspicuously white-furfuraceous (incl. pedicel and sepals); floral Bra ovate, 7  3 mm, acuminate, veined, stramineous, subspreading to reflexed; Fl 12–14 mm long and ∅, somewhat campanulate, strongly sweet-fragrant; Ped inconspicuous, 2  3.5–4 mm ∅; Sep broadly ovate to suborbicular, 5  4 mm, ecarinate, yellow with green tip to pale orange; Pet broadly obovatespatulate from narrow base, 10  6 mm, upper part spreading to spreading-ascending, yellow; St shorter than the petals but distinctly exposed at anthesis, connivent; Fil yellow, connate over the whole length and forming a tube 8 mm long Anth narrowly subtriangular, 2 mm, dark purplish-brown, basally connate; Ov 2.5  1 mm, yellowish-white; Sty 1 mm, yellowish; Sti conduplicate-spiral, yellow; Fr and Se unknown. A rarely collected species, easily overlooked due to its small size. — [F. Krapp] D. ibicuiensis Strehl (Bromélia 4(4): 14–16, ills., 1999). Type: Brazil, Rio Grande do Sul (Strehl 1098 [HAS 31720]). — Distr: Brazil (C Rio Grande do Sul); rocky outcrops, among grasses and herbs.

922

[2/3b] Ros 20-leaved, 60 cm ∅; L 40  2 cm, sheath not described, L lamina narrowly triangular, light green, flexible, slightly contorted, the inner ones erect, the outer spreading, glabrous adaxially, striate-lepidote abaxially, margins subentire with widely spaced minute Sp only; Inf 2 m, simple or sometimes branched, erect; peduncle 70 (30–140) cm, yellowishgreen, glabrous; peduncular Bra long-lanceolate, acute, 150  12 mm and longer than the internodes (lowest), the others much smaller, lepidote at the base, margins membranous, sparsely irregularly spinose, soon drying; fertile Inf part densely to sparsely flowered, robust, 55 (40–80) cm, slightly lepidote to glabrous, yellowishgreen; floral Bra narrow, slightly carinate basally, ending in a long acute pungent tip, 40 mm (lowest) or shorter (uppermost), lemon-yellow, finely serrate; Fl spreading, tubular with spreading limb, 20–24 mm; Ped 6–7 mm; Sep narrow, ovate, 10–15 mm, apex rounded, ending in a short spine, slightly carinate, yellow; Pet free, tongueshaped, 15–20 mm, distally spreading, greenishyellow; St included; Fil of their length free above the common tube; Fr 13–15  10 mm, dark brown at the apex, otherwise almost black, shiny; Se base acute, apex rounded, with a lateral wing, 4 mm wide, brown. Similar to D. dusenii. D. ibiramensis Reitz (Sellowia 14: 104, fig. 4, 1962). Type: Brazil, Santa Catarina (Reitz & Klein 2635 [HBR, US]). — Lit: Hmeljevski & al. (2011: genetics, conservation). Distr: Brazil (C-E Santa Catarina: Ibirama); ledges along streams, and rock islands, seasonally inundated; only known from the region of the type locality. I: Smith & Downs (1974: 552); Reitz (1983: t. 48). [1/2] Ros solitary or forming small groups with age, acaulescent; L 30–40  2.2 cm, rigid, sheath not described, L lamina narrowly triangular, glabrous above, cinereous-lepidote abaxially, sinuate-serrate, Sp 4 mm; Inf 1 m, paniculate; peduncle glabrous; peduncular Bra erect, foliaceous (lowest) to 15 mm (upper), serrate; fertile Inf part few- to 6-branched, subglabrous or sparsely lanate distally, Br ascending, laxly flowered; primary Bra like the upper peduncular

F. Krapp and U. Eggli

bracts, much shorter than the sterile branch bases; floral Bra triangular, recurved, pungent, shorter than the sepals, serrate; Fl suberect to reflexed; Ped 3 (–5) mm; Sep broadly elliptic, 7–9  4 mm, rounded, entire, coriaceous, glabrous; Pet broadly ovate, 14  8 mm, ecarinate with undulate margins, deep orange; St included; Fil free above the short common tube; Sty 2 mm. Hmeljevski & al. (2011) studied the genetic diversity and mating system of this rare endemic, which is confined to a 4 km large rocky outcrop along the Rio Itajaí, and which is classified as critically endangered. The flowers are pollinated by the carpenter bee Xylocopa brasilianorum as well as by the hummingbird Thalurania glaucopsis. D. incana O. B. C. Ribeiro & Leme (J. Bromeliad Soc. 65(1): 23–26, ills. (incl. p. 18), 2015). Type: Brazil, Minas Gerais (Ribeiro 268 [BHCB, HB]). — Distr: Brazil (Minas Gerais: Nova Lima); iron-rich rock outcrops in Campo Rupestre vegetation, 1310 m; known only from the region of the type population. [3a] Ros 12 cm ∅, sometimes asymmetrical, offsetting and forming dense colonies; L 8–10, dense and sometimes secund, subspreading to suberect, sheath 1.5–2  3–4 cm, adaxially whitish, adaxially castaneous, glabrous towards the base, L lamina narrowly triangular, very succulent and rigid, 4.5–6  1–1.4 cm, densely and coarsely cinereous-tomentose on both faces, tip a pungent spine, margins laxly spinose, Sp narrowly triangular, straight to slightly retrorse, 0.5–1 mm, 5–8 mm apart; Inf 25–55 cm, erect, simple; peduncle 25–35 cm, sparsely white-lepidote, pale orange to brownish-red; peduncular Bra 5–10  5 mm, with suborbicular base and a long lanceolate-caudate lamina, erect, stramineous, veined and with raised midvein; fertile Inf part 6–10 cm, subdensely 11- to 16-flowered, rachis reddish, glabrous, distally slightly flexuose; floral Bra broadly ovate to triangular, acuminate to acute, 4.5–5  4.5–5 mm, subspreading to suberect, finely veined, stramineous, slightly crenulate; Fl 12–13 mm, porrect-spreading,  secund, tubular, unscented; Ped 3.5–5 mm, stout, yelloworange, glabrous, curved; Sep broadly ovate to

Dyckia BROMELIACEAE

suborbicular, 4–5.5  3.5–4 mm, orange to reddish, convex, obtuse and apiculate, ecarinate, glabrous except the margins with recurved fimbriate trichomes; Pet broadly obcordate with narrowed base, 7.5  6.5–7 mm, orange-yellow, emarginate, ecarinate, (nearly) erect, basally connate for 1.5 mm; St included and much shorter than the petals; Fil free above the common tube; Anth oblong-ovate, (almost) straight, base bilobed; Ov suboblong-ovoid, 2.5  1.5 mm, yellow; Sty 1 mm, distinct, yellow; Sti conduplicate-spiral, yellow; Fr and Se unknown. Another member of the D. saxatilis complex, and compared with D. brachyphylla, D. macedoi and D. oligantha (here treated as synonym of D. saxatilis) in the protologue. — [U. Eggli] D. inflexifolia Guarçoni & M. Sartori (Ann. Bot. Fenn. 49: 407–410, ills., 2012). Type: Brazil, Minas Gerais (Guarçoni & Sartori 1474 [VIC, R]). — Lit: Büneker & al. (2016: with ills.). Distr: Brazil (C Minas Gerais: Serro); Campo Rupestre vegetation on iron-rich igneous rocks (“pedra canga”) and rocky soil; only known from the small population at the type locality. [2/3a] Ros solitary, dense, 40-leaved, 70 cm ∅, 40 cm high; L 30–35  2.4–3 cm, outer reflexed, youngest inflexed, strongly succulent, sheath elliptic, 3–3.3  5.9–6.3 cm, white, yellowish-cream at the apex adaxially, densely white-lepidote at the apex abaxially, dentate at the apex, L lamina narrowly triangular, indistinctly canaliculate, acute, with pungent tip, green, adaxially completely glabrous, abaxially densely white-lepidote (basally) to pubescent, margins rather densely spinose, Sp irregularly curved, 2.4–3.5 mm, amber with a brown apex, 7–17 mm apart; Inf 0.95–1.45 m, erect, simple or with up to 6 branches; peduncle erect, 25–67 cm, red, densely rusty-tomentose; peduncular Br linear-triangular to triangular with suborbicular base, erect and acuminate and pungent (upper), carinate, succulent (lowest), 36–65  3–6 mm (lowest) to 14–20  6–8 mm (upper), margins vinaceous adaxially (lowest), stramineous (upper), margins (lowest) or completely (upper) densely rusty-lepidote abaxially, fimbriate basally, entire apically; main axis of the fertile

923

Inf part to 45-flowered, 24–56 cm, branches 8to 10-flowered, subdense, erect, rachis red, densely rusty-tomentose, basal-most branches with delayed development, suberect, curved or not, slightly flexuose, 4.5–21 cm; primary Bra similar to the upper peduncular bracts but slightly smaller, 13–15  5–6 mm; floral Bra carinate, triangular, convex, acuminate, pungent, 11–18  5–8 mm, stramineous, densely rustytomentose abaxially, entire to inconspicuously fimbriate; Fl 18–20 mm, ascending to spreading; Ped 5 mm, robust, red; Sep ovate to broadly triangular, 7.2–11  6.3–8.7 mm, obtuse, convex, strongly succulent, red to orange, densely rustytomentose abaxially, fimbriate; Pet rhomboid to elliptic, 11–14  8–11 mm, otuse to retuse, strongly succulent, undulate after anthesis, orange, scarcely tomentose, margins in the upper 1= 2 fimbriate; St included; Fil free above the 1.3–2.9 mm high common tube, 7.4–8.8 mm, yellowish-orange; Anth lanceolate to elliptic, yellow, 3–3.4 mm, base sagittate, apex otusely apiculate; Ov pyramidal, 4.9–5.2 mm, yellow to greenish-yellow; Sty 0.7–0.8 mm, orange to brownish-orange towards the apex, Sti orange, 0.6–0.7 mm; Fr ovoid, acute, 14  10 mm, dark brown. Part of the D. sordida complex together with D. nobilis and D. ursina (Büneker & al. 2016). D. insignis Hassler (Annuaire Conserv. Jard. Bot. Genève 20: 316, 1919). Type: Paraguay, Concepción (Fiebrig 4615 [G, F [photo]]). — Distr: NE Paraguay (Concepción); dry open rocky ground. I: Smith & Downs (1974: 515).  Dyckia grandiflora Mez (1919) (nom. illeg., Art. 52.1); incl. Dyckia insignis [?] flaviflora Hassler (1919); incl. Dyckia insignis var. macrantha Hassler (1919); incl. Dyckia insignis var. obtusiflora Hassler (1919). [3a] Ros shortly caulescent, stem 6–8 cm; L 20–35  1.5 cm, sheaths forming a bulb, broad, brown and lustrous abaxially, glabrous, L lamina linear-triangular, filiform-attenuate, appressedly white-lepidote, laxly serrate, Sp acicular, 3 mm; Inf 30–60 cm, simple; peduncle slender, 25–50 cm, striate, glabrous; peduncular Bra long-acuminate from a triangular-ovate base, the

924

lower 6–7 cm, exceeding the internodes, the upper 2 cm, shorter than the internodes, serrate like the leaves; fertile Inf part laxly 3- to 7-flowered, all parts including the flowers  lepidote; floral Bra like the upper peduncular bracts (lower) to triangular-ovate and filiform-attenuate and shorter (upper), entire; Fl erect to spreading; Ped 7–11 mm; Sep oblong-ovate, 10–12 mm, obtuse, finely fimbriate-serrulate; Pet erect, ellipticoblong, 22–28 mm, subacute or obtuse, red to yellowish (brownish-red in the protologue of D. grandiflora); St exserted 10 mm; Fil free above the 1 mm high common tube; Anth linear, subsagittate at the base, strongly recurved; Ov conical, 10 mm; Sty 25 mm; Fr stoutly ovoid, long-beaked, 20  14 mm; Se with a dorsal falciform hyaline wing, 4 mm. D. irmgardiae L. B. Smith (Phytologia 15: 150, t. 7, figs. 8 and 9, 1966). Type: Brazil, Rio Grande do Sul (Smith & al. s.n. [US]). — Distr: Brazil (NE Rio Grande do Sul); ecology not recorded. I: Smith & Downs (1974: 535). [3a] Ros spreading, very many-leaved, with a short and stout rhizome; L to 56  1.8 cm, subrigid, sheath not described, L lamina very narrowly triangular, soon glabrous adaxially, closely appressedly pale-lepidote abaxially, laxly serrate, Sp antrorsely curved, black, 2 mm; Inf to 1 m, simple, erect; peduncle densely ferrugineouslanate; peduncular Bra densely imbricate, narrowly triangular from a broadly ovate base, ferrugineous-lanate, serrulate; fertile Inf part densely many-flowered, 30 cm, densely ferrugineous-lanate; floral Bra like the upper peduncular bracts, mostly exceeding the flowers, serrulate; Ped 5 mm, stout, compressed; Sep ovate, 10 mm, acuminate, the posterior ones alate-carinate; Pet lamina subrhombic, 15 mm, colour not described; St included; Fil free above the common tube; Sty  none. Similar to D. reitzii. — [F. Krapp] D. irwinii L. B. Smith (Phytologia 14: 483, t. 1, figs. 16–19, 1967). Type: Brazil, Mato Grosso (Irwin & al. 16923 [US, NY]). — Distr: Brazil (E Mato Grosso); open Cerrado scrub and border of fields. I: Smith & Downs (1974: 552).

F. Krapp and U. Eggli

[1] Ros not described; L 60–70  2–2.5 cm, sheaths suborbicular, 4 cm, castaneous with pale margins, L lamina linear-triangular, with pungent tip, appressedly whitish-lepidote on both faces, glabrescent adaxially, margins laxly serrate, Sp 2 mm, brown; Inf to 2.25 m, laxly few-branched; peduncle pale-lepidote, glabrescent; peduncular Bra subfoliaceous (lower) to narrowly triangular from a broadly ovate base (upper), much shorter than the internodes, serrulate; fertile Inf part 1 m, finely cinereous-lepidote, glabrescent; Br suberect, slightly flexuous, slender, elongate; primary Bra like the upper peduncular bracts, much shorter than the sterile bracteate branch bases; floral Bra broadly ovate, apiculate, mostly shorter than the pedicels; Fl laxly arranged, secund, curved-ascending; Ped to 5 mm, slender; Sep broadly ovate, 6–8 mm, obtuse; Pet suberect, broadly obovate, 9 mm or more, colour not described; St included; Fil connate above the common tube; Sty  none; Fr ovoid, acute, 15 mm; Se with a broad rounded subfalcate wing. D. joanae-marcioi P. J. Braun & al. (Bromelie 2008(1): 33–46, ills., 2008). Type: Brazil, Minas Gerais (Esteves Pereira & Braun 493 [UFG, HAL]). — Distr: Brazil (Minas Gerais: Mato Verde); rocky ground, between shrubs and grasses, 900 m. [3] Ros many-leaved, 20–30 cm ∅, 10–15 cm high, usually with some offsets; L 8–17  1–2 cm, 5–15 mm thick, lamina succulent, very hard, stiff, breaking, adaxially flat to concave, apex pungent and brown, both faces grey- to sordidly white-lepidote, margins spinose-serrate, Sp brown, lower ones white-lepidote; Inf to 50 cm but usually shorter, simple, erect to ascending, stout; peduncle reddish, to 32 cm, sparsely white-lepidote; fertile Inf part 12–18 cm, 5- to 10-flowered, reddish, sparsely lepidote; floral Bra to 6 mm, very soon drying, later translucent; Fl erect in bud, then slightly nutant, 15–20  10 mm; Ped 3–5 mm, thick; Sep broadly spatulate, to 13  5–8 mm, basally connate, apical 7–10 mm free, succulent, convex, red to orange, white-lepidote, compressing the perianth, densely covered with thick nectar drops; Pet 16–17  8–10 mm, reddish-orange to dark orange, upper

Dyckia BROMELIACEAE

margins rolled back towards the end of anthesis; St included or shortly exserted; Fil 7–8 mm, free above the common tube (Pinangé & al. 2017: fig. 3); Anth yellow, 3 mm, strongly arching outwards; Ov incl. Sty 10 mm, light yellow; Fr 13  12–15 mm, blackish-brown; Se lentiform, slightly flattened in the hilum-micropylar region, 5–6 mm ∅, light brown. Compared with D. marnier-lapostollei and D. braunii, and very similar. Fruit size has been taken from the table in the protologue, and appears incorrectly given in the description. D. jonesiana Strehl (Bromeliaceae 42(5): 8–9, figs. 1–4, 2009). Type: Brazil, Rio Grande do Sul (Silva & Strehl 1172 [HAS]). — Distr: Brazil (S-C Rio Grande do Sul: Caçapava do Sul); on or between sandstone. [3b] Ros with only 7 leaves, 15 cm ∅, with a bulbous base, rhizomatous and forming sparse groups; L 10–15  1.2–1.5 cm, sheath ovate, 2 cm wide, white, L lamina succulent, triangular-acute, rather recurved, with hard and pungent tip, easily breaking, dark green to copper-coloured, densely white-lepidote esp. abaxially, very laxly serrate, Sp slender, 1–2 mm; Inf 60–80 cm, simple; peduncle erect, 40–60 cm; peduncular Bra triangular, amplexicaul, acute, membranous, longer (lower) or shorter (upper) than the internodes, whitelepidote; fertile Inf part lax, 20 cm, rachis slightly lepidote or glabrous, reddish; floral Bra membranous, ovate, 5–12  11–13 mm, ecarinate or slightly carinate, orange to castaneous, whitelepidote esp. apically; Ped absent or short; Sep narrowly ovate, apex acute or roundish, ecarinate, covered by the bracts, 8–16 mm, orange-reddish, glabrous, margins membranous; Pet free, tongueshaped, 20–24 mm, carinate, obtuse; St included; Fil connate above the 7 mm high common tube; Anth narrowly triangular, recurved; Ov 15 mm; Fr dehiscent to the base, 13–14  10 mm, dark castaneous to nearly black, shining; Se acute at the base, apex rounded, with lateral wing, 4 mm, castaneous. The rather small-sized plants are compared with D. remotiflora in the protologue. — [F. Krapp]

925

D. julianae Strehl (Vidalia 2(2): 2730, figs. 1–4, 2004). Type: Brazil, Rio Grande do Sul (Strehl 1427 [HAS]). — Distr: Brazil (E Rio Grande do Sul); on rock outcrops and stony ground, between grasses. [3a] Ros squarrose, 50- to 80-leaved, 20–30 cm ∅, subcaulescent, offsetting and forming dense clumps; L to 14–15  1.5 cm, sheath 2.5  3 cm, white with brown spots adaxially, both faces glabrous, L lamina narrowly acuminate, recurved, with acute pungent tip, somewhat succulent, pale green, white-lepidote on both faces, margins densely regularly spinose, Sp 2 mm, antrorse, 10–15 mm apart; Inf 80–100 cm, simple or sometimes branched; peduncle 20–30 cm; peduncular Bra almost verticillate near the peduncle base, leaf-like (lowest), 2–5  1–1.5 cm, lamina long-triangular, apex long-acuminate, white-tomentose, margins serrate; fertile Inf part 20–40 cm, axis whitetomentose; floral Bra ovate-triangular, acute, 5–12  5 mm, abaxially puberulous, somewhat carinate, (sub-) entire; Ped absent and Fl sessile, tubular to narrowly funnel-shaped, 16  4 mm; Sep narrowly ovate, 6–8  3–4 mm, ecarinate or slightly carinate towards the apex, basally green, distally brown, slightly white-puberulous abaxially; Pet ovate, 13–16  5 mm, (sub-) erect, yellow; St included; Fil yellow, filiform, free above the common tube; Anth yellow, visible in the throat; Ov 5  3 mm, yellow-green or brown; Sty 6 mm. D. kranziana Leme (Phytotaxa 16: 12–14, ills., 2011). Type: Brazil, Mato Grosso (Kranz 122 [RB, HB]). — Distr: Brazil (Mato Grosso); Cerrado vegetation, on reddish soil or rocks. [1/3b?] Ros dense, 12-leaved, basally offsetting; L 29–32  2–3.5 cm, stiff, slightly succulent, sheath inconspicuous, L lamina narrowly triangular, attenuate and strongly canaliculate towards the apex, suberect-arcuate, acuminate and with pungent tip, green, abaxially densely white-lepidote, adaxially subdensely whitelepidote, subdensely to laxly spinose, Sp narrowly triangular, straight to slightly retrorse, 2.5–5 mm, with castaneous tip, 6–15 mm apart; Inf to 85 cm, simple or sometimes with 3 late-developing branches; peduncle erect, 65 cm, green, sparsely

926

white-lanate to glabrous mainly towards the base; peduncular Bra acuminate, lowest sublinear and canaliculate-carinate and exceeding the internodes, upper ovate-caudate and shorter than the internodes, 18–25  10–12 mm, green, stramineous at anthesis (upper), inconspicuously white-lepidote to glabrous, veined, lowest laxly spinulose, upper densely finely denticulatecrenulate; fertile Inf part densely 38-flowered, 10–30 cm, often deflexed-ascending, greenish to pale orange, densely white-lanate; Br when present spicate, 3–6 cm, densely 5- to 11-flowered, 10–12 cm; primary Bra narrowly lanceolate, acuminate-caudate, white-lepidote, veined; floral Bra ovate, acuminate, 7–15  6–8 mm, stramineous at anthesis, white-lepidote, veined, margins remotely denticulate-crenulate to entire; Fl suberect, 13–14 mm, narrowly ovate with narrow throat, unscented; Ped 2 mm, pale orange, densely white-lanate; Sep broadly ovate, 6–7  6 mm, obtuse-emarginate, ecarinate, strongly convex, yellowish-orange, obscurely white-lanate, entire, sparsely fimbriate; Pet broadly obovatespatulate, 10  8.5 mm, obtuse-emarginate, ecarinate, orange, glabrous; St equalling the petals to slightly exserted; Fil 1 mm connate above the 1 mm high common tube, 7.5–8 mm, pale orange; Anth narrowly subtriangular-sagittate, 3 mm, recurved at anthesis, acuminate; Ov narrowly suboblong, 4 mm; Sty 1 mm, pale yellow, Sti 1 mm, orange, margins scalloped; Fr subglobose, broadly acute and shortly beaked, 15  11 mm, olivaceous-castaneous, lustrous; Se nearly circular, obtuse, strongly flattened, 4–5  3.5–4.5 mm. Similar to D. velascana. D. lagoensis Mez (in Martius, Fl. Bras. 3(3): 483, 1894). Type: Brazil, Minas Gerais (Warming 2171 p.p. [C, F [photo]]). — Distr: Brazil (Minas Gerais); open Cerrado scrub and fields, on soil or on rocks. I: Smith & Downs (1974: 535). [3b] Ros acaulescent; L to 40  1.2–1.6 cm, sheath not described, L lamina narrowly triangular, green, canescent or fuscescent, sublepidote abaxially, laxly serrate, Sp slender, 1–2 mm; Inf to 1 m, simple; peduncle lepidote or finally glabrous; peduncular Bra broadly ovate with long

F. Krapp and U. Eggli

narrowly triangular lamina, equalling or exceeding the internodes, serrulate; fertile Inf part few-flowered, minutely and fugaciously lepidote; floral Bra suberect to spreading, suborbicular, lamina long narrowly triangular, exceeding the sepals and the lowest flowers, fimbriate-serrulate; Fl erect or suberect, to 12 mm; Ped 1–2 mm, stout; Sep ovate, broadly subacute or rounded, mucronulate or obtuse, 6–9 mm, slightly fimbriate; Pet erect, lamina broadly obovate, obtuse, slightly undulate, orange, subcarinate, slightly exceeding the sepals; St included; Fil 7 mm connate above the common tube; Anth narrowly triangular, recurved; Sty very short. D. leptostachya Baker (Gard. Chron., ser. nov. 1884(2): 198, 1884). Type: Brazil, Paraná (Anonymus s.n. [K, GH [photo]]). — Distr: Brazil (Minas Gerais, Santa Catarina, Paraná, Mato Grosso, Rio Grande do Sul), Paraguay (Caaguazú, Itapúa), Bolivia (Chuquisaca, Santa Cruz), Argentina (Chaco, Corrientes, Entre Ríos); dry fields or open scrub (Cerrado vegetation). I: Smith & Downs (1974: 549); Reitz (1983: t. 51, as D. remotiflora var. montevidensis). Incl. Dyckia leptostachia Baker (s.a.) (nom. inval., Art. 61.1); incl. Dyckia boliviensis Mez (1896); incl. Dyckia conspicua Mez (1896); incl. Dyckia hassleri Mez (1903); incl. Dyckia apensis Mez (1919); incl. Dyckia hassleri fa. gracilis Hassler (1919); incl. Dyckia hassleri ssp. basispina Hassler (1919); incl. Dyckia hassleri var. arenosa Hassler (1919); incl. Dyckia hassleri var. montana Hassler (1919); incl. Dyckia longifolia Mez (1919); incl. Dyckia rojasii Mez (1919); incl. Dyckia hassleri var. typica Hassler (1919) (nom. inval., Art. 26.1). [2/3a] Plants very variable as to size; Ros 15leaved; L 40–100  1–3 cm, L lamina arching, narrowly triangular, long-attenuate, densely white-lepidote, soon glabrous adaxially, Sp slender, curved, 3–4.5 mm; Inf 50–150 cm, simple or few-branched; peduncle very slender, minutely lepidote or glabrous; peduncular Bra very broadly ovate, apiculate, much shorter than the internodes, entire; fertile Inf part 12–16 cm, rachis sparsely lepidote or glabrous; floral Bra spreading, very broadly ovate, abruptly contracted to an

Dyckia BROMELIACEAE

acuminate point, 3–8 (–12) mm, much exceeded by the sepals; Fl suberect to spreading, 13–23 mm; Ped very short, stout; Sep broadly ovate, 6–12 mm, obtuse,  carinate, smooth, soon glabrous; Pet erect, broad, obtuse or emarginate, minutely crenulate, red-orange; St usually exserted; Fil free above the common tube; Anth acute, mucronate, recurved; Sty sometimes long or partially divided. D. limae L. B. Smith (Phytologia 20: 179, t. 2, figs. 9–11, 1970). Type: Brazil, Pernambuco (Lima 61-3996 [IPA, US [photo]]). — Distr: Brazil (Pernambuco); in compact nearly barren sandy soil. I: Siqueiro Filho & Leme (2007: 315). [1/2] Ros not described; L 24  1.5 cm, curved, sheath 3.5 cm wide, stramineous to dark brown, densely serrulate at the apex, L lamina narrowly triangular, with pungent tip, appressedly pale-lepidote, laxly serrate, Sp slender, antroretrorsely uncinate, 2 mm, brown; Inf 70–80 cm, depauperately compound with 1 or 2 small suberect branches, or simple; peduncle  straight, glabrous; peduncular Bra except the lowest triangular, remote, small, finely serrulate; fertile Inf part lax, rachis slender, flexuous, compressed, to 25 cm, all parts incl. pedicels and sepals appressedly pale-lepidote; floral Bra ovate, 6 mm, acuminate, slightly exceeding the pedicels (lowest), serrulate (lowest); Fl secund; Ped to 5 mm in fruit, rather stout; Sep ovate, 7 mm, broadly acute and apiculate, sulcate; Pet elliptic-rhomboid, 12 mm, obtuse, carinate, orange; St included; Fil 7 mm, connate for 2–4 mm with the petals but free above the common tube (or completely connate according to Fig. 3 of Pinangé & al. (2017)); Sty 1 mm; Fr subglobose, 15 mm. Similar to D. velascana in general appearance. D. lindevaldae Rauh (Trop. subtrop. Pfl.-welt 65: 12–17, figs. 6 and 7, 1988). Type: Brazil, Goiás (Esteves Pereira s.n. in Rauh 67425 [HEID]). — Distr: Brazil (N Goiás); terrestrial on weathered soil. I: Braun & al. (2008). [3b] Ros many-leaved, 15–20 cm ∅, to 4 cm high, acaulescent or shortly caulescent, solitary or usually in groups, often dividing dichotomously;

927

L to 10  3 cm, sheath to 1.5  3.5 cm, greenishwhite, castaneous at the base abaxially, glabrous, entire, L lamina narrowly triangular, acuminate, pungent, slightly succulent, strongly reflexed, both faces very densely tomentose (easily wiped away) giving a snow-white overall colour, serrate, Sp retrorse, to 4 mm, yellowish-brown, lanate, 5 mm apart; Inf to 30 cm, simple; peduncle slender, curved-ascending, to 15 cm, brownishgreen, glabrous; peduncular Bra subfoliaceous (lower) or smaller (upper), oblong, triangular, acuminate, shorter than the internodes, glabrous at the broadened base, elsewhere sparsely lepidote, soon drying; fertile Inf part spirally 12flowered, rachis slender, flexuous, to 20 cm, reddish-brown, sparsely lepidote; floral Bra spoon-shaped, acute, carinate, 7 mm, equalling or slightly shorter than the sepals, brownishgreen, initially sparsely lepidote, later glabrous, veined, slightly serrate; Fl erect-spreading, 10–12 mm, subsessile; Sep ecarinate, strongly convex, obtuse, 6–7  4 mm, nearly glabrous, margins and tip ciliate; Pet broadly round to rhombic, 9–11 mm, erect, ecarinate, obtuse, pale orange-red, yellow towards the base, ciliate at the tip; St slightly exserted; Fil very shortly connate, white; Ov cylindrical, polygonally sulcate, 4 mm; Sty short, yellow, Sti shortly papillate; Fr and Se not described. Similar to D. marnier-lapostollei var. marnierlapostollei. D. linearifolia Baker (Handb. Bromel., 131, 1889). Type: Brazil, Minas Gerais (Saint Hilaire 1010 [P, GH [photo]]). — Distr: Brazil (Minas Gerais, São Paulo); open scrub, savanna. [3a] Ros not described; L 40–55  1 cm, sheath suborbicular, 5 cm, dark castaneous, L lamina sublinear, strongly appressedly brown- or whitishlepidote abaxially, laxly serrate towards the base, Sp slender, curved, 1.5 mm; Inf 40 cm and more, simple; peduncle very slender, glabrous; peduncular Bra ovate, acuminate, entire, remote; fertile Inf part subdense, many-flowered, completely glabrous; floral Bra spreading to reflexed, triangular-ovate, exceeding the pedicels but much shorter than the sepals; Fl spreading at anthesis, then erect; Ped to 7 mm, rather slender;

928

Sep broadly ovate, to 8 mm, obtuse; Pet lamina suberect, narrowly elliptic, 15 mm, obtuse, carinate, colour not described; St included; Fil free above the common tube; Anth linear, mucronulate, nearly straight; Ov not described; Sty  as long as the ovary; Fr and Se not described. — [F. Krapp] D. lunaris Leme (Selbyana 30(2): 131–133, fig. 2, 2010). Type: Brazil, Goiás (Leme 6359 [HB, RB]). — Distr: Brazil (Goiás); ecology not recorded. [3b] Ros dense, 30-leaved, with elongated basal shoots; L 29–40  2.7–3.4 cm, strongly coriaceous, succulent, sheath wider than the lamina, L lamina narrowly triangular,  flat to slightly canaliculate, strongly recurved, apex long acuminate, subulate and pungent, green, sometimes yellowish-green basally and apically, adaxially glabrous, abaxially veined and inconspicuously white-lepidote, margins laxly spinose, Sp narrowly triangular,  straight to antrorsely uncinate, 6–9 mm, glabrous, greenish to reddish with yellowish tip, 10–40 mm apart; Inf 55–90 cm, simple, erect; peduncle 25–45 cm, green, subdensely white-lanate to glabrescent; peduncular Bra foliaceous to subfoliaceous, strongly recurved to suberect, upper ones long acuminate-caudate from a subtriangular-ovate base, distinctly exceeding the internodes, yellowish-green with red apex, upper ones stramineous at anthesis, densely veined, glabrous, margins laxly spinose; fertile Inf part 50- to 80-flowered, subdense to dense, 22–45 cm, rachis straight, (sub-) densely white-lanate; floral Bra long acuminate-caudate from an ovate base (lowest) to subtriangular-ovate and acuminate (upper), spreading to suberect-ascending, ecarinate, 30–55  10–14 mm (lowest) to 12–30  7–10 mm (upper), stramineous at anthesis, glabrous to inconspicuously white-lepidote, distinctly veined, subdensely and minutely spinulose; Fl spreading to suberect at anthesis, becoming erect afterwards, 19–23 mm, campanulate, unscented; Ped 3–7 mm, stout, orange, densely white-lanate; Sep ecarinate,  acute, adaxial ones asymmetrical-curved, subtriangular-ovate, abaxial one symmetrical, ovate, distinctly convex, 10–11  5–6 mm, orange to

F. Krapp and U. Eggli

reddish, subobscurely white-lanate, entire, fimbriate; Pet broadly obovate, 11–13  7 mm, apex rounded to cucullate, ecarinate, orange, abaxially subdensely white-lepidote; St included; Fil 4–5 mm connate above the 2 mm high common tube, 6–7 mm, apex pale orange; Anth ovate, 3 mm, strongly spirally recurved at anthesis, base sagittate, acute; Ov narrowly subpyramidal, 5.5 mm, yellowish; Sty 2 mm, Sti conduplicatespiral, 1.5 mm, orange, minutely crenulate; Fr broadly ellipsoid-ovoid, shortly beaked, 14–15  9–10 mm, dark castaneous, lustrous; Se flat, asymmetrical, subcuneate, base obtuse, 4–5  3 mm. Similar to D. burle-marxii. The only other species with asymmetrical sepals (and thus somewhat zygomorphic flowers) is D. pontesii. D. lutziana L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 107, t. 107, 1943). Type: Brazil, sine loco (Foster 1144-B [GH]). — Distr: Brazil (without recorded locality); never recollected. I: Smith & Downs (1974: 576). [3b] Ros not described; L incompletely known, inner leaves to 10 cm, lamina linear, appressedly pale-lepidote on both faces, laxly serrulate; Inf to 80 cm, simple, very slender; peduncle glabrous; peduncular Bra remote, broadly ovate, acuminate, membranous, small, reddish; fertile Inf part lax, straight, to 9 cm, subglabrous; floral Bra like the peduncular bracts but apiculate, shorter than to barely exceeding the pedicels; Fl suberect to spreading; Ped 4–5 mm, slender; Sep elliptic, 8–9 mm, obtuse; Pet lamina broad, to 12 mm, orange-yellow when dry; St slightly exserted; Fil high-connate above the common tube; Ov not described; Sty slender, 2 mm. D. macedoi L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 2: 195, 1952). Type: Brazil, Minas Gerais (Macedo 2974 [US]). — Distr: Brazil (Minas Gerais); rocky fields. [3a] Ros spreading, with a stout probably short stem; L 6–8  1.4 cm, suberect-recurved, sheath broad, >3 cm wide, L lamina narrowly triangular, with pungent tip, adaxially soon glabrous, abaxially appressedly cinereous-lepidote and inconspicuously veined, margins laxly

Dyckia BROMELIACEAE

serrate, Sp nearly straight, 1.5 mm, dark; Inf 30 cm, simple; peduncle very slender, glabrous; peduncular Bra broadly ovate, acuminate, shorter than the internodes, 7 mm (uppermost), serrulate; fertile Inf part laxly 10-flowered, 4.5–5.5 cm, glabrous; floral Bra like the upper peduncular bracts,  equalling the pedicels; Fl spreading or reflexed; Ped 5 mm, slender; Sep oblong-elliptic, 5 mm, broadly obtuse, strongly convex; Pet shape not described, 7 mm, reddish drying to goldenyellow; St included; Fil free above the 0.5 mm high common tube; Sty  none. D. nana is compared with this species in its protologue. D. machrisiana L. B. Smith (Contr. Sci. Nat. Hist. Mus. Los Angeles County 17: 7, fig. 5, 1957). Type: Brazil, Goiás (Dawson 14153-A [R, US [photo]]). — Distr: Brazil (Goiás); fields and open Cerrado scrub. [3a] Ros not described; L 25  1.5 cm, sheath suborbicular, 3 cm wide, glabrous, L lamina linear-triangular, appressedly white-lepidote on both faces, margins laxly serrulate, Sp acicular, 1 mm; Inf 80 cm, simple; peduncle size not described, sparsely pale-lepidote; peduncular Bra broadly ovate, thin, abruptly contracted into a linear-triangular apex, all but the lowest much shorter than the internodes; fertile Inf part lax, slender, flexuose, sparsely pale-lepidote; floral Bra broadly ovate, acuminate, to 6 mm; Fl spreading or divergent; Ped 2–4 mm, stout; Sep ovate, 7 mm, obtuse, ecarinate, rather fleshy; Pet erect, elliptic, 10 mm, ecarinate, dark blackish, orange externally; St included; Fil free above the short common tube; Ov not described; Sty  none. — [F. Krapp] D. macropoda L. B. Smith (Phytologia 14: 485, t. 1, figs. 25–28, 1967). Type: Brazil, Minas Gerais (Pereira 1622 p.p. [RB, US [photo]]). — Distr: Brazil (Minas Gerais: Diamantina region); rocky fields. I: Smith & Downs (1974: 568). [3b] Ros not described; L 25  1.5 cm, sheath suborbicular, 3 cm, stramineous, L lamina narrowly triangular, attenuate to a fine rigid point, smooth and glabrous adaxially, appressedly

929

whitish-lepidote abaxially (appearing ferrugineous from soil deposits), laxly serrate, Sp retrorsely uncinate, 1.5 mm; Inf 50 cm, simple; peduncle very short, finely ferrugineous-lepidote; peduncular Bra long-acuminate from an ovate base, much shorter than the internodes (uppermost), subentire; fertile Inf part laxly many-flowered, 25 cm, all parts incl. pedicels and sepals finely ferrugineous-lepidote; floral Bra like the peduncular bracts, exceeding the sepals (lowest), entire; Fl spreading; Ped to 10 mm, slenderly cylindrical; Sep broadly ovate, 7 mm, rounded or apiculate, strongly convex, the posterior  carinate; Pet broadly rhomboid, 14 mm, carinate, orange; St included, Fil almost wholly connate; Ov not described; Sty very short. — [F. Krapp] D. maracasensis Ule (Bot. Jahrb. Syst. 42: 197–198, 1909). Type: Brazil, Bahia (Ule 7019 [B, F [photo]]). — Distr: Brazil (Bahia: Maracás region); rocks. I: Smith & Downs (1974: 542). [1] Ros dense; L to 20 cm, sheath suborbicular, short, L lamina narrowly triangular, attenuate, with pungent tip, densely lepidote abaxially, margins laxly serrate, Sp curved, 2 mm; Inf 1 m or more, racemose or paniculate, or simple; peduncle subangular, densely ferrugineous-flocculose towards the apex; peduncular Bra erect, lanceolate-ovate with long, narrowly triangular, serrulate lamina, foliaceous and imbricate (lower) to shorter than the internodes (upper); fertile Inf part ferrugineous-flocculose; primary Bra like the upper peduncular bracts, much shorter than the long compressed bases of the branches; floral Bra broadly ovate, apiculate, slightly exceeding the pedicels, finely serrate; Fl suberect or sometimes spreading and secund; Ped 5 mm, stout; Sep broadly ovate, 6 mm, obtuse; Pet broadly elliptic, 9 mm, obtuse, yellow; Fil 2 mm connate (or completely connate according to Fig. 3 of Pinangé & al. (2017)) above the common tube; Ov 4.5 mm; Sty 1.5 mm. — [F. Krapp] D. maritima Baker (Handb. Bromel., 136, 1889). Type: Brazil, Rio Grande do Sul (Tweedie s.n. [K, GH [photo]]). — Lit: Waldemar & Irgang (2003: ecology). Distr: Brazil (S Santa Catarina, E Rio Grande do Sul), open rocky or turfy ground,

930

mostly near the coast. I: Smith & Downs (1974: 510); Reitz (1983: t. 47); Klein V. & Klein (2014: 76–77).  Prionophyllum maritimum (Baker) Mez (1896); incl. Dyckia tomentosa Mez (1896). [1] Ros spreading, many-leaved; L 60 to 4 cm, thick, rigid, sheath suborbicular, dark castaneous, glabrous, L lamina narrowly triangular, attenuate, with pungent tip, glabrous adaxially, minutely pale-lepidote abaxially, conspicuously veined, margins very laxly serrate, Sp broad, curved, 3–5 mm; Inf to 2 m and more, 3 branched; peduncle straight, glabrous; peduncular Bra imbricate, foliaceous (lower) to narrowly triangular (upper), much reduced; fertile Inf part densely tomentose-lepidote; Br broadly spreading, many-flowered, subdense, to 25 cm; floral Bra spreading to reflexed, subulate-acuminate from a broad base, scarcely 3 mm; Fl perfect and unisexual, suberect to spreading; Ped very short; Sep broadly ovate, 5 mm, acute, greenish, densely brown-furfuraceous; Pet obtuse, 7–8 mm, yellow; St hardly exserted; Fil free above the very short common tube; Ov pryamidal, 3-sulcate; Sty longer than the ovary; Fr 5 mm, ovoid, brown to almost black, dehiscing to below 1=2 , shortly rostrate; Se oblong with a narrow lateral wing widened at the ends. Belongs to the “Prionophyllum” complex. A naturally-occurring variegated plant was recently described as cultivar ‘Leopoldo Witeck’ (Lawn 2011). Waldemar & Irgang (2003) found a facultative association of the species with termites, and it grows esp. well on termite mounds. D. marnier-lapostollei L. B. Smith (Bromeliad Soc. Bull. 16: 102, ills., 1966). Type: Brazil, Goiás? (Anonymus ex cult. Marnier-Lapostolle s.n. [US]). — Lit: Rauh (1985: 51–58); Braun & Esteves Pereira (2005). Distr: Brazil (SE Goiás). [3a] Ros acaulescent, spreading, 10-leaved, usually solitary or occasionally in small groups; L 12  4 cm, sheath suborbicular, 2 cm, glabrous, L lamina triangular, thick, strongly recurved, densely pale-lepidote on both faces and with silvery appearance, margins densely serrate, Sp coarse, retrorsely curved; Inf 50–100 cm, simple, erect; peduncle compressed, very slender,

F. Krapp and U. Eggli

glabrous; peduncular Bra broadly ovate, apiculate, small, entire, remote; fertile Inf part lax, few-flowered, slender, slightly flexuous, 19 cm, nearly glabrous; floral Bra broadly ovate, acuminate, shorter than the sepals, entire; Fl subsessile, spreading, 12 mm; Sep broadly ovate, 7 mm, rounded, cucullate, ecarinate; Pet subrhomboid, 12 mm, limb only slightly distinct from the claw, obtusely carinate, cucullate, orange; St equalling the petals; Fil free above the common tube; Ov not described; Sty  none. The description has been amended from data provided by Braun & Esteves Pereira (2005). These authors report hybrids with D. machrisii, with  intermediate characters. D. marnier-lapostollei var. estevesii Rauh (Trop. subtrop. Pfl.-welt 53: 55–56, figs. 26–28, 1985). Type: Brazil, Goiás (Esteves Pereira s.n. in Horst 5 [HEID]). — Distr: Brazil (E to SE Goiás); rocks, in Campo Rupestre vegetation. – Fig. 7. [3a] L less densely lepidote, bluish-grey, partly glabrous, Sp to 6 mm; Inf to 80 cm, very slender; Fl scent less pronounced; Anth visible at anthesis. D. marnier-lapostollei var. marnier-lapostollei — Distr: Brazil (E to SE Goiás); rocks, in Campo Rupestre vegetation, 1000 m and more. I: Rauh (1985: 51–58); Braun & Esteves Pereira (2005); Braun (2018: 72). Incl. Dyckia rauhii hort. (s.a.) (nom. inval., Art. 29.1). [3a] L densely white-lepidote, silvery, Sp to 3 mm; Inf to 50 cm; Fl scent strong, fish-like; Anth included at anthesis. D. martinellii B. R. Silva & Forzza (Novon 14: 168–170, ills., 2004). Type: Brazil, Rio de Janeiro (Martinelli 14413 [RB]). — Distr: Brazil (Rio de Janeiro: Paratimirim); granitic outcrops; narrowly endemic. [3b] Ros 60 cm ∅, with stout basal offshoots; L 23–38  2.2–2.6 cm, suberect-arcuate, sheath elliptic, 2.5–3  5–6 cm, castaneous abaxially, whitish adaxially,  entire, L lamina lanceolate, apex attenuate, whitish abaxially, light green adaxially, spinose, Sp straight, patent-

Dyckia BROMELIACEAE

931

Fig. 7 Dyckia marnierlapostollei var. estevesii. (Copyright: U. Eggli)

antrorse, 3–4 mm, light green with a castaneous apex, 10–15 mm apart; Inf 70–100 cm, simple; peduncle erect, 55–70 cm (to 97 cm according to the protologue), green, sparsely white-lepidote; peduncular Bra subfoliaceous (lowest) to lanceolate-attenuate (upper), densely arranged, clasping the peduncle, erect, chartaceous, 11–12.5  0.9–1.2 mm (lowest), exceeding to rarely equalling the internodes (upper), stramineous, glabrous to sparsely lepidote, spinulose to inconspicuously serrulate; fertile Inf part 38- to 65-flowered, lax, straight, 30–43 cm, orange, sparsely white-lepidote; floral Bra elliptic to elliptic-attenuate, acuminate, exceeding to equalling the middle of the sepals, 4–12  3–6 mm, castaneous, white-lepidote, entire; Fl patent at anthesis, suberect afterwards; Ped 1–2 mm, to 4 mm in fruit, cylindrical, orange, sparsely lepidote; Sep ovate, 7–9  4–6 mm, apex rounded, orange, apex castaneous, whitelepidote; Pet obtrullate, 9–12  9 mm, apex emarginate, erect, orange, glabrous; St included; Fil 1.5 mm connate (completely connate according to Fig. 3 of Pinangé & al. (2017)) above the 2.5 mm high common tube; Anth slightly sagittate at the base, attenuate and reflexed at the apex, 3 mm; Ov 7 mm; Fr 15 mm; Se alate, 3 mm.

Similar to D. pseudococcinea, and compared with D. espiritosantensis in the protologue of the latter. D. mauriziae Esteves & Hofacker (Bromelie 2011(1): 39–42, ills., 2011). Type: Brazil, Goiás (Carneiro MBR-41 [UFG]). — Distr: Brazil (Goiás); steep slopes in yellow gravelly latosol. [3b] Ros solitary or in small groups, with secund prostrate leaves, stem to 20  5 cm; L 39  3.7 cm, inner ones straight, soon curved, coriaceous, rigid, succulent, margins narrow, sheath 2.6  4.8 cm, white-yellowish, canescent-lepidote, L lamina flat, occasionally slightly concave, apex long-attenuate with a hard, pungent, light-brown, 9 mm long tip, greenishgrey to pale green, both faces finely veined and with canescent scales, margins greenish-brown, serrate, Sp hard, pungent, antroretrorse-uncinate, to 4.8 mm, 22 mm apart; Inf to 1.8 m, usually simple, in cultivation rarely paniculate, stout, erect; peduncle to 110 cm, pale greyish- or brownish-green, sparsely light-brown-lepidote; peduncular Bra linear to lanceolate, suberect or curved, foliaceous, long attenuate, acuminate, pungent, 130  17 mm (lower) to 34  13 mm (uppermost), pale green to greenish-brown, canescent-lepidote, finely veined, laxly spinose, teeth

932

0.3–1.8 mm; fertile Inf part 90 cm, reddish-rose, densely 20-flowered, conspicuously furrowed, with creamy papillae-like trichomes; floral Bra like the peduncular bracts but shorter, narrowly triangular, 19  8 mm (lowest) to 4.8  3.8 mm, pale orange, soon stramineous, with dense colourless papillae-like trichomes, finely veined, with ciliate margins; Fl pointing slightly sidewards, after pollination directed upwards, 21 mm, smaller towards the inflorescence tip, fragrant with slight sour scent; Ped 3.7 mm, red; Sep lanceolate to ovate, 9.5  6.5 mm, succulent, carinate, bright orange-reddish, with papillae-like trichomes; Pet oblong, 11.6  7.2 mm, obtuse, rigid, carinate apically, orange to orange-red, brilliant; St just visible in the throat; Fil 4–6 mm connate above the common tube, 9–11 mm, cream-coloured; Anth yellow, 4.6  1.3 mm, recurved at anthesis; Ov narrowly subpyramidal, 7  3 mm, whitish-yellow; Sty 2.3 mm, yellow; Sti fringe-like, yellow, 1.6 mm; Fr ellipsoid to ovoid, acuminate, 12–15  9–11 mm, dark brownish, lustrous. Similar to D. estevesii due to the fan-like and seemingly distichous leaf arrangement, and also compared with D. mirandana in the protologue. D. mello-barretoi L. B. Smith (Phytologia 7: 109, t. 1, figs. 16–19, 1960). Type: Brazil, Minas Gerais (Mello Barreto 2122 [BHMG, US [photo]]). — Distr: Brazil (Minas Gerais); sand and rocky fields. I: Smith & Downs (1974: 549). [2/3a?] Ros not described; L >30  1.5 cm, sheath not described, L lamina narrowly triangular, attenuate to an abruptly acute pungent tip, pale-lepidote between the veins, margins very laxly serrate, Sp slender, spreading, retrorsely curved, 4 mm; Inf probably >1 m, subsimple to compound; peduncle inconspicuously palelepidote at the nodes; peduncular Bra ovate, acuminate, the upper remote; fertile Inf part with a short 2-flowered Br at the base, lax, flexuous, appressedly stellately ferrugineous-lepidote, rachis soon  glabrous; floral Bra reflexed, ovate and acuminate (lower) or suborbicular and apiculate (upper), 11 mm, nearly equalling (lower) to as long (upper) as the sepals; Fl spreading; Ped 3 mm, stout; Sep broadly ovate,

F. Krapp and U. Eggli

8 mm, obtuse, erose; Pet lamina broadly obovate, to 13 mm, emarginate, ecarinate, colour not described; St included; Fil free above the common tube; Anth narrowly triangular, nearly straight, 3 mm; Ov not described; Sty simple, 2 mm. Similar to D. sordida, and compared with D. espiritosantensis in the protologue of the latter. — [F. Krapp] D. mezii Krapp (Ann. Bot. Fenn. 50(1): 73, 2013). Type: Brazil, Minas Gerais (Glaziou 17280-A [B, F [photo]]). — Distr: Brazil (Minas Gerais); ecology not recorded, not recently recollected.  Dyckia argentea Mez (1894) (nom. illeg., Art. 53.1). [3a] Ros not described; L to 30  2 cm, sheath suborbicular, large, L lamina narrowly triangular, abruptly acute, with pungent tip, coarsely lustrous-silvery-lepidote, laxly serrate, Sp coarse, uncinate, 3.5 mm; Inf 50–70 cm, simple; peduncle  furfuraceous towards the base, glabrous elsewhere; peduncular Bra ovate with acuminate tip, shorter than the internodes, serrulate; fertile Inf part sublax, many-flowered, glabrous; floral Bra lanceolate-triangular (lower) or ovate and acute (upper), exceeded by the sepals; Fl spreading at anthesis, later erect, 11 mm; Ped to 5 mm; Sep elliptic, 6 mm, obscurely apiculate; Pet erect, cuneiform-obovate, broadly rounded, scarcely or not at all carinate, orange; St included; Fil free above the common tube; Anth linear, acuminate, strongly recurved; Sty slightly shorter than the ovary. See Braun & al. (2008) for a discussion of the name (as D. argentea) and the possible origin of the material on which it based. D. microcalyx Baker (Handb. Bromel., 133, 1889). Type: Paraguay (Balansa 696 [P, GH [photo]]). — Distr: S Brazil, Paraguay, NE Argentina; dry rocky slopes and river banks, seasonally inundated. [1/2] Ros not described; L 20–150  1.5–3 cm, sheath scarcely wider than the lamina, L lamina narrowly triangular, pale-lepidote esp. abaxially, often glabrescent, typically subdensely

Dyckia BROMELIACEAE

serrate but varying to entire, Sp 6 mm; Inf 40–200 cm, few-branched or rarely simple; peduncle erect, slender, glabrescent; peduncular Bra ovate with long acuminate pungent lamina, shorter than the internodes (at least the uppermost); fertile Inf part many-flowered, to 80 cm, soon glabrous, Br spreading, densely flowered, 6–26 cm; primary Bra like the peduncular bracts, small, usually shorter than the sterile branch bases; floral Bra broadly ovate or suborbicular, apiculate, 2 mm, much exceeded by the sepals; Fl 6–13 mm, subsessile; Sep suborbicular, 3–6 mm; Pet obovate, suberect, ecarinate, orange; St exserted or  equalling the petals; Fil free above the short common tube; Sty  equalling the ovary. — [F. Krapp] D. microcalyx var. microcalyx — Distr: S Brazil (Mato Grosso, Paraná), Paraguay; dry rocky slopes. I: Rauh & Gross (1991: 9). Incl. Dyckia microcalyx var. inermis Hassler (1919); incl. Dyckia microcalyx var. micrantha Hassler (1919); incl. Dyckia minutiflora Mez (1919). [1/2] L to 150 cm, lamina relatively long and narrow; Inf mostly compound; Sep 3–4 mm. — [F. Krapp] D. microcalyx var. ostenii L. B. Smith (Contr. Gray Herb. 104: 73, t. 3, fig. 16, 1934). Type: Argentina, Misiones (Osten & Rojas 8097 [MVM, GH [photo]]). — Distr: N Argentina (Misiones); rocky banks; only known from the type locality. I: Smith & Downs (1974: 549). [2] L scarcely over 20 cm, lamina 3 cm wide, strongly spinose-serrate; Inf simple; Sep 6 mm. — [F. Krapp] D. milagrensis Leme (Harvard Pap. Bot. 4(1): 164–166, fig. 21, 1999). Type: Brazil, Bahia (Nahoum s.n. in Leme 3391 [HB]). — Distr: Brazil (Bahia); Campo Rupestre vegetation, on rocks. – Fig. 8. [2/3a] Ros propagating by basal shoots, dense, many-leaved; L 12–15  1–1.5 cm, thickly succulent, sheath not described, L lamina very narrowly triangular, subulate-attenuate, spinescentacuminate with pungent tip, suberect to spreading,

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purplish-green, obscurely white-lepidote on both faces, partially glabrescent adaxially, veined, laxly spinose, Sp opposite, narrowly triangular,  straight, flat, 4–6 mm, densely white-lepidote; Inf 120 cm, simple or with 1 or 2 branches; peduncle erect, 70 cm, ferrugineous, inconspicuously white-furfuraceous to glabrescent; peduncular Bra foliaceous (lowest) to narrowly triangular and acuminate, erect, 12–20  4–6 mm (upper), white-lepidote, veined, laxly spinulose; fertile Inf part 45 cm, 46-flowered, subdense, suberect, subdensely white-furfuraceous; Br 10–15 cm, 4- to 7-flowered,  red; primary Bra like the upper peduncular bracts; floral Bra triangular, acuminate, 4–10  3–4 mm, white-furfuraceous, veined, entire; Fl subspreading at anthesis, erect afterwards, the upper slightly secund, 15–16 mm, unscented; Ped 4–5 mm; Sep broadly ovate, 6  6 mm, acute, ecarinate, reddish-orange, subdensely white-furfuraceous, entire with longfimbriate apex; Pet broadly obovate-spatulate, 11  4 mm, rounded, ecarinate, suberect, orange, glabrous; St just included; Fil free above the 2 mm high common tube (completely connate according to Fig. 3 in Pinangé & al. (2017)), complanate, 6 mm; Anth 2 mm,  straight, base shortly sagittate, apex obtuse; Ov narrowly subpyramidal, 6 mm; Sty 8 mm,  equalling the anthers, Sti orange. Similar to D. mezii and D. pectinata. — [F. Krapp] D. minarum Mez (in Martius, Fl. Bras. 3(3): 483, t. 91, 1894). Type: Brazil, Minas Gerais (Regnell II-283 [S, US]). — Distr: Brazil (Goiás, Minas Gerais, Santa Catarina, São Paulo); on rocks or open rocky ground, 900–1300 m. I: Reitz (1983, t. 49). [3b] Ros acaulescent, dense, with bulbous base; L numerous, irregularly spreading, sheath suborbicular, large, pale brown, glabrous but apically lepidote, L lamina 20–35  1 cm, lineartriangular, abruptly acute with thickened pungent tip, green to brownish-green, canaliculate, rigid, thick, densely pale-lepidote esp. abaxially, laxly serrate, Sp curved, 2 mm; Inf 25–75 cm, simple, erect; peduncle rather stout, tomentulose, glabrescent; peduncular Bra broadly ovate with

934

F. Krapp and U. Eggli

Fig. 8 Dyckia milagrensis. (Copyright: F. Krapp)

long acuminate tip, equalling or exceeding the internodes, serrulate towards the apex; fertile Inf part usually lax at anthesis, 30- to 40-flowered, 7–35 cm, white-furfuraceous; floral Bra spreading or reflexed, like the peduncular bracts, equalling or exceeding the flowers (lowest), usually serrulate; Fl  spreading; Ped to 4 mm; Sep broadly elliptic, 7–9 mm, acute, densely lanatelepidote; Pet suborbicular, 11–14 mm, carinate, suberect, orange-yellow; St included; Fil 2 mm connate above the common tube; Anth narrowly triangular, acuminate, recurved; Ov massively pyramidal-trigonous; Sty very short; Fr trigonous-ellipsoid, 14 mm, almost black, glossy; Se 4 mm, basally pointed, alate towards the tip. D. mirandana Leme & Z. J. G. Miranda (J. Bromeliad Soc. 59(2): 75–78, ills., 2009). Type: Brazil, Goiás (Miranda s.n. in Leme 6380 [HB]). — Distr: Brazil (Goiás); sandy soil accumulating among quartzitic outcrops, Campo Rupestre vegetation. [3a] Ros  distichously 10- to 15-leaved, solitary or scarcely offsetting; L 28–43  1 cm, coriaceous, sheath suborbicular-ovate, 3.4  4–5 cm, dark castaneous, lepidote, L lamina sublinear-attenuate, distinctly canaliculate, suberect-arcuate, apex long acuminate-caudate, pungent, green to reddish towards the apex,

abaxially inconspicuously white-lepidote, adaxially glabrescent towards the apex, distinctly veined on both faces, margins glabrous, sparsely spinose, Sp uncinate, antroretrorse, 0.5–1 mm, castaneous, glabrous, 15 mm apart; Inf 20–25 cm, simple; peduncle erect with curved apex, 8 cm, dark purplish wine-red to blackish, glabrous; peduncular Bra erect, subcarinate, narrowly triangular, long acuminate-caudate, 12–18  6–8 mm, exceeding to equalling the internodes, dark purplish wine-coloured, sparsely to densely whitelepidote towards the apex, veined; fertile Inf part 9-flowered, deflected but rachis straight, 6.5 cm, dark purplish wine-red to blackish, glabrous; floral Bra subtriangular, acuminate, subcarinate, membranous, suberect, 7–10  5–6 mm, dark purplish wine-red, sparsely white-lepidote, veined, inconspicuously denticulate to entire; Fl suberect, 12 mm, tubular, strongly sweetfragrant; Ped 1.5 mm, stout; Sep acuminate, convex, subcarinate, 6–6.5  3.5–4 mm, dark purplish wine-red, glabrous, entire; Pet broadly obovate-spatulate from a narrower base, 10  7.5 mm, apex obtuse and apiculate to subacute, ecarinate, yellow; St exserted; Fil free above the 1.5–3 mm high common tube (according to the protologue, completely connate according to Fig. 3 of Pinangé & al. (2017)), yellow, 8 mm; Anth narrowly subtriangular, 2 mm, base

Dyckia BROMELIACEAE

distinctly sagittate, acute, apically strongly recurved; Ov ovoid, 4  2 mm, greenish; Sty 1 mm, yellowish, Sti subentire, yellow; Fr and Se not described. — [F. Krapp] D. mitis A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 36: 51, t. 2, 1929). Type: Argentina, Misiones (Spegazzini s.n. [BA]). — Distr: N Argentina (Misiones); ecology not recorded. I: Smith & Downs (1974: 570). [3b] Ros not described; L 18  1 cm, lamina attenuate, with pungent tip, laxly serrate, Sp curved, 3 mm, 4–6 mm apart; peduncle slender, 27 cm, glabrous; Inf 30 cm, simple; peduncular Bra amplexicaul at the base, shorter than the internodes; fertile Inf part 5–10 cm, rachis furfuraceous; floral Bra suborbicular, apiculate, shorter than the sepals, veined, lepidote; Fl sessile; Sep suborbicular, 6  8–10 mm, broadly rounded; Pet rhombic, 14 mm, rounded at the apex, erect, 14 mm, colour not described; St slightly exserted; Fil highly connate above the 3 mm high common tube; Anth slightly recurved, 3–3.5 mm; Ov pyramidal, 5 mm; Sty 6 mm. Insufficiently known. — [F. Krapp] D. montezumensis Leme (Phytotaxa 67: 23–25, ills., 2012). Type: Brazil, Minas Gerais (Leme & Oliveira 8315 [RB]). — Distr: Brazil (Minas Gerais); shrubby Cerrado vegetation. [2/3b] Ros laxly 10-leaved; L 40–53  2.2–2.5 cm, thickly coriaceous, sheath suborbicular, 2.5  4.5 cm, whitish towards the base, glabrous, L lamina narrowly triangular, distinctly canaliculate, arcuate, apex long acuminatecaudate, dark green to reddish wine-coloured, densely white-lepidote (esp. abaxially), distinctly veined, margins sparsely white-lepidote to glabrous, sparsely spinose, Sp narrowly triangular, antroretrorse, 1.5–2.5 mm, dark castaneous, glabrous, 7–20 mm apart; Inf to 155 cm, simple or branched with 1 branch at the base; peduncle 72 cm, dark green to dark purplish wine-red, inconspicuously white-lepidote to glabrous, smooth; peduncular Bra broadly ovate to suborbicular at the base, narrowly sublineartriangular, carinate, acuminate-caudate, erect,

935

10–40  7–10 mm, much shorter than the internodes, stramineous, inconspicuously whitelepidote, strongly veined, remotely spinulose (lowest) to entire (upper); fertile Inf part 25flowered, lax to subdense, distichous to subpolystichous, rachis straight to flexuous towards the apex, 35 cm, dark purplish wine-red, subdensely white-lepidote; Br suberect, 4-flowered,  polystichous, 7.5 cm, rachis flexuous, sterile base 4 cm; primary Bra like the basal floral bracts; floral Bra broadly ovate, acute to acuminate, ecarinate, 3–7  2–4.5 mm, inconspicuously white-lepidote, veined, entire; Fl subspreading, 16–18 mm, campanulate; Ped 4 mm, dark purplish wine-red to nigrescent, subdensely white-lepidote; Sep ovate, 6  5 mm, distinctly convex, apex obtuse to often emarginate, blackish wine-red, white-lepidote with glabrous apex; Pet broadly spatulate, 13  9–11 mm, obtuseemarginate, ecarinate, orange, apex wine-colored; St deeply inserted; Fil completely connate above the 2 mm high common tube, 9 mm, orange; Anth sublinear, 4 mm, base bilobed, apiculate; Ov suboblong-ovoid, 7–8  2.5 mm, greenish; Sty 1 mm, yellowish, Sti papillose, orange; Fr and Se unknown. Closely related to D. atratiflora. D. monticola L. B. Smith & Reitz (Sellowia 14: 104, fig. 3, 1962). Type: Brazil, Santa Catarina (Reitz & Klein 4789 [US, HBR]). — Distr: Brazil (Minas Gerais, Santa Catarina); open slopes, savanna, on rocks, 1250–1500 m. I: Smith & Downs (1974: 519). [1/2?] Ros dense, many-leaved; L 23–27  1–1.3 cm,  curved, sheath 3 cm wide, brown, lustrous and lepidote towards the apex adaxially, dull and glabrous elsewhere, L lamina linear, rigid, canaliculate, acute and mucronulate, appressedly cinereous-lepidote, soon glabrous adaxially, margins laxly serrate, Sp slender, antrorsely curved, 2.5 mm, brown; Inf 70 cm, with 6 basal branches (according to the protologue, simple according to Fig. 5 in Pinangé & al. (2017)); peduncle subterete, 40 cm, very dark when dry, flocculose, glabrescent; peduncular Bra broadly ovate, foliaceous-laminate or the uppermost acuminate,  equalling the upper

936

internodes, cinereous-lepidote, serrulate; fertile Inf part 28 cm, many-flowered, completely densely ferrugineous-flocculose including pedicel and sepals; Br few-flowered with long sterile base; primary Bra like the upper peduncular bracts, to 17 mm; floral Bra acuminate from a broadly ovate base, spreading, slightly shorter than the sepals, serrulate; Fl spreading, to 15 mm; Ped 5 mm; Sep ovate-elliptic, to 10 mm, acute, incurved, rugose when dry, serrulate; Pet distinctly unguiculate, obovate, curved-spreading towards the apex, orange; St included; Fil 3 mm connate (completely connate according to Fig. 3 in Pinangé & al. (2017)) above the common tube; Anth narrowly triangular, 3.5 mm, recurved towards the apex; Ov slenderly ovoid; Sty  none, Sti 3-parted. — [F. Krapp] D. nana Leme & al. (Rodriguésia 61(1): 36–37, figs. 4G–H and 5H–N, 2010). Type: Brazil, Minas Gerais (Leme & al. 7484 [RB, HB]). — Distr: Brazil (Minas Gerais: Diamantina); on white quartzite soil in grassy Campo Rupestre vegetation. [3b] Ros dense, 8- to 10-leaved; L 6–8  1.2–1.4 cm, coriaceous, sheath 2.5–3  2.5–3.5 cm, castaneous abaxially, pale adaxially, glabrous towards the base, densely white-lepidote apically, L lamina narrowly triangular, distinctly canaliculate, suberect to  erect and slightly secund, acuminate with pungent tip, green, obscurely white-lepidote, distinctly veined, laxly spinulose, Sp subtriangular-acicular, spreading to slightly antrorse, 0.5–1 mm, 4–10 mm apart; Inf 19–29 cm, simple, erect; peduncle 15–20 cm, green, glabrous, smooth; peduncular Bra erect, subcarinate, long lanceolate-acuminate-caudate from a broadly subtriangular base, 5–11  3–3.5 mm, stramineous, sparsely white-lepidote, veined, microscopically denticulate to  entire; fertile Inf part 4- to 6-flowered, subdense to lax, 2.5–5 cm, rachis  straight, smooth, greenish to orange-yellow, glabrous; floral Bra broadly ovate-subtriangular, acuminate-caudate, subcarinate, subspreading to suberect, 4–4.5  3.5–4 mm, stramineous towards the apex, finely veined, fimbriate, remotely and irregularly denticulate to entire; Fl spreading and secund, 12–13 mm,

F. Krapp and U. Eggli

subtubular to slightly campanulate, unscented; Ped 3.5–5 mm, slender, terete, glabrous, yelloworange, curved; Sep broadly ovate, 4–5.5  3.5–4 mm, convex, apex obtuse and remotely and irregularly apiculate, ecarinate, orange, glabrous, retrorsely-curved fimbriate; Pet  broadly obcordate from a distinctly narrowed base, 7.5  6.5–7 mm, apex broadly emarginate, ecarinate, orange; St included; Fil 1.5 mm connate above the 1.5 mm high common tube, complanate, yellow; Anth oblong-ovoid, 2.5 mm, base sagittate, acute, yellow,  straight; Ov suboblong-ovate, 2.5  1.5 mm, yellow; Sty 1 mm, yellow, Sti shortly crenulate, lacerate, yellow. Similar to D. consimilis and D. macedoi. D. nervata Rauh (Trop. subtrop. Pfl.-welt 60: 21–24, figs. 13 and 14, 1987). Type: Brazil, Bahia (Rauh 56443 [HEID]). — Distr: Brazil (Bahia); quartzitic rocks, 1000 m. [3a] Ros 7-leaved, solitary or in groups, stem short, surrounded by old sheaths, slightly thickened at the base; L to 18 to 1 cm, sheath broadly ovate, 2  1.8 cm, white, densely lepidote abaxially, glabrous adaxially, strongly veined, L lamina erect (younger) to recurved (older), sturdy, strongly canaliculate, acute, with pungent tip, reddish-green, adaxially sparsely lepidote, abaxially densely white-lepidote, veined, Sp retrorse, 2 mm, brownish; Inf to 12 cm, simple; peduncle erect, 6 cm, olive-green, white-lanatelepidote; peduncular Bra amplexicaul at the base, shorter than the internodes, appressed to the peduncle, soon drying; fertile Inf part lax, 5- to 7-flowered, erect, 5 cm, white-lanate; floral Bra with a broad sheath, lamina narrowly lanceolate, recurved, shorter than the sepals, olive-green, lanate, entire; Fl suberect; Ped short, stout; Sep free, broadly ovate, 5  5 mm, obtuse, ecarinate, brilliant orange, abaxially sparsely lanate; Pet erect, spatulate, 10  5 mm, obtuse, ecarinate, brilliant orange; St included; Fil free above the 5 mm high common tube, yellowish; Ov 5 mm, white; Sty 2 mm, orange; Sti erect, shorter than the anthers. Compared with D. oligantha (here treated as synonym of D. saxatilis) in the protologue. — [F. Krapp]

Dyckia BROMELIACEAE

D. niederleinii Mez (in Martius, Fl. Bras. 3(3): 474, 1894). Type: Argentina, Misiones (Niederlein 229 p.p. [B, F [photo]]). — Distr: N Argentina (Misiones); open fields, terrestrial or on rocks. I: Smith & Downs (1974: 555). Incl. Dyckia missionum Mez (1894); incl. Dyckia missionum var. breviflora Hassler (1919). [1] Ros not described; L 25–50  1–3.5 cm, sheath not described, L lamina narrowly triangular, sparsely pale-appressed-lepidote abaxially, laxly serrate, Sp slender, curved, 4 mm; Inf 100–120 cm, few-branched, glabrous; peduncle slender; peduncular Bra broadly ovate with long-attenuate entire tips, shorter than the internodes; fertile Inf part with erect branches, elongate, very laxly flowered; primary Bra only a little larger than the floral bracts; floral Bra broadly ovate or suborbicular, apiculate, 5–8 mm, those of the branches much shorter than the sepals, those of the main inflorescence axis longer than the sepals, lustrous; Fl suberect, 13 mm; Ped 2–4 mm, very stout; Sep suborbicular, 5–7 mm, obtuse to shortly mucronate, carinate; Pet lamina spreading, elliptic, 13–15 mm, obtuse or emarginate, undulate, colour not described; St exserted; Fil highly connate for 6 mm above the common tube; Anth sublinear, 2 mm, acute, recurved; Sty  equalling the ovary. D. nigrospinulata Strehl (Bromeliaceae 42(5): 9–13, figs. 5–9, 2009). Type: Brazil, Rio Grande do Sul (Strehl 1602 [HAS]). — Distr: Brazil (Rio Grande do Sul); on rocks in a river. I: Klein V. & Klein (2014: 80). [1] Ros open, 60- to 200-leaved, 1.5 m ∅; L 70–80  5–6 cm, sheath suborbicular, dark castaneous, glabrous, L lamina triangular, attenuate, stiff, with a strong pungent black tip, dark green, centre lighter, adaxially glabrous, abaxially lepidote, striate, Sp predominantly antrorsely curved, 5–7 mm, dark, 30–70 mm apart; Inf 2–2.5 m, compound, 3–4 branched; peduncle strong, green, glabrous; peduncular Bra broadly ovate with long-attenuate tip, 80–100  12–20 mm (lower) to 10–40  7–10 mm (upper), entire or lower ones with 3 mm long teeth towards the apex; fertile Inf part with easily removable white-woolly tomentum, 25–30 cm;

937

Br laxly many-spiked, at acute angles, somewhat pendent with ascending tip, 50–60 cm, 2–5 cm apart; floral Bra triangular, acute, 1–2 mm, becoming shorter upwards, with brown tips, later stramineous,  entire; Fl pointing upwards in bud but spreading at anthesis, later sometimes turned backwards, sessile, narrowly tubular; Sep free, rounded, 5–6  1–2 mm, green-yellow with a small spot at the apex; Pet free, erect, rounded, 7–10  1–2 mm, yellow; St exserted; Fil free, linear; Ov incl. style as long as the stamens; Fr 6–8  3–4 mm, black, glossy; Se elongate, 3  1 mm, light castaneous. Part of the “Prionophyllum complex”, and similar to D. maritima. The large size is exceptional for the genus Dyckia. D. nobilis Büneker & al. (Phytotaxa 244(1): 60–63, ills., 2016). Type: Brazil, Minas Gerais (Büneker & al. 296 [HDCF, RB]). — Distr: Brazil (Minas Gerais: Serro); on sedimentary rocks; only known form the type locality. [2/3b] Ros solitary, with short stem, 28–55 cm ∅; L 10–30, inner straight, outer suberect to recurved, sometimes secund, sheath 5  4.5 cm, suborbicular, white with brown-greenish base, L lamina 25–80  2.1–6.2 cm, narrowly triangular, stiff and succulent, upper face flat or canaliculate, reddish-green, densely whitecinereously lepidote on both faces, tip extended into a 6 mm long spine, margins unarmed or serrate with laxly arranged Sp to 4 mm; Inf 60–170 cm tall, simple or paniculate, erect; peduncle 30–90 cm, reddish, basally whitetomentose, above brownish-ferruginous-tomentose; peduncular Bra 50 mm, shorter than the internodes, basal ones leaf-like, upper ones erect, triangular, 10–25  13–26 mm; fertile Inf part 25–95 cm, with up to 8 (sub-) erect to upcurved branches at the base, laxly 27- to 110-flowered; primary Bra ovate-triangular; floral Bra ovatetriangular, 8–15  10–21 mm, brownishferruginous tomentose-lepidote, entire, basal ones sometimes longer than the flowers, upper ones shorter; Fl 15 mm, porrect or slightly reflexed, tubular; Ped 2–3 mm, tomentose; Sep ovate-elliptic, convex, 5–7  7–9.5 mm, orange with reddish base, thick and succulent, basally

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connate for 3 mm, sparsely ferruginoustomentose; Pet suborbicular-rhomboidal, 10–13  18–20 (?) mm, orange, sparsely lepidote near the base; St included; Fil narrowly triangular, yellow-orange, to 9 mm, connate for 1 mm above the 2 mm long common tube; Ov 5  2.5 mm, subcylindrical, yellowish; Sty 2 mm, orange; Sti conduplicate-spiral; Fr globoseovoid, glossy brown or black; Se discoid, with hyaline wing. Close to D. sordida and D. inflexifolia. The basally connate sepals are unusual in the genus. The measurements given in the protologue for the floral bracts and the petals, both described as potentially broader than long, are not supported by the drawings in fig. 2 of the protologue. — [U. Eggli] D. odorata L. B. Smith (Phytologia 10: 485, t. 2, figs. 8 and 9, 1964). Type: Brazil, Goiás (Dawson 14578 [US, UC]). — Distr: Brazil (Goiás: Chapada de Veadeiros); ecology not recorded; known only from the type collection. I: Smith & Downs (1974: 515). [3b] Ros not described; L 15  0.5 cm, sheath triangular-ovate, stramineous, smooth, sublustrous, sparsely lepidote towards the apex, L lamina narrowly triangular, appressedly white-lepidote, soon glabrous adaxially, laxly serrate, Sp slender, spreading, pale, 2 mm; Inf 10 cm, sessile without peduncle, simple, axis subdensely whiteflocculose, with some flowers at the base, then with some aborted flowers in the middle, and well-developed flowers distally; floral Bra broadly ovate with long narrowly triangular serrate lamina (lower) to merely apiculate (upper), lower ones exceeding the flowers; Fl described as dimerous, spreading, strongly sweet-fragrant; Ped 3 mm, slender; Sep suborbicular, 5–6 mm, very broadly acute and minutely apiculate, palemargined when dry; Pet lamina spreading, broadly rounded, 10 mm, orange; St much shorter than the petals; Fil almost completely connate; Anth sagittate, curved, 1.5 mm; Ov slenderly conical; Sty short but obvious and separate. The sessile inflorescence without distinct peduncle and the dimerous flowers are notable and induce speculations that the taxon might be

F. Krapp and U. Eggli

based on an aberrant and/or depauperate specimen. D. orobanchoides Mez (in Martius, Fl. Bras. 3 (3): 475, 1894). Type: Brazil, Minas Gerais (Tamberlik s.n. [W †, B [lecto]]). — Distr: Brazil (Minas Gerais: without exact locality); ecology not recorded; only known from the type collection. [1?] Only known from the fragmentary lectotype; L unknown; peduncle glabrous; peduncular Bra ovate, acuminate, much shorter than the internodes, entire; Inf few-branched, sparsely shorttomentulose, Br ascending, subdensely flowered; floral Bra broadly ovate, acute, shorter than the pedicels, fimbriate at the apex; Fl spreading; Ped 3 mm; Sep broadly ovate-elliptic, 7 mm, apiculate, fimbriate, appressed-tomentulose; Pet suborbicular, 11 mm, obtuse, carinate, erect, colour not described; St equalling or slightly exceeding the petals; Fil free above the common tube; Anth sublinear, acuminate, strongly recurved; Sty  none. Very insufficiently known and apparently never recollected. D. paraensis L. B. Smith (Phytologia 13: 150, t. 7, figs. 10–12, 1966). Type: Brazil, Pará (Fróes 30030 [IAN, US [photo]]). — Distr: Brazil (Pará: Araguáia region); sandstone ledges, 600 m; only known from the type collection. I: Smith & Downs (1974: 576). [3b] Ros subbulbous, many-leaved; L 50  2.5–3 cm, sheath broad, 4 cm, L lamina recurved-spreading, very narrowly triangular, rigid, soon glabrous adaxially, closely appressedly cinereous-lepidote abaxially, very laxly serrate, Sp subspreading, 1.5 mm, black; Inf 70 cm, simple; peduncle erect, very slender, soon glabrous; peduncular Bra remote, acuminate from a broadly ovate base, very small, entire; fertile Inf part very lax, >20 cm, glabrous; floral Bra like the upper peduncular bracts but apiculate, to 5 mm; Fl spreading; Ped 3 mm, rather stout; Sep elliptic-oblong, 6 mm, broadly rounded, ecarinate, rather thin; Pet very broadly rounded, 10 mm, carinate, colour not described but presumably orange; St  equalling the petals;

Dyckia BROMELIACEAE

Fil shortly (to completely?, Pinangé & al. (2017: fig. 3)) connate above the common tube; Sty  none. Compared with D. duckei in the protologue. D. pauciflora L. B. Smith & Read (Phytologia 38: 138, t. 6, 1977). Type: Brazil, Goiás (Hatschbach 39429 [US, MBM]). — Distr: Brazil (Goiás); peak of a rock dome, 800 m. [3b] Ros dense; L 8  1.7 cm, sheath broad, 1.5 cm, L lamina recurved-spreading, narrowly triangular, both faces appressedly cinereouslepidote, glabrescent adaxially, laxly serrate, Sp recurved, 3 mm, black; Inf 50 cm, stout, simple; peduncle green, glabrous; peduncular Bra acuminate or apiculate, very small, slightly longer (lower) to much shorter (upper) than the internodes, many, remote; fertile Inf part lax, 6-flowered, rachis geniculate, 5 cm, glabrous; floral Bra suborbicular, apiculate, 3 mm,  equalling the pedicels; Fl subspreading; Ped slender; Sep broadly elliptic, 6 mm, obtuse; Pet very broadly rounded, 9 mm, ecarinate, orange; St included; Fil connate above the common tube; Sty  none. Similar to D. paraensis. — [F. Krapp] D. paucispina Leme & Esteves (Vidalia 1(1): 28–30, ills., 2003). Type: Brazil, Mato Grosso do Sul (Esteves Pereira 378 [HB, UFG]). — Lit: Braun & Esteves Pereira (2006a). Distr: Brazil (C Mato Grosso do Sul); on rocks in Cerrado vegetation. [2/3a] Ros 10- to 16-leaved, forming small and dense clumps, stem subglobose, 4  3 cm; L 10–18  1.8–2 cm, very rigid, succulent, sheath wider than the lamina, broadly subreniform, greenish, inconspicuously lepidote at the apex, L lamina narrowly triangular, strongly recurved,  slightly canaliculate, subulate towards the very pungent tip, long-acuminate, dark green to reddish, adaxially subdensely white-lepidote, abaxially subobscurely whitelepidote abaxially, veined, margins glabrous to slightly white-floccose, entire or basally irregularly and very remotely spinose, Sp slightly uncinate, predominantly retrorse, 2–3 mm; Inf 70–105 cm, pseudosimple to laxly 2 (rarely

939

3) branched, erect; peduncle 45–58 cm, dark green, glabrescent towards the base, subdensely white-lanate towards the apex; peduncular Bra narrowly long-triangular to ovate-triangular from a suborbicular base, acuminate to acuminate-caudate, erect, distinctly shorter than the internodes, stramineous, white-floccose to glabrescent, veined, entire; fertile Inf part very slightly angled, 15 cm (simple) and 23-flowered or to 30 cm (compound), straight, reddish, densely white-lanate at anthesis; primary Br 2–7, laxly arranged and late-developing, 7–10 cm, 11to 15-flowered, subdense to dense mainly towards the apex, suberect, rachis flexuose to geniculate, pale orange, secondary Br incompletely known; primary Bra like the upper peduncular bracts; floral Bra suberect, ecarinate, lowest ovateacuminate and 6–7  4–5 mm, upper suborbicular-apiculate, to 3–5  4 mm, stramineous towards the apex, white-lanate to glabrescent, veined, entire; Fl suberect and somewhat upwardly secund, 16–18 mm, tubular with slightly narrowed throat, unscented; Ped 2–3 mm, stout; Sep broadly ovate to ovate-elliptic, 6–9  5–6 mm, apex obtuse and inconspicuously apiculate, ecarinate, pale orange, densely whitelepidote, fimbriate, entire or minutely crenulate; Pet broadly obovate-spatulate, 11–13  9–10 mm, obtuse-emarginate, base very narrow, ecarinate, erect at anthesis, bright orange, abaxially white-floccose towards the base to glabrous; St slightly exceeding the petals; Fil free above the 2–3 mm high common tube, 10–12 mm, pale orange; Anth sublinear to subtriangular, 4–4.5 mm, strongly recurved, distinctly visible at anthesis, base sagittate, apiculate; Ov suboblongovoid to narrowly pyramidal, 5.5–6 mm, yellowish; Sty 1 mm, orange, Sti 0.5 mm, orange, irregularly recurved. Similar to D. burchellii and D. pulquinensis. — [F. Krapp] D. pectinata L. B. Smith & Reitz (Phytologia 14: 486, t. 1, figs. 29–32, 1967). Type: Brazil, Minas Gerais (Pabst 4129 [HB, US [photo]]). — Distr: Brazil (Minas Gerais: Conceição do Rio Verde to Cambuquira); dry sandstone outcrops. I: Smith & Downs (1974: 570).

940

[3b] Ros not described; L 14  0.8 cm, sheath broadly ovate, stramineous basally, L lamina very narrowly triangular, subulate-attenuate, appressedly cinereous-lepidote on both faces,  glabrescent adaxially, subdensely pectinate-serrate, Sp flat, spreading, 6 mm; Inf 75 cm, simple; peduncle glabrous; peduncular Bra subfoliaceous (lower) to narrowly triangular (upper), thin, irregularly shorter than the internodes, serrulate; fertile Inf part laxly many-flowered, 21 cm, glabrous; floral Bra like the upper peduncular bracts, lowest ones exceeding the sepals, serrulate; Fl spreading to reflexed; Ped 3 mm, broadly obconical, constricted at the base; Sep broadly ovate, 7 mm, rounded, entire or apiculate, rugose when dry as if formerly fleshy, little if at all carinate; Pet obovate, 9 mm, carinate, red-orange; St included; Fil shortly (to completely?, Pinangé & al. (2017: fig. 3)) connate above the short common tube; Sty none. Similar to D. dawsonii. — [F. Krapp] D. pernambucana L. B. Smith (Phytologia 20: 179, t. 2, figs. 12–14, 1970). Type: Brazil, Pernambuco (Lima 65-4275 [IPA, US [photo]]). — Lit: Siqueiro Filho & Leme (2007: 316–318, with ill.). Distr: Brazil (Pernambuco); gneiss outcrops. Incl. Dyckia rupestris W. Till & Morawetz (1990). [2/3b] Ros 30-leaved, dense, with a subglobose base to 8  8 cm; L to 33  2.1 cm, very rigid and succulent, sheath broadly subreniform, 2  4 cm, dark brown towards the base, subentire, L lamina narrowly triangular, nearly flat, arcuate, canaliculate towards the subulate pungent tip, dark greenish wine-red, distinctly veined, adaxially inconspicuously white-lepidote to glabrescent, abaxially densely cinereouslepidote, margins irregularly densely to laxly serrate, Sp antrorse, uncinate, 0.5–1.5 mm, 2–25 mm apart; Inf to 80 cm, compound or simple; peduncle  straight, 39–60 cm, sparsely to subdensely pale lanate; peduncular Bra erect, subfoliaceous (lower) or narrowly triangular (upper), shorter than the internodes, (sub-) densely pale-lepidote, serrulate; fertile Inf part lax, 25–45 cm, main axis 18- to 37-flowered, suberect, appressedly whitelepidote; primary Bra like the upper peduncular

F. Krapp and U. Eggli

bracts, much shorter than the long sterile branch bases; primary Br suberect, 14–21 cm, laxly 8- to 14-flowered, with a 2.5–7 cm long sterile base; floral Bra triangular-ovate, acuminate, (sub-) densely lanate, basal ones narrowly triangular, 10–12  5–6 mm, minutely denticulate, upper  triangular, 5–8  4–6 mm, entire or microscopically denticulate; Fl 16–20 mm, suberect to slightly secund, campanulate, weakly fragrant; Ped 3–5 mm, stout; Sep broadly ovate, 6.5–9  6–7 mm, acute to narrowly obtuse and apiculate, ecarinate, orange, subdensely lanate; Pet broadly obovate-spatulate, 10–12  8–10 mm, obtuse, ecarinate, reddish-orange; St included; Fil 2–2.5 mm (to completely?, Pinangé & al. (2017: fig. 3)) connate above the common tube; Anth triangular, 2 mm, straight to slightly recurved; Ov suboblong, 5 mm, yellow; Sty 2 mm; Sti conduplicate-spiral, 1 mm, orange; Fr subglobose, 12–14  12 mm, dark brown, glossy, shortly rostrate; Se flat, asymmetrically subcuneate, 4  2.5 mm. Compared with D. encholirioides in the protologue, but the similarity is superficial (Siqueiro Filho & Leme 2007: 318). Rather, it is related to D. limae and similar to D. secunda. D. rupestris is synonymized on the base of Siqueiro Filho & Leme (2007: 316–318), and the description has been amended according to this source. D. piauiensis Esteves & Gouda (Phytotaxa 164(4): 296–300, figs. 1 and 2A–F, 2014). Type: Brazil, Piauí (Esteves Pereira 375 [UFG]). — Distr: Brazil (Piauí: Canto do Burití); dry Cerrado vegetation, rocky soil or on rocks, in full sun or partial shade, 335 m; only known from the type locality. [3b] Ros acaulescent, densely 19- to 25-leaved, 45–54 cm ∅, with bulbous base, with few basal offsets; L coriaceous, succulent, sheath broadly ovate, 2.5  3.8–4.6 cm, fairly succulent, both faces white and brilliant, distally somewhat brownish, margins minutely dark-spined, L lamina first erect, later curved and spreading, triangular-lanceolate, 21–26  2.8–3.4 cm, green, both faces finely veined, basal part densely appressedly furfuraceous-lepidote between the veins, adaxially glabrous distally, margins laxly

Dyckia BROMELIACEAE

serrate, Sp mostly retrorsely uncinate, 0.5–4.3 mm, pale yellow and brown-tipped, 3–12 mm apart (distally to 27 mm); Inf 35–120 cm, simple, erect to  flexuous; peduncle 30–100 cm, sparsely lepidote; lower peduncular Bra leaf-like and to 7 cm, upper 0.8–4.2 cm, narrowly triangular to sublinear, with pungent tip, carinate, (pale) green, soon stramineous, sparsely serrulate; fertile Inf part 5.5–14 cm, rose-red, 7- to 16-flowered, sparsely white-lepidote; floral Bra broadly ovate, 8  2 mm, acuminate, rose-red, carinate; Fl 15  8 mm, spreading and slightly secund, broadly tubular; Ped stout, 3 mm (to 8 mm in fruit); Sep broadly ovate, 7–9  7 mm, rounded or emarginate, strongly convex, incurved, ecarinate, red, smooth, sparsely white-lepidote to glabrous, margins minutely spinose; Pet shortly spatulate, 10–13  9 mm, rounded or emarginate, slightly cucullate, orange; St included; Fil highly connate for 7–9 mm above the common tube; Anth 3 mm, yellow; Ov narrowly ovoid, 6–7  3 mm, pale yellow; Sty 1 mm; Sti 2 mm, crisped, orange with yellow margins; Fr ovoid, 14–17  11–13 mm, dark brown, glossy. Compared with the similar D. pernambucana, differing by larger retrorse leaf marginal spines, shorter inflorescences and characters of the flowers. — [F. Krapp] D. piracanjubensis Esteves & Gouda (J. Bromeliad Soc. 66(3): 172–179, ills., 2017). Type: Brazil, Goiás (Esteves Pereira 355 [UFG 50537]). — Distr: Brazil (S Goiás: along the Rio Piracanjuba); isolated flat limestone rock outcrops, in full sun, 650 m. [3b] Ros 67 cm ∅, solitary or forming dense clumps; stem to 17  6.5 cm, hidden by the leaf sheaths; L 40, spreading, the inner suberect to upward-secund, sheath 2.5–3.2  3.7 cm, succulent, both faces glossy, base white, dark honeycoloured above, margins spinose, L lamina sublinear-triangular, 35–53  3 cm, strongly canaliculate, pale green to brownish-green, finely veined, densely canescent-lepidote esp. abaxially, becoming glabrescent towards the tip, tip flat, dark brown, glossy, margins spinose, Sp uncinate, antrorse except for the distal ones, to 3.5 mm, becoming smaller upwards, brown from green

941

base, 3–14 mm apart; Inf 1.8–2 m tall, simple; peduncle to 100 cm, straight to slightly sinuous, sparsely cinereous-tomentose; lower peduncular Bra leaf-like, to 9.5 cm, upwards becoming smaller, linear-lanceolate, carinate, pungent, serrate; flowering part 72 cm, cinereously whitishlepidote; lower floral Bra triangular-lanceolate, 15  7 mm, upper only 3 mm, all carinate, veined, serrulate, sparsely cinereous-tomentose, pungent; Fl 16  7 mm ∅, subsessile to shortly pedicellate,  horizontally porrect, cup-shaped with slightly flaring limb; Sep triangularlanceolate, 7  6 mm, orange-yellowish, free, fleshy, densely white-tomentose, margins ciliate; Pet 13  9 mm, orange-red, basally connate for 1–3 mm, entire, rounded or slightly emarginate, glabrous; St included; Fil whitish to pale yellow, to 7.6 mm, 2–3.5 mm connate above the common tube; Anth 3  1 mm, dorsifixed near the base, strongly recurved at anthesis; Ov 4=5 superior, narrowly subprismatic, 6  2.5 mm, gradully merging into the style; Sty yellow basally, orange distally, including stigma 3 mm; Sti lobes 1.7 mm, orange; Fr broadly ovoid, 13–16  9–13 mm ∅, glossy dark brown, somewhat beaked; Se not described. Compared with D. goiana, but also similar to D. formosensis (with fewer leaves, longer fertile inflorescence part, obtuse sepals and longer filaments) and D. cangaphila (smaller rosettes and shorter leaves, shorter and occasionally compound inflorescences, and pedicellate smaller flowers). — [U. Eggli] D. platyphylla L. B. Smith (Phytologia 19: 283, t. 1, figs. 9–11, 1970). Type: Brazil, Bahia? (Foster 2489 [US]). — Distr: Brazil (Bahia?); not known from the wild with certainty. I: Smith & Downs (1974: 575). [3b] Ros not described; L to 23  5 cm, sheath suborbicular, 5 cm, yellowish, glabrous, L lamina narrowly triangular, thick, succulent, glabrous adaxially, appressedly whitish-lepidote abaxially, margins serrate, Sp slender, antroretrorsely curved, 3 mm, brown; Inf 80 cm, simple; peduncle slender, much compressed at the base, somewhat flexuous, glabrous; peduncular Bra broadly ovate and acuminate, small, much shorter than the

942

internodes, sparsely pale-lepidote, entire or subentire; fertile Inf part lax, many-flowered, 28 cm, glabrous; floral Bra broadly ovate, acuminate,  equalling the sepals (lower); Fl mostly suberect; Ped 1–2 mm, stout; Sep broadly ovate, 8 mm, rounded and cucullate, ecarinate; Pet elliptic, 11 mm, yellow; St included; Fil connate above the common tube; Sty none. Similar to D. aurea. — [F. Krapp] D. polyclada L. B. Smith (J. Bromeliad Soc. 39(5): 206–207, figs. 7 and 8, 1989). Type: Brazil, Rio Grande do Sul? (Foster 3096 [US, B, GH, NY]). — Distr: Brazil (Rio Grande do Sul?); ecology not recorded; only known from the type. [1] Ros clustering, acaulescent; L >40  2 cm, erect to spreading, sheath pale, L lamina finely white-lepidote between the veins on both faces, margins laxly serrate, Sp curved, 5 mm, 20–25 mm apart; Inf to 2 m, seemingly terminal, laxly 2–3 branched; peduncle erect, stout; peduncular Bra subfoliaceous, exceeding the internodes but exposing most of the axis; fertile Inf part densely white-flocculose; Br usually with 1 or 2 branches near the base, spreading to recurved, 50 cm, densely many-flowered; primary Bra very narrowly triangular, much shorter than the sterile branch bases; floral Bra ovate, acute, 4 mm; Fl spreading-ascending, narrowly elongately barrel-shaped; Ped very short; Sep suborbicular, 4 mm; Pet spatulate, not further described, yellow; St and Sti slightly exserted. Described on the base of cultivated material without clear provenance. Apparently related to D. selloa of the “Prionophyllum complex”. D. pontesii Büneker & al. (Rodriguésia 66(2): 500–503, ills., 2015). Type: Brazil, Rio Grande do Sul (Büneker & Witeck 212 [HDCF, RB, SMDB]). — Distr: S Brazil (S Rio Grande do Sul: Pinheiro Machado); conglomerate rock outcrops. [1/3a?] Ros 18–60 cm ∅, offsetting and rhizomatous; L 50, suberect to spreading, sheath suborbicular, 3  4.5 cm, white, glossy, L lamina narrowly triangular, 15–35  1.2–3.5 cm, stiff and succulent, straight to curved, adaxially

F. Krapp and U. Eggli

slightly canaliculate, green, sparsely lepidote only near the base, otherwise (sub-) glabrous, abaxially veined-striate, white-lepidote between the veins, tip a pungent spine to 6 mm, margins serrate-spinose, Sp slender and often inconspicuous, curved, 1–4 mm, laxly arranged, brown or yellowish; Inf 0.6–1.7 m tall, simple or branched; peduncle robust, 35–75 cm, green, subglabrous to densely white-flocculose-tomentose; basal peduncular Bra leaf-like, to 12 cm, as long as or longer than the internodes, erect, serrate, upper ones ovate to broadly elliptic, 2–6.5  1.3 – 2.6 cm, entire or inconspicuously serrate; fertile Inf part 25–75 cm, laxly to subdensely 25- to 150-flowered, axis green or brownish-green, subglabrous to densely white-flocculose-tomentose; Br when present 1–7, erect or suberect-arcuate, near the base of the main axis; primary Bra like the upper peduncular bracts; floral Bra ovatetriangular, 10–21  11–35 mm, the lower as long or longer than the flowers, the upper  as long, bulging, basally green, above castaneous, subglabrous to white-tomentose; Fl sessile, 20 mm, suberect; Ped absent; Sep ovatetriangular, 6–9  11–15 mm, brownish-yellow with greenish base, glabrous or sparsely tomentose-lepidote towards the tip, the adaxial ones carinate, the abaxial one ecarinate or slightly carinate; Pet obtrullate, 18–20  10–13 mm, erect with slightly flaring tips, yellow, glabrous, tip rounded-cucullate; St included; Fil free above the common tube; Anth not described; Ov 7  4 mm, whitish-yellow; Sty 11 mm; Sti not described; Fr ovoid, size not described, castaneous to black, very glossy. The flowers are slightly asymmetrical (termed “zygmomorphic” in the protologue) when observed from the bottom because the adaxial sepals are carinate and closer together than the ecarinate or slightly carinate abaxial sepal. The species is overall similar to D. dusenii and D. elisabethae. The only other species reported to have asymmetrical sepals (and thus zygomorphic flowers) is D. lunaris. — [U. Eggli] D. pottiorum Leme (Phytotaxa 67: 26–28, ills., 2012). Type: Brazil, Mato Grosso do Sul (Leme & al. 8579 [RB, HB]). — Distr: Brazil

Dyckia BROMELIACEAE

(Mato Grosso do Sul); shallow soil on flat or slightly inclined rock outcrops. I: Braun (2018: 71). [3a] Ros distichously 12- to 18-leaved; L nearly prostrate, 23–41  1.5–2.5 cm, coriaceous, sheath reniform, 2.5  4 cm, with castaneous base, greenish at the apex, base glabrous, towards the apex densely whitelepidote, L lamina sublinear-attenuate, strongly U-canaliculate, apical portion recurved, apex long acuminate-caudate, pungent-spinescent, green or bronze to reddish, green specimens densely white-lepidote with glabrous apex, bronze and reddish specimens densely and coarsely white-lepidote abaxially and less dense adaxially, veined, margins glabrous, sparsely (apically) to densely spinose, Sp narrowly triangular to acicular, straight to prevailingly retrorseuncinate, 1–3 mm, castaneous, 1–7 mm apart; Inf 23–52 cm, simple, erect; peduncle 10–34 cm, green to dark purplish wine-red, glabrous; peduncular Bra erect, subcarinate, long acuminate-caudate from a suborbicular base, 8–21  3–5 mm, densely white-lepidote to glabrous, veined, minutely spinulose; fertile Inf part densely 11- to 18-flowered, 6–11 cm, slightly flexuous, orange towards the apex, glabrous; floral Bra subtriangular-ovate, acuminate, ecarinate, suberect to subspreading, 4–8  3–4 mm, stramineous, thin-textured, glabrous, veined, inconspicuously denticulate to entire; Fl subspreading at anthesis and suberect-secund afterwards, 14–15 mm, subtubular, unscented; Ped 2–3 mm, stout; Sep ovate, 6  5 mm, rounded and inconspicuously apiculate, ecarinate, orange to yellowish-orange, glabrous, entire; Pet broadly spatulate from a narrower base, 11  9 mm, apex truncate and inconspicuously emarginate, ecarinate, orange to yellowish-orange; St  equalling the petals; Fil free above the 1.5 mm high common tube, yellowish; Anth sublinear, 4 mm, base distinctly bilobed, apiculate, slightly recurved near the apex; Ov ovoid, 4.5  2 mm, yellowish; Sty 1.5 mm, yellowish; Sti orange; Fr broadly fusiform, 12–15  8–10 mm, dark greenish-castaneous; Se suborbicular, flat, 3.5  3 mm, castaneous. Compared with D. burchellii and D. coximensis in the protologue. — [F. Krapp]

943

D. princeps Lemaire (Jard. Fleur. 3: tt. 224–225 + text, 1853). Type: Brazil, Minas Gerais (Anonymus s.n. [[icono] l.c. tt. 224–225]). — Distr: Brazil (Minas Gerais); ecology not recorded. I: Smith & Downs (1974: 515). Incl. Dyckia gigantea K. Koch (1874) (nom. illeg., Art. 52.1). [2?] Ros dense, many-leaved, with a very short and stout rhizome covered with old sheaths; L 35–60  3 cm, arching-recurved, sheath suborbicular, large, L lamina narrowly triangular, flat, with pungent tip, glabrous adaxially, densely appressedly pale-lepidote abaxially, margins laxly serrate, Sp 3 mm; Inf 1 m or taller, amply compound; peduncle stout, short, ferrugineoustomentulose; peduncular Bra subfoliaceous, much exceeding the internodes, serrulate; fertile Inf part ferrugineous-tomentulose; Br elongate, laxly flowered; primary Bra like the upper peduncular bracts, shorter than the sterile branch bases; floral Bra spreading, lanceolate-triangular, attenuate, exceeding the sepals; Fl spreading; Ped to 15 mm, stout; Sep broadly ovate, 9–10 mm, acute or apiculate, convex, ecarinate; Pet spreading, suborbicular, 25 mm, apiculate, red-orange; St included; Fil shortly connate above the common tube; Anth triangular, acuminate, strongly recurved; Sty very short. — [F. Krapp] D. pseudococcinea L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 108, t. 109, fig. 1, 1943). Type: Brazil, Rio de Janeiro (Foster 1144 [GH]). — Lit: Mendes & al. (2012: conservation). Distr: Brazil (Rio de Janeiro, São Paulo, Paraná); swampy meadows, on the ground or on rocks. I: Smith & Downs (1974, 542). [3b] Ros dense, many-leaved; L 30  1–1.2 cm, sheath broadly elliptic, 3 cm, pale, glabrous, L lamina linear, attenuate, adaxially glabrous, appressedly whitish-lepidote abaxially, margins laxly serrate, Sp slender, spreading or retrorsely curved, 2 mm; Inf 60–100 cm, simple; peduncle slender,  glabrous at anthesis; peduncular Bra narrowly triangular from an ovate base, the uppermost slightly shorter than the internodes, minutely serrulate; fertile Inf part lax, 17–25 cm,  glabrous; floral Bra ovate, acuminate,  equalling the sepals (lowest), minutely serrulate; Fl

944

spreading at anthesis, then suberect; Ped 3–4 mm; Sep ovate, 7–9 mm, acute, cucullate; Pet broadly elliptic, 12 mm, obtuse, ecarinate, suberect, orange; St much shorter than the petals; Fil 2 mm connate above the common tube; Sty very short. Available information on the geographical range of this species is conflicting. Material identified as this species has been extensively reported from the Brazilian states as detailed above, but according to Mendes & al. (2012), as well as the Brazilian Red List for plants, D. pseudococcinea is endemic to the restingas of Maricá, Rio de Janeiro, and is classified as critically endangered. D. pulquinensis Wittmack (Meded. RijksHerb. [Leiden] 29: 88, 1916). Type: Bolivia, Santa Cruz (Herzog 1849 [L, B, F [photo]]). — Distr: Bolivia (Santa Cruz); dry rocky slopes, 1900 m. I: Smith & Downs (1974: 552). [2] Ros not described; L 14–25  0.6–1 cm, sheath suborbicular, 3 cm, dark castaneous, L lamina linear-triangular, pungent, appressedly cinereous-lepidote, glabrescent adaxially, margins very laxly serrate, Sp stout, retrorsely curved, 5 mm; Inf 80 cm, few-branched; peduncle very slender, very sparsely white-lepidote; peduncular Bra triangular-ovate, acute, much shorter than the internodes, upper ones 8 mm; Inf branches flexuous, very slender, laxly and secundly flowered; primary Bra like the upper peduncular bracts, much shorter than the sterile bracteate branch bases; floral Bra broadly ovate, apiculate, 3 mm; Fl 15 mm, glabrous, subsessile; Sep broadly ovate-elliptic, 8 mm, apiculate, fuscous when dry; Pet broadly obovate, 14  6 mm, rounded, golden-yellow; St slightly exserted; Fil free above the 1.5 mm high common tube; Anth narrowly triangular, 4.5 mm, distinctly recurved towards the apex; Ov slenderly ellipsoid, 9 mm; Sty none. D. crassifolia is sometimes treated as synonym of this species. — [F. Krapp] D. pumila L. B. Smith (Phytologia 13: 151, t. 7, fig. 13, 1966). Type: Brazil, Goiás (Irwin & Soderstrom 7365 [US, NY]). — Distr: Brazil (Goiás); steep rocky scree. I: Smith & Downs (1974, 565).

F. Krapp and U. Eggli

[3a] Ros nearly acaulescent, densely manyleaved; L 13  0.7 cm, sheath suborbicular, 2.5 cm wide, brown, glabrous, laxly and minutely serrate apically, L lamina linear-triangular, pungent, thick, white-lepidote, glabrous adaxially with age, margins laxly serrate, Sp slender, spreading, 2 mm, brown; Inf 18 cm, simple; peduncle erect, slender, very finely whitelepidote, soon glabrous; peduncular Bra ovate, attenuate, small, much shorter than the internodes, obscurely serrulate; fertile Inf part lax, few-flowered, 2–5 cm, very sparsely white-stellate-lepidote; floral Bra like the peduncular bracts but entire, to 6 mm; Ped 2 mm, broadly obconical; Sep broadly elliptic, 6 mm, rounded and apiculate, the posterior ones carinate; Pet very broadly subacute, 10 mm, orange; St barely exserted; Fil free above the short common tube; Sty  none; Fr subglobose, apiculate, 13 mm, dark castaneous. Similar to D. duckei. — [F. Krapp] D. racemosa Baker (Handb. Bromel., 132, 1889). Type: Brazil, Goiás (Gardner 4015 [K, K [photo]]). — Distr: Brazil (Goiás); dry hills. [3b] Ros not described; L >40  >2 cm, sheath not described, L lamina narrowly triangular, covered with a membrane of fused scales abaxially, margins laxly serrate, Sp 1.5 mm; Inf >85 cm, simple; peduncle very slender, glabrous; peduncular Bra remote, broadly ovate, acuminate, entire; fertile Inf part lax, >15 cm, minutely furfuraceous; floral Bra broadly elliptic, obtuse or sometimes minutely mucronate, to 3 mm; Fl ascending, 10 mm, glabrous; Ped to 6 mm; Sep elliptic-ovate, 6 mm, obtuse; Pet erect or suberect, subrhombic, obtuse, slightly carinate, red; St included; Fil high-connate above the common tube; Anth narrowly triangular, acuminate, slightly recurved; Sty as long as the ovary. D. racinae L. B. Smith (J. Bromeliad Soc. 38 (6): 248–249, figs. 5 and 6, 1988). Type: Brazil, Rio Grande do Sul? (Foster 3095 [US, B, GH, NY]). — Lit: Dorneles & al. (2014: with ills.). Distr: Brazil (Rio Grande do Sul: near São Pedro do Sul); open Pampa vegetation on shallow stony soil.

Dyckia BROMELIACEAE

[1] Ros spreading, acaulescent; L >55  4 cm, sheath pale, L lamina obscurely lepidote abaxially between the veins, glabrous adaxially, margins laxly serrate, Sp curved, 5 mm; Inf to >2 m, seemingly terminal, laxly amply 2 branched; peduncle erect; peduncular Bra subfoliaceous, exceeding the internodes but exposing most of the peduncle; Inf branches glabrous, lax, somewhat secundly flowered, 50 cm; primary Bra very narrowly triangular, much shorter than the sterile branch bases; floral Bra attenuate from a broadly ovate base,  equalling the sepals, enlarging in fruit to 25 mm; Ped very short at anthesis, to 6 mm in fruit, stout; Sep broadly ovate-lanceolate, 6–7  3–4 mm, rounded and apiculate, glabrous to sparsely pilose; Pet obovate to spatulate, 12–13  5–6 mm, obtuse, glabrous, goldenyellow with paler margins; St just included; Fil free above the common tube; Anth triangular with curved apex and cordate base; Fr ovoid, beaked, black; Se with asymmetric lateral wings. Part of the “Prionophyllum complex”, and similar to D. selloa. Dorneles & al. (2014) provide an amplified description based on recently collected material. They argue that the species is close to D. cabrerae, which differs by larger flowers on longer pedicels. D. ragonesei A. Castellanos (Lilloa 10: 454, fig. 2, 1944). Type: Argentina, Santa Fé (Castellanos s.n. [BA 19465, US [photo]]). — Distr: N Argentina (Entre Ríos, Santa Fé, Santiago del Estero), Bolivia (Santa Cruz), Paraguay (Boquerón, Amambay, Canindeyú); rocky slopes. I: Smith & Downs (1974, 555). [2/3b?] Ros not described; L 40  1.2 cm, sheath broadly triangular, curved, semiamplexicaul, 3  6 cm, L lamina linear-triangular, serrate, Sp curved, 4 mm, brown; Inf 60–80 cm, paniculate or simple; peduncle elongate, furfuraceous; peduncular Bra deltoid, shorter than the internodes; fertile Inf part yellowfurfuraceous; Br subdensely flowered; floral Bra broadly triangular, to 7 mm, tomentose-lepidote; Fl 16 mm, sessile; Sep ovate, 8 mm, obtuse, margins hyaline; Pet 14–15 mm, broadly and irregularly rounded, undulate, golden-yellow; St distinctly exserted; Fil 3–5 mm connate above the

945

2 mm high common tube; Anth oblong, apiculate, straight, 5 mm; Sty 1 mm; Fr pyramidal, 17  12 mm; Se with a falcate wing, 4–5  5 mm. Usually cited as endemic for Argentina, but also reported from Bolivia and Paraguay as shown above (E. Gouda, pers. comm., Dec. 2016). D. rariflora Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): 1195, 1830). Type: Brazil, Minas Gerais (Martius s.n. [M, F [photo]]). — Distr: Brazil (Minas Gerais); rocky fields, savanna, terrestrial or on rocks; 1000 m. I: Smith & Downs (1974: 552). [3b] Ros not described; L to 14  1 cm, sheath broadly ovate, large, L lamina narrowly triangular, attenuate, with pungent tip, sparsely pale-appressed-lepidote abaxially, margins laxly serrate, Sp stout, curved, 3 mm; Inf to 50 cm, simple or branched; peduncle somewhat lepidote; peduncular Bra broadly ovate, acuminate, shorter than the internodes, entire or serrulate; fertile Inf part with a few short branches at the base, lax or subdense towards the apex, few-flowered, soon glabrous; floral Bra reflexed, broadly ovate, apiculate, 7–10 mm, equalling or shorter than the sepals; Fl spreading; Ped very short but distinct; Sep broadly ovate, 6 mm, broadly subacute; Pet suberect, broad, 9 mm, obtuse, carinate, orange; St included; Fil high-connate above the common tube; Anth elliptic, apiculate, straight or slightly recurved; Sty 1=5 as ong as the ovary. Some early illustrations purporting to represent this species in fact show D. remotiflora (Smith & Downs 1974: 526). D. reitzii L. B. Smith (Anais Bot. Herb. “Barbosa Rodrigues” 2: 14, tt. 1–3, 1950). Type: Brazil, Santa Catarina (Reitz 2690 [US, HBR]). — Distr: Brazil (Minas Gerais, Paraná, Santa Catarina, Rio Grande do Sul); sandstone ledges and rocky soil, 1500–2500 m. I: Reitz (1983: t. 50); Klein V. & Klein (2014: 77, 79). [3b] Ros dense, 100-leaved; L to 38  1–2 cm, sheath suborbicular, 3.5 cm wide, brown, soon glabrous, laxly serrulate, L lamina triangularlinear, attenuate, with pungent tip, smooth and glabrous adaxially, cinereous and strongly veined abaxially, margins laxly serrate, Sp curved,

946

1.5–3 mm, antrorse, 10 mm apart; Inf 50 cm, simple; peduncle slender, ferrugineous-flocculose; peduncular Bra densely imbricate, lanceolate, acuminate, cinereous-flocculose, serrulate; fertile Inf part densely 30-flowered, 9–12 cm, ferrugineous-flocculose to subglabrous; floral Bra like the peduncular bracts, exceeding the flowers; Fl subspreading, some with additional petals; Ped 5 mm, stout; Sep ovate, 8–9 mm, acute and cucullate, densely ferrugineous-tomentose; Pet rhomboid, 16 mm, somewhat spreading, yellow or reddish-yellow; St included; Fil highconnate above the common tube; Ov pyramidal; Sty short; Sti slightly contorted. D. remotiflora Otto & A. Dietrich (Allg. Gartenzeitung 1: 129, 1833). Type: Brazil (Sellow s.n. [B, K]). — Distr: Brazil (Minas Gerais, Paraná, São Paulo, Santa Catarina, Rio Grande do Sul), Uruguay, N Argentina (Entre Ríos).  Dyckia rariflora var. remotiflora (Otto & A. Dietrich) Baker (1889). [3a] Ros offsetting, rhizome short and stout; L 10–25  0.8–1.2 cm, arching, flat, sheath broadly ovate or suborbicular, large, L lamina narrowly triangular, with pungent tip, evenly dark green, appressedly pale-lepidote esp. abaxially, laxly serrate, Sp slender, curved, 1–3 mm; Inf to 1 m, simple; peduncle stout, sparsely tomentulose at the nodes; peduncular Bra suborbicular, the lowest laminate, all but the lowest much shorter than the internodes; fertile Inf part lax, 12–20 cm, sparsely tomentulose when young; floral Bra broadly ovate, shorter than the sepals; Fl divergent to spreading; Ped very short, stout; Sep ovate, apiculate,  carinate; Pet lamina spreading, trapeziform, obtuse, carinate, dark orange; St included; Fil free above the common tube; Anth narrowly triangular, recurved; Sty as long as or longer than the ovary. Some early illustrations labelled D. rariflora in fact show D. remotiflora (Smith & Downs 1974: 526). D. remotiflora var. angustior L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 108, 1943). Type: Brazil, Rio Grande do Sul (Bornmueller 351 [GH]). — Distr: Brazil (Rio Grande do Sul); ecology not recorded.

F. Krapp and U. Eggli

[3a] Upper peduncular Bra and floral Bra acuminate. — [F. Krapp] D. remotiflora var. montevidensis (K. Koch) L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 108, 1943). Type: Uruguay, Montevideo (Sellow s.n. [B]). — Distr: Brazil (Rio Grande do Sul), Uruguay, N Argentina (Entre Ríos); dry open rocky ground, to 50 m.  Dyckia montevidensis K. Koch (1874)  Dyckia rariflora var. montevidensis (K. Koch) Baker (1889). [3a] Upper peduncular Bra and floral Bra with broad apiculate tip; Sep nearly or quite straight, 6–8 mm; Pet 11–17 mm. Formerly also cited to occur in Santa Catarina, but the references relate to D. leptostachya (Reitz 1983: 516–517). — [F. Krapp] D. remotiflora var. remotiflora — Distr: Brazil (Minas Gerais, Paraná, São Paulo, Rio Grande do Sul), Uruguay; rocky fields, savanna, on rocks or terrestrial. – Fig. 9.

Fig. 9 Dyckia remotiflora var. remotiflora (Birolini s.n.: Uruguay; Maldonado, Punta Ballena). (Copyright: U. Eggli)

Dyckia BROMELIACEAE

Incl. Dyckia rariflora var. cunninghamii Baker (1889); incl. Dyckia vaginosa Mez (1894). [3a] Upper peduncular Bra and floral Bra with broad apiculate tip; Sep cucullate, 8–10 mm; Pet 17–23 mm. — [F. Krapp] D. remotiflora var. tandilensis (Spegazzini) Cabrera (Fl. Prov. Buenos Aires 4(1): 449, 1968). Type: Argentina, Buenos Aires (Spegazzini 219 [LP]). — Distr: Argentina (Buenos Aires); ecology not recorded; only known from the type collection and apparently extinct.  Dyckia montevidensis var. tandilensis Spegazzini (1901). [3a] Upper peduncular Bra and floral Bra with broad acuminate tip; Sep cucullate; Pet 25  to 2.4 cm, sheath not described, L lamina very narrowly triangular, appressedly cinereous-lepidote on both faces but becoming  glabrous adaxially, margins laxly serrate, Sp retrorsely curved, 2 mm, brown; Inf 90 cm, simple; peduncle very slender, glabrous; peduncular Bra remote, triangular, acuminate, lower ones subfoliaceous, upper ones small, subentire; fertile Inf part very laxly secund-flowered, to 23 cm, sparsely and fugaciously pale-lepidote; floral Bra acute, distinctly exceeding the pedicels (lowest), entire; Fl spreading; Ped 3 mm, cylindrical; Sep ovate, 7 mm, rounded, thin, brownish; Pet apparently rather fleshy, lamina broadly rhomboid, 12 mm, ecarinate, brown; St included; Fil short-connate above the common tube; Sty 1 mm. The laxly secund inflorescence and the brown flowers are notable. D. secundifolia Leme (Phytotaxa 67: 31–33, ills., 2012). Type: Brazil, Mato Grosso (Kranz 129 [RB, HB]). — Distr: Brazil (Mato Grosso); shallow soils in depressions and crevices of sandstone rock outcrops. [3b] Ros dense, 25- to 30-leaved; L 8–11  1–1.4 cm, coriaceous, sheath suborbicular, L lamina narrowly triangular,  flat, suberect to  erect and unilaterally curved, acuminate, with pungent tip, light green, adaxially

950

subdensely to laxly white-lepidote, abaxially subdensely white-lepidote, veined, margins glabrous, laxly to subdensely spinose, Sp narrowly triangular, straight or sometimes slightly antrorse, 0.5–1 mm, glabrous, pale castaneous, 4–10 mm apart; Inf 55–62 cm, simple, erect; peduncle 30 cm, reddish towards the apex, sparsely and inconspicuously white-lepidote to glabrous; peduncular Bra erect, obtusely if at all carinate, broadly triangular from a slightly gibbose base, long laminate to acuminate, 10–21  4–7 mm, reddish to stramineous, subdensely white-lepidote to glabrescent, veined, remotely denticulate to entire; fertile Inf part 14- to 22-flowered, lax, straight, 23–25 cm, rachis orange-red, subdensely but inconspicuously white-lepidote to glabrescent; floral Bra ovate, acuminate-caudate (lowest) to narrowly acute and apiculate (upper), suberect to  spreading at anthesis, ecarinate, 5–9  3.5–5 mm, orange-red, apex stramineous, subdensely and inconspicuously white-lepidote, finely veined, subentire to entire; Fl suberect to  spreading at anthesis, erect afterwards, 16–17 mm, tubular, unscented; Ped 2.5–3 mm, orange, sparsely white-lepidote to glabrous; Sep broadly ovate, 7–8  6–7 mm, emarginate, ecarinate, strongly convex, orange-red, subdensely white-lepidote to glabrous, entire, densely fimbriate; Pet subrhomboid-orbicular, 11–13  11 mm, obtuse-emarginate, ecarinate, orange to reddishorange, glabrous, inconspicuously crenulate; St equalling the petals or slightly shorter; Fil 8–10 mm, 1.5–2 mm connate above the 2 mm high common tube, pale orange towards the apex; Anth narrowly subtriangular, 3–3.5 mm, strongly recurved at anthesis, base bilobed, apiculate; Ov narrowly suboblong, 6 mm, pale yellow; Sty 1 mm; Sti 1.5 mm, orange, minutely crenulate-lacerate. Compared with D. coximensis in the protologue. D. selloa (K. Koch) Baker (Handb. Bromel., 136, 1889). Type: Uruguay? (Sellow 3339 [B, GH, NY, US]). — Distr: Brazil (Rio Grande do Sul), Uruguay?; sandstone cliffs. I: Smith & Downs (1974: 512).

F. Krapp and U. Eggli

 Prionophyllum selloum K. Koch (1874); incl. Dyckia grandifolia Baker (1889); incl. Dyckia macracantha Baker (1889); incl. Dyckia myriostachya Baker (1889). [1] Ros acaulescent; L rigid, arching, sheath conspicuous, reniform (outer) to ovate (inner), castaneous abaxially, glossy, glabrous except abaxially near the apex, margins minutely spinose, L lamina to 30  3.3 cm, thinly to densely cinereous-lepidote esp. abaxially, with pungent tip, margins laxly serrate, Sp stout, deltoidcuspidate, straight to retrorse, 6 mm; Inf >70 cm, compound and 2–3 branched; peduncle 45–60 cm, stout, glabrous; peduncular Bra leaflike, much exceeding the internodes, appressedly lepidote, margins strongly spinose; fertile Inf part 30 cm, glabrous, Br numerous, spicate, the lowest to 20 cm, those with perfect flowers lax or sometimes flowers subverticillate, those with pistillate flowers dense in the apical 1=2 ; primary Bra narrowly lanceolate, attenuate, much shorter than the branches, usually entire, quickly deciduous; floral Bra broadly ovate, apiculate, 2–3 mm; Fl divergent to spreading, sessile, perfect Fl to 13 mm, pistillate Fl 6 mm; Sep suborbicular or broadly ovate, 5 mm (perfect flowers) or 2 mm (pistillate flowers), apiculate, together forming an urceolate structure that embraces the petals; Pet obovate to lanceolate, somewhat spreading, 11–12 mm (perfect flowers) or 5–5.5 mm (pistillate flowers), acute or obtuse, yellow; St  equalling the petals or slightly shorter; Fil free above the short common tube; Anth 2.5 mm or 70 cm, simple; peduncle slender, glabrous; peduncular Bra remote, triangular, pungent, serrulate; fertile Inf part lax, 20 cm, sparsely furfuraceous or glabrous; floral Bra spreading to reflexed, ovate with long triangular lamina, 8–10 mm, exceeded by the sepals, minutely puberulent; Fl spreading or reflexed at anthesis, 12 mm; Ped short but distinct, stout; Sep elliptic, 6.5 mm, subobtuse, glabrous; Pet lamina erect, obovate, broadly rounded, scarcely if at all carinate, colour not described; St included; Fil high-connate above the common tube; Anth subtriangular, acute, strongly recurved. Insufficiently known, and described on the base of material without known provenance. D. sickii L. B. Smith (Arq. Jard. Bot. Rio de Janeiro 15: 330, fig. m–q, 1958). Type: Brazil, Pará (Sick B-613 [RB, US]). — Distr: Brazil (Pará); on rocks. I: Smith & Downs (1974: 565). [3a] Ros not described; L to 20  0.7 cm, sheath suborbicular, 2 cm wide, brown abaxially, soon glabrous, subdensely serrulate, L lamina very narrowly triangular, with pungent tip, glabrous adaxially, sulcate when dry, finely appressedly whitish-lepidote abaxially or the veins glabrous, margins laxly serrate, Sp slender, curved, 1.5 mm; Inf 45 cm, simple; peduncle erect, very slender, glabrous; peduncular Bra broadly ovate with a very narrow triangular lamina, very small, entire, remote; fertile Inf part very lax, slender, slightly flexuous, 20–22 cm, glabrous; floral Bra short-acuminate from a broadly ovate base, 7 mm, entire; Fl divergent or more often spreading; Ped 4 mm, cylindrical; Sep

951

broadly ovate, 5 mm, sometimes erose at the apex, ecarinate, very obscurely lepidote; Pet broadly obovate, 10 mm, obtusely carinate, orange-yellow; St very slightly exserted; Fil free above the common tube; Anth oblong, strongly curved; Ov slenderly pyramidal; Sty very short. — [F. Krapp] D. silvae L. B. Smith (Phytologia 13: 151, t. 7, figs. 14 and 15, 1966). Type: Brazil, Pará (Silva 784 [IAN]). — Distr: Brazil (Pará); on rocks in “Campo” vegetation. I: Smith & Downs (1974: 542). [3a] Ros spreading, many-leaved, acaulescent; L 25  2 cm, rigid, thick, sheath suborbicular, 3 cm wide, brown, nearly glabrous, L lamina linear-triangular, filiform-attenuate, appressedly white-lepidote, glabrous adaxially with age, margins laxly serrate, Sp slender, spreading, 3 mm, brown; Inf 66 cm, simple; peduncle erect, slender, very sparsely white-lepidote, soon glabrous; lower peduncular Bra leaf-like, upper broadly ovate-attenuate, small but nearly all exceeding the internodes except the uppermost; fertile Inf part subdensely many-flowered, 18 cm, very sparsely white-lepidote; floral Bra like the uppermost peduncular bracts, lowest  equalling the sepals, serrulate; Ped 3 mm, subcylindrical, thick, strongly constricted at the base and appearing articulate; Sep 6–7 mm, ovate, obtuse with a wide thin margin; Pet to 17  6 mm, broadly rounded, strongly carinate, curved-spreading, yellow; St completely included; Fil free above the short common tube; Sty none. — [F. Krapp] D. simulans L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 108, t. 110, 1943). Type: Brazil, Minas Gerais (Foster 570 [GH]). — Distr: Brazil (Minas Gerais); rocky fields, 1300 m. I: Smith & Downs (1974: 530). [3a] Ros not described; L 7–8  0.8–1 cm, rigid, sheath suborbicular, large, L lamina narrowly triangular, with pungent tip, appressedly cinereous-lepidote, margins laxly serrate, Sp spreading, 2 mm; Inf to 30 cm, simple; peduncle slender, furfuraceous, soon glabrous; peduncular Bra ovate, long-acuminate, exceeding the

952

internodes, serrulate; fertile Inf part insufficiently known, probably becoming lax, 4 cm when young, densely furfuraceous; floral Bra broadly ovate, acute,  equalling the flowers (lowest), minutely serrulate; Fl spreading; Ped 3 mm, stout; Sep broadly ovate, 9–10 mm, obtuse, cucullate; Pet suborbicular, slightly exceeding the sepals, reddish, glabrous; St included; Fil free above the 2 mm high common tube; Sty very short. — [F. Krapp] D. sordida Baker (Handb. Bromel., 132, 1889). Type: Brazil, Minas Gerais (Saint Hilaire 402 [P, GH [photo]]). — Lit: Guarçoni & al. (2012). Distr: Brazil (Minas Gerais); rocky slopes, 1300–1400 m. Incl. Dyckia duarteana L. B. Smith (1967). [3a] Ros usually caespitose, 32- to 50-leaved; L 50 cm or longer, sheaths suborbicular, 4 cm, dark castaneous, L lamina very narrowly triangular, green with reddish margins, adaxially sparsely lepidote, abaxially densely or sparsely and inconspicuously lepidote, margins almost entire, Sp only few, 1 mm, almost obsolete; Inf to 1 m, simple; peduncle stout, sparsely floccose; peduncular Bra subfoliaceous, shorter than the internodes (upper), entire; fertile Inf part manyflowered, very lax, 40 cm, densely ferrugineous-tomentose; floral Bra reflexed, broadly ovate, acute, slightly shorter than the sepals; Fl spreading or reflexed, to 18 mm; Ped 3–5 (–15) mm; Sep broadly elliptic, 8–11 mm, obtuse, free or inconspicuously connate at the base, densely brown-ferrugineously lepidote; Pet erect, conspicuous, limb suborbicular and slightly flaring, neither undulate nor carinate, colour not described but presumably orange; St included; Fil free above the common tube; Anth short, subtriangular, apiculate, scarcely recurved; Sty very short; ovules with narrow asymmetric wings. The description above was completed from data presented by Büneker & al. (2016), but some doubts persist as to the pedicel length, said to be 3–5 (–15) mm long in the key on p. 58, but the illustrations on p. 64 show almost sessile flowers. D. sordida forms a complex with D. inflexifolia and D. ursina (Guarçoni & al. 2012), as well as with D. nobilis (Büneker & al. 2016). These last

F. Krapp and U. Eggli

authors argue that other species such as D. mellobarretoi and D. elata are also similar. All species are endemic to the Serra do Espinhaço of Minas Gerais. The complex is characterized by peduncles, inflorescence axes and sepals covered with brown-ferrugineous trichomes, orange petals, (sub-)triangular antepetalous filaments, and filaments free above the common tube. D. spinulosa L. B. Smith & Reitz (Phytologia 14: 486–487, t. 1, figs. 33–36, 1967). Type: Brazil, Minas Gerais (Duarte 7409 [HB, US [photo]]). — Distr: Brazil (Minas Gerais); savanna, on rocks; only known from the type collection. I: Smith & Downs (1974: 570). [3b] Ros not described; L 35  1.6 cm, sheath suborbicular, 3 cm, pale brown, L lamina lineartriangular, appressedly cinereous-lepidote on both faces, becoming  glabrous adaxially, margins laxly serrate, Sp mostly retrorsely uncinate, 1 mm; Inf 85 cm, simple; peduncle sparsely and finely pale-lepidote; peduncular Bra ovate, the lower ones dense and with long linear lamina, the upper acuminate, shorter than the internodes, serrulate; fertile Inf part laxly many-flowered, to 27 cm, finely white-lepidote at first; floral Bra like the upper peduncular bracts, equalling the middle of the sepals (lowest), serrulate; Fl curvedascending; Ped 5 mm, slender, angled, not constricted; Sep ovate, 7 mm, rounded and apiculate, strongly convex; Pet rhomboid, 10 mm, colour not described but presumably orange; St included; Fil short-connate above the common tube; Sty  none. — [F. Krapp] D. stenophylla L. B. Smith (Phytologia 14: 487, t. 1, figs. 42–45, 1967). Type: Brazil, Goiás (Irwin & al. 9728 [US, NY]). — Distr: Brazil (Goiás); soil-filled rock crevices, 1175 m. I: Smith & Downs (1974: 575). [3b] Ros dense, globose, many-leaved; L to 16  0.5 cm, sheath suborbicular, 2 cm, brown, L lamina linear, finely subulate-attenuate, appressedly whitish-lepidote on both faces, soon glabrous, margins laxly serrate, Sp slender, recurved, 2 mm; Inf to 30 cm, simple; peduncle glabrous; peduncular Bra broadly ovate, linearlaminate (lowest) or acuminate, remote, small,

Dyckia BROMELIACEAE

microscopically serrulate; fertile Inf part laxly few-flowered, to 10 cm, fugaciously whitelepidote; floral Bra like the upper peduncular bracts, equalling the sepals (lowest) or shorter, microscopically serrulate; Ped 1 mm, stout but distinct; Sep broadly ovate, 6 mm, rounded; Pet lamina broadly elliptic, 8–10 mm, orange; St included; Fil connate above the common tube; Sty none. — [F. Krapp] D. stolonifera P. J. Braun & Esteves (Cact. Succ. J. (US) 81(6): 301, ills. (p. 303), 2009). Type: Brazil, Mato Grosso do Sul (Esteves Pereira 627 [UFG]). — Distr: Brazil (Mato Grosso); borders of rock outcrops, 150 m. [3a] Ros with 19 living leaves, stem 8  2.9 cm, semi-underground, with 1 - 6 stolons emerging above the ground and to 16 cm long, becoming submerged, forming new rosettes 5–16 cm apart from the mother plant; L 38  2.6 cm, rigid, coriaceous, first slightly erect, later recurved-spreading, sheath 1.8  4.7 cm, cream-coloured to brownish adaxially, brilliant dark brownish and apically lepidote abaxially, L lamina triangular-lanceolate, olivegreen to dark purple-greenish, concave, with blunt yellowish tip, densely veined, adaxially slightly glossy and sparsely lepidote, abaxially greyish-lepidote, Sp predominantly retrorse, 2.7 mm,  cream at the base, dark brown at the tip, 4.3–16 mm apart; Inf to 95 cm, simple, erect; peduncle 55 cm, pale greyish-green with a brownish hue; peduncular Bra hard, carinate, amplexicaul (upper ones semi-amplexicaul), lower ones to 21  8 mm, adaxially scatteredlepidote, veined, margins cream, membranous; fertile Inf part slightly contorted, 24 cm, rachis rose to reddish,  glabrous; floral Bra 4  2.3 mm, rose, brilliant; Fl spreading to erect, 20 mm, subsessile; Sep elongate-oval, 8–12  7.2 mm, succulent, carinate, reddish-orange, margins pale; Pet 17  11 mm, abaxially orange, basally more reddish, margins delicately yellow; St exserted; Fil free above the 5 mm high common tube, erect, to 15 mm, white-yellowish; Anth yellow, slightly triangular, 4–5  1.4 mm, acute, recurved; Ov slightly triangular-elongate, 7  2 mm, white with a very pale yellowish hue; Sty

953

1 mm, Sti complex, 2.9 mm, apex fringed, yellow; Fr glossy dark brownish. D. strehliana H. Büneker & R. Pontes (Revista Brasil. Bioci. 11(3): 284–289, figs. 1 – 2, 2013). Type: Brazil, Rio Grande do Sul (Büneker & al. 101 [HDCF, SMDB, SP]). — Distr: Brazil (Rio Grande do Sul); basaltic river banks, periodically flooded. [2/3a] Ros 16- to 54-leaved, 8–19 cm ∅, rhizomatous; L 6–15  1.1–2.6 cm, rigid, succulent, sheath elliptic to orbicular, 2–3.5  2.5–4.4 cm, white, both faces glossy, L lamina triangular, straight or inflexed, with reddish pungent tip, veined abaxially, usually entire, rarely spinose with 1–10 spines per margin; Inf 35–120 cm, simple or rarely with 1–4 suberect branches; peduncle erect, slender, 19–60 cm, tomentose; peduncular Bra leaf-like, ovate to elliptic, longer (lower) to shorter than the internodes (upper); fertile Inf part 13- to 34-flowered (40–60 when branched), 18–40 cm, rachis white-tomentose; floral Bra ovate-triangular, slightly carinate, apex attenuate, 11–16  6–11 mm, stramineous; Fl suberect at anthesis, 20 mm; Ped absent or short; Sep ovate to elliptic, 8–10  4–8 mm, ecarinate to slightly carinate, yellow or green, tomentose; Pet erect, obovate to obtrullate, 14–18  6–11 mm, yellow; St included or equalling the petals; Fil shortly connate above the common tube; Ov 5  3 mm, whitish-yellow; Fr 15–20 mm. Similar to D. brevifolia and D. distachya. Plants are periodically flooded in their native habitat. D. subinermis Mez (in Martius, Fl. Bras. 3(3): 487, 1894). Type: Argentina, Misiones (Niederlein 1205 [B, F [photo]]). — Distr: N Argentina (Misiones); ecology not described. [3a] Ros acaulescent, densely leaved; L to 20  7 cm (inner ones) or wider, sheath broad, dark brown, glabrous, L lamina linear, attenuate, subdensely closely appressedly pale-lepidote abaxially, entire or very laxly serrate, Sp few, 0.5 mm; Inf 50 cm, simple; peduncle rather stout, puberulous towards the apex; peduncular Bra long-acuminate from a broadly ovate base,

954

to 2 cm, shorter than the internodes, entire; fertile Inf part lax, puberulous-lepidote; floral Bra reflexed, ovate, acuminate, 7 mm, much shorter than the sepals, entire; Fl erect, 18 mm, glabrous; Ped 4 mm, stout, angled, articulate; Sep ovate, 9.5 mm, rounded and mucronate; Pet suberect, suborbicular, 17 mm, rounded at the apex, undulate, carinate, yellow when dry; St included; Fil free above the common tube; Anth narrowly triangular, strongly recurved; Sty none and Sti sessile, strongly spirally contorted; Fr subellipsoidtrigonous, 15  6 mm, glossy brown, acute; Se 2  3 mm, basally somewhat acute, apically broadly rounded, with hyaline wing. D. sulcata Guarçoni (Phytotaxa 188(3): 170–172, ills., 2014). Type: Brazil, Minas Gerais (Guarçoni & Paixão 1663 [VIC]). — Distr: Brazil (Minas Gerais: Serro); in sand on quartzitic rock outcrops, 1000 m; only known from the type locality. [3a] Ros 20–43 cm ∅; L 10–27, asymmetrically spreading-arcuate to erect, sheath suborbicular, 1.5–4.3  2.3–6.7 cm, white to cream, upper 1=3 of the abaxial face white-lepidote and inconspicuously aculeate, L lamina very narrowly triangular, 9.9–17.8  0.9–1.4 cm, flat to shallowly canaliculate, very succulent and rigid, green, or basally rarely wine-red on the upper face, upper face sparsely white-lepidote in the lower 1=2 , lower face strongly ribbed and densely white-lepidote at the base esp. between the ribs, apex acute and pungent, margins spinose except the apical 1=3 , Sp patent to slightly retrorse, dark brown, 0.7–1.4 mm, 10–20 (–26) mm apart; Inf (24–) 38–62 (–71) cm, simple, erect; peduncle 16–48 cm, green to brown, white-lepidote; peduncular Bra shorter than the internodes, lower 8–60  1–9 mm, upper 6–14  4–8.6 mm, ovate, long-acuminate, spine-tipped, carinate, abaxially white-tomentose; fertile Inf part laxly 5- to 18-flowered, rachis orange, white- to cinereously tomentose becoming glabrous; floral Bra reaching to the middle of the sepals, ovatetriangular and long-acuminate, spine-tipped, 3.8–7.8  2.8–5.9 mm, orange to stramineous with green base, white-tomentose, fimbriate; Fl 14–19 mm, slightly reflexed or rarely patent,  tubular with flaring mouth; Ped 2.4–6.4 mm,

F. Krapp and U. Eggli

stout, orange, white-tomentose; Sep triangular to ovate, 5.9–8.9  4.4–6.9 mm, orange, fleshy, obtuse to acute, white-lepidote, fimbriate; Pet obovate to somewhat obtrullate, 9.7–13.1  6.7–10.6 mm, orange, obtuse to slightly emarginate, crenulate; St included; Fil yellow, to 9.5 mm, free above the 1.6–3.3 mm long common tube; Anth elliptic; Ov oblong, orange, 5.3–10.3 mm; Sty orange, 0.9–2.1 mm; Sti conduplicate-spiral, 0.6–0.7 mm, yellow; Fr ovoid, dark brown, 11.4–13.7  9.2–10.3 mm; Se not described. Part of the D. saxatilis complex, and compared with D. saxatilis and D. brachyphylla (both with symmetrical rosettes and leaves without spines in the apical ). — [U. Eggli] D. tenebrosa Leme & H. Luther (Selbyana 19 (2): 183–184, fig. 1, 1998). Type: Brazil, Minas Gerais (Leme & al. 2895 [HB, SEL]). — Distr: Brazil (Minas Gerais); rocky fields, terrestrial, 1000–1300 m. [3a] Ros not described; L 14  1.5 cm, rigid, thick, sheath not described, L lamina spreading-recurved, very narrowly triangular, attenuate towards the involute-subulate apex, acuminate-caudate, green to reddish, adaxially glabrous, abaxially densely white-lepidote and veined, margins laxly spinose, Sp retrorse, uncinate, 1.5–2 mm; Inf 30–40 cm, simple, erect; peduncle 20–30 cm, yellowish-green, subdensely white-floccose; peduncular Bra linear-triangular from a broadly ovate base, long-acuminate, erect, distinctly carinate, 10–20  7 mm, green at the base, reddish at the apex, veined, white-floccose to glabrescent, remotely denticulate-crenulate, apical ones like the floral bracts; fertile Inf part 4- to 9-flowered, lax, 9 cm, slender, flexuous, completely white-sublanate at anthesis except the petals; floral Bra broadly ovate-triangular, subcarinate, acuminate-caudate (lowest) to shortly acuminate (upper), 6–10  5–8 mm, veined, entire but upper ones remotely denticulate near the apex; Fl suberect to spreading at anthesis, 16 mm; Ped 2–3 mm, stout; Sep ovate, 7–8  5 mm, apex narrowly obtuse, ecarinate, fimbriate, reddish-green, entire; Pet broadly spatulate, 12  9 mm, narrowly obtuse, base very narrow, ecarinate, subspreading at anthesis,

Dyckia BROMELIACEAE

reddish-brown to wine-red with orange base, glabrous; St included; Fil free above the 2 mm high common tube, 6 mm, pale orange; Anth very narrowly triangular, 2.5 mm, strongly recurved, base sagittate, acuminate; Ov narrowly subpyramidal, 5 mm, green; Sty 2–4 mm, yellow, Sti elliptic, 1 mm, wine-red; Fr and Se not described. Similar to D. machrisiana and compared with D. oligantha as well as D. hilaireana (here both treated as synonyms of D. saxatilis). D. tenuis Mez (in Martius, Fl. Bras. 3(3): 484, 1894). Type: Brazil, Goiás (Gardner 3479 [BM, K]). — Distr: Brazil (Goiás, Mato Grosso); ecology not recorded. Incl. Dyckia morreniana Mez (1894); incl. Dyckia kuntzeana Mez (1896). [2/3a] Ros not described; L 20–40  1.5–1.7 cm, sheath small, L lamina narrowly triangular, attenuate, with pungent tip, appressedly pale-lepidote abaxially, margins laxly serrate, Sp slender, spreading to spreading-reflexed, 1.5–2 mm; Inf 40–50 cm, simple or racemose; peduncle slender, minutely furfuraceous or soon glabrous; lower peduncular Bra subfoliaceous, upper ones  as long as the internodes and entire or sparsely serrulate; fertile Inf part few-flowered, lax, minutely furfuraceous; floral Bra divergent to spreading, (broadly) ovate with narrowly triangular tips, usually  equalling the flowers (lower), entire; Fl erect or suberect, 10–12 mm; Ped to 3 mm, stout; Sep ovate-elliptic, 6–7 mm, acute or obtuse, glabrous or subglabrous; Pet lamina suberect, broadly acute or obtuse, reddish; St exserted; Fil free above the common tube; Anth recurved; Sty very short or none. Described on the base of incomplete material, and insufficiently known. D. tobatiensis Hassler (Annuaire Conserv. Jard. Bot. Genève 20: 309, 1919). Type: Paraguay, Cordillera (Hassler 2099 [G, F [photo]]). — Distr: C Paraguay (Cordillera); denuded hill slopes. [1] Ros not described; L 40–50  3 cm, thickly fleshy-coriaceous, sheath not described, L lamina narrowly triangular, both faces shortly brownish-

955

furfuraceous-lepidote, margins strongly serrate, Sp antrorsely curved, strong, 6–8 mm, wholly lepidote; Inf to 95 cm, paniculately branched; peduncle 50–80 cm, glabrous; peduncular Bra much shorter than the internodes; fertile Inf part glabrous, Br 4–5, laxly flowered, divergent, 15 cm; primary Bra ovate, mucronulate, 10 mm; floral Bra subreflexed, triangular-ovate, apiculate, to 5 mm,  equalling or shorter than the pedicels; Fl 17 mm; Ped 4–5 mm; Sep ovate, 6.5 mm, evenly rounded at the apex; Pet subrhomboidovate, 17  8 mm, obscurely carinate, goldenyellow; St equalling or slightly exceeding the petals; Fil free above the common tube; Anth narrowly elliptic, 3.5  1.5 mm, acute, barely recurved; Ov pyramidal, 10 mm; Sty exserted; Sti globose, contorted. D. tomentella Mez (Repert. Spec. Nov. Regni Veg. 16: 69, 1919). Type: Paraguay (Fiebrig 4046 [B, GH]). — Distr: N Paraguay (Presidente Hayes?, Amambay?); ecology not recorded. [3a] Ros and L not described; Inf >80 cm, simple; peduncle erect, very slender, 60 cm; peduncular Bra acuminate from a very broadly ovate base, much shorter than the internodes, prominently striate, entire; fertile Inf part rather few-flowered, sublax, straight, 20 cm, appressedly cinereous-lepidote; floral Bra subspreading, very broadly ovate, 7 mm, apiculate, not much shorter than the sepals, entire; Fl suberect to erect, subsessile; Sep very broadly elliptic, 6 mm, rounded and emarginate at the apex; Pet erect, claws very short, lamina very broadly trapeziform-orbicular, to 10 mm, very broadly rounded and slightly emarginate, ecarinate, golden-yellow, with minutely undulate margins; St 2 mm longer than the petals; Fil broad, free above the common tube; Sty very short. The original description is based on very incomplete material. Tropicos (accessed Dec. 2016) lists two recent collections from Paraguay, whose collection notes report caespitose rosettes and yellow flowers. D. trichostachya Baker (Handb. Bromel., 133, 1889). Type: Brazil, Minas Gerais (Sellow Brom. Paris 59 [P, GH [photo]]). — Distr: Brazil

956

(Minas Gerais); rocky fields, terrestrial or on rocks. I: Smith & Downs (1974, 523). Incl. Dyckia micracantha Baker (1889). [2] Ros not described; L 50  >2 cm, sheath not described, L lamina narrowly triangular, densely lepidote esp. abaxially, margins laxly serrate, Sp 1.5 mm; Inf to 1.3 m, few-branched at the base or pseudo-simple with branch buds in the axils of the lower bracts; peduncle stout, 100 cm, soon glabrous; peduncular Bra broadly ovate with narrowly triangular acuminate lamina, exceeding the internodes, serrulate; fertile Inf part 30 cm, sparsely ferrugineous-tomentose; primary Bra much shorter than the branches; floral Bra broadly ovate, acute, lower ones equalling or exceeding the flowers and serrulate; Fl suberect to spreading, 12–13 mm; Ped short, stout; Sep broadly elliptic, 7–12 mm, obtuse,  glabrous; Pet erect, suborbicular, ecarinate, colour not described; St included; Fil free above the common tube; Anth linear, slightly recurved; Sty very short. — [F. Krapp] D. tuberosa (Vellozo) Beer (Fam. Bromel., 157, 1857). Type: Brazil (Vellozo s.n. [[icono]: Vellozo, Fl. Flumin. Icon. 3: t. 136, 1835]). — Lit: Vosgueritchian & Buzato (2006: ecology, with ills.). Distr: Brazil (Minas Gerais, Paraná, São Paulo, Santa Catarina); dry fields and open scrub, Cerrado vegetation, 800–1300 m. I: Smith & Downs (1974: 565); Reitz (1983: t. 52).  Tillandsia tuberosa Vellozo (1829); incl. Dyckia coccinea Mez (1894). [3a] Ros many-leaved, with bulbous base, acaulescent, slowly caespitose; L 15–50  1 cm, erect to spreading, sheath broadly ovate or suborbicular, large, persistent and forming a bulb, L lamina straight to  curved, narrowly triangular, abruptly acute and pungent, usually involute, glabrous adaxially, appressedly pale-lepidote abaxially, laxly and evenly serrate, Sp curved, dark; Inf 35–100 cm, simple; peduncle slender, lepidote or glabrescent; peduncular Bra erect, broadly ovate with narrowly triangular thick lamina, much shorter than the internodes (upper), serrulate; fertile Inf part lax, few- to manyflowered,  glabrous; floral Bra spreading or reflexed, lancolate-triangular, acuminate, usually

F. Krapp and U. Eggli

shorter than the sepals, serrulate; Fl spreading to slightly downwards-pointing at anthesis, broadly tubular, erect in fruit; Ped 2–3 mm; Sep ovate, 5–7 mm, acute or obtuse, strongly convex; Pet claw broad, lamina suberect, broadly obovate, 9–14 mm, carinate-complicate, reddish or orange; St included; Fil free above the common tube; Anth narrow, recurved; Ov 5 mm, narrowly pyramidal; Sty 1 mm; Sti contorted. Vosgueritchian & Buzato (2006) found 3 species of hummingbirds as pollinators of the flowers. The inflorescences are also visited by several species of ants. Only 55% of the plants of the study population in São Paulo State flowered, and asexual reproduction appears to be important. D. tubifilamentosa Wanderley & G. Sousa (Hoehnea 41(2): 315–319, figs. 1–2, 2014). Type: Brazil, Piauí (Wanderley & al. 2630 [SP, TEPB]). — Distr: Brazil (Piauí); rocky soil or rock outcrops in Caatinga and transitions to Cerrado vegetation. [3b] Ros in small clumps or solitary; L distichous or secund, 13–17 cm, sheath broad, oval, 2–2.5  3–4 cm, fleshy, whitish to greenish, entire, L lamina straight to curved, narrowly triangular with mucronate tip, green or brownish to vinaceous, cinereous-lepidote, strongly spinose, Sp 1–2 mm, brown and brittle, erect to retrorse; Inf 22–58 cm, simple; peduncle 10–50 cm, green to vinaceous, glabrous; lowest peduncular Bra subfoliaceous, creamy-green, entire, the remaining peduncular Bra 20  8–20 mm, shorter than the internodes, ovate to filiform, apex attenuate, reddish-brown, entire; fertile Inf part lax, 2to 10-flowered, secund or rarely distichous, slender, slightly geniculate; floral Bra ovate, apex caudate, 3–5 mm, green to reddish-brown, entire; Fl sessile to subsessile, 28–32 mm; Sep ovate, 8–11 mm, apiculate, green with translucent dots and dark reddish wrinkles at the base, entire; Pet oblong-lanceolate, 17–20 mm, acute, slightly cucullate, green or with dark reddish wrinkles at the apex, entire; St exserted, 20–28 mm; Fil entirely connate above the common tube, white becoming vinaceous to purplish; Anth sagittiform, connivent, divergent after anthesis, 5–6 mm,

Dyckia BROMELIACEAE

creamy-yellow; Ov 10 mm; Sty 11 mm; Sti spirally-conduplicate; Fr 14  9 mm, brown; Se (round-) ovoid to falciform, 4.2  2.5 mm, light to dark brown, with corky wings. Unique in the genus by having small green petals, and a long-exserted white staminal tube that becomes wine-red to purplish after anthesis. The spirally-conduplicate stigma lobes, as well as petal colour, are reminiscent of species of Encholirium. D. tweediei Mez (in Martius, Fl. Bras. 3(3): 485, 1894). Type: Argentina, Santiago del Estero (Tweedie s.n. [K]). — Distr: N Argentina (Santiago del Estero, Salta); terrestrial or on rocks. [3a] Ros acaulescent; L 50  1.5 cm or longer, sheath broad, thick, dark brown, L lamina narrowly triangular, rigid, canaliculate, densely pale-lepidote abaxially, margins subdensely serrate, Sp slender, 2 mm; Inf 1 m or more, simple; peduncle slender, subglabrous; peduncular Bra very broadly ovate, abruptly acute, much shorter than the internodes, entire; fertile Inf part rather lax, subsecund, tomentellouslepidote; floral Bra subspreading, broadly ovate, to 6 mm, abruptly acute or subrounded, shorter than the sepals; Fl ascending or the uppermost erect, 19 mm, subsessile; Sep elliptic-ovate, to 8 mm, incised-emarginate, lepidote-tomentellous; Pet erect, broadly obovate, 15–16 mm, rounded, ecarinate,  undulate, yellow; St exserted; Fil free above the common tube; Anth triangular, acute, slightly recurved; Ov narrowly bottle-shaped, with six longitudinal furrows; Sty thick, as long as the ovary; Sti with 3 free spreading lobes; Fr not described. D. uleana Mez (in A. & C. de Candolle, Monogr. Phan. 9: 517, 1896). Type: Brazil, Goiás (Ule 3134 [Herb. Taubert †?, B]). — Distr: Brazil (Goiás); ecology not recorded. [3b] Ros not described; L 50  3.5 cm, lamina narrowly triangular, with pungent tip, densely pale-lepidote abaxially, margins densely finely serrate, Sp to 2 mm; Inf 1 m, simple; peduncle stout, furfuraceous-lepidote; peduncular Bra broadly ovate, acuminate, with pungent tip, much shorter than the internodes, entire; fertile

957

Inf part subdense but interrupted at the base, densely ferrugineous-lepidote; floral Bra spreading or reflexed, ovate, acuminate to acute, to 10 mm, equalling or shorter than the sepals; Fl somewhat erect to divergent, to 13 mm, subsessile; Sep broadly ovate-elliptic, 7 mm, obtuse, abaxially densely tomentose; Pet suberect, rhomboid, subobtuse, carinate, colour not described; St included; Fil high-connate above the common tube; Anth linear, subacute, recurved; Sty very short; Fr not described. Compared with D. edwardii in the protologue of the latter, and description completed from the tabular comparison. D. ursina L. B. Smith (Arq. Bot. Estado São Paulo 2(1): 109, t. 111, 1943). Type: Brazil, Minas Gerais (Foster 636 [GH]). — Lit: Büneker & al. (2016: with ills.). Distr: Brazil (Minas Gerais: Serra do Cipó); dry open rocky ground, 1100–1400 m; only known from a single population. I: Smith & Downs (1974: 517). [2/3a] Ros usually solitary; L usually arching and secund, to 60  3–3.5 cm, sheath suborbicular, glabrous towards the base, laxly serrulate, L lamina very narrowly triangular, margins very laxly serrate, Sp slender, curved, 3 mm, lepidote basally, glabrous elsewhere; Inf >1 m, simple or few-branched; peduncle stout, brownlanate; peduncular Bra erect, subfoliaceous (lower) to narrowly triangular (upper), longer or shorter than the internodes; fertile Inf part sublaxly flowered, very densely dark brown-lanate throughout; floral Bra broadly ovate, 8–20 mm, short-acuminate, the lower  equalling the sepals, entire; Fl spreading, (sub-) sessile; Sep elliptic, to 15 mm, acute, margins wholly obscured by the brown-lanate indumentum; Pet narrowly elliptic and without a distinct claw, only slightly exceeding the sepals, orange, lanate abaxially; St included; Fil free above the short common tube; Sty simple, very short. The inflorscences are conspicuously densely brown-lanate and typical for the D. sordida complex (Büneker & al. 2016). D. velascana Mez (in Martius, Fl. Bras. 3(3): 476, 1894). Type: Argentina, La Rioja (Hieronymus

958

& Niederlein 66 [B, CORD]). — Distr: N to N-C Argentina (Catamarca, Córdoba, La Rioja, San Luis, San Juan, Tucumán, Jujuy, Salta); rocky slopes, 750–3000 m. I: Smith & Downs (1974: 544); Subils (2009: 348). [1] Ros offsetting, forming dense colonies; L 23–50  4 cm, spreading, sheath suborbicular, 5 cm, pale, L lamina narrowly triangular, appressedly pale-lepidote abaxially, margins spinose, Sp antrorsely uncinate, 4 mm; Inf to 1 m, paniculate, lax; peduncle 60–70 cm, erect, lepidote; peduncular Bra erect, broadly ovate, lowest narrowly triangular and serrulate, apical ones apiculate, much shorter than the internodes; Inf branches very densely flowered, short; primary Bra cinereous-lepidote-pubescent, shorter than the sterile branch base; floral Bra reflexed, acuminate from a broadly ovate base, to 5 mm, puberulous; Fl subsessile, spreading, to 14 mm; Sep ovate, to 8 mm, rounded and mucronulate, glabrous and lustrous with a ferrugineouspuberulous base; Pet erect, subreniform, 10–12 mm, broadly rounded and minutely emarginate, not undulate, ecarinate, sulphur-yellow; St exserted; Fil free above the common tube; Anth sublinear, acute; Ov 6 mm, oblong and narrowly pyramidal; Sty 0.5 mm; Se 2.5 mm, with a circular wing. No other species of the genus reaches similarly high altitudes. D. velloziifolia Mez (Repert. Spec. Nov. Regni Veg. 16: 70, 1919). Type: Paraguay, Caaguazú (Hassler 9576 [B]). — Distr: SE Paraguay (Caaguazú); campos. [3a] Only known from fragmentary material; Ros dense; L to 12  0.4 cm, lamina first erect, then irregularly curved-spreading, subulate to acicular, glabrous, entire or older ones with a few minute Sp basally; Inf to 50 cm, simple (?); peduncle stout, length not described but much exceeding the leaves, glabrous; peduncular Bra very broadly ovate, acuminate, all short-laminate, much shorter than the internodes; fertile Inf part very lax at least at the base, stout, glabrous; floral Bra like the peduncular bracts but with shorter lamina, suberect to erect, to 12 mm, exceeding the flowers, violet-tinged when dry; Fl erect, to

F. Krapp and U. Eggli

11 mm; Ped short; Sep very broadly elliptic, 6 mm, rounded and with a few teeth at the apex, submembranous; Pet erect, broadly rounded and slightly emarginate, ecarinate, lemon-yellow; St much shorter than the petals; Fil free above the very short common tube, very broad and almost foliaceous; Sty very stout. D. vestita Hassler (Annuaire Conserv. Jard. Bot. Genève 20: 315, 1919). Type: Paraguay, Concepción (Fiebrig 5311 [G, F [photo]]). — Distr: Paraguay (Concepción); campos. [1/2?] Only known from fragments; Ros and L unknown; Inf with 7 branches, >60 cm, Br erect or suberect, sublaxly flowered, to 30 cm, fulvous-tomentose; primary Bra ovate, acuminate, fulvous-tomentose; floral Bra ovate, acuminate, to 11 mm, equalling the sepals, fulvous-tomentose; Fl erect to spreading, to 20 mm, subsessile; Sep oblong-ovate, 11 mm, rounded at the apex, tomentose; Pet ovate, 18 mm, subacute, erect,  carinate, pale yellow; St equalling the petals; Fil 3.5 mm (to completely?, Pinangé & al. (2017: fig. 3)) connate above the common tube; Anth linear, acute, sagittate at the base, strongly recurved; Ov pyramidal, 13 mm; Sty none. — [F. Krapp] D. vicentensis Strehl (Bromeliaceae 42(5): 13–15, figs. 10–13, 2009). Type: Brazil, Rio Grande do Sul (Strehl 1375 [HAS]). — Distr: Brazil (C-W Rio Grande do Sul: São Vicente do Sul); on rocks. [2/3] Ros acaulescent, 30- to 40-leaved, 20–25 cm ∅; L 8–12  1–1.3 cm, recurved to spreading, tough, sheath broad, 1  4 cm, white, bright, glabrous, L lamina green, sometimes reddish basally, with pungent tip, glabrous and shiny adaxially, cinereous-lepidote abaxially, sometimes densely white-lepidote basally, marginal Sp slightly hooked, predominantly retrorse, 2 mm, with brown tip and green base, 5 mm apart; Inf 70–100 cm, simple or rarely branched; peduncle erect, green to reddish, white-lepidote and lanate; peduncular Bra long-triangular, asymmetrical, few, concentrated at the base of the stem; primary Bra narrowly linear to subtriangular from an ovate base, lower ones longer than the

Dyckia BROMELIACEAE

internodes, veined, slightly serrate, upper ones shorter and  inflated and scale-like; floral Bra spreading, ovate, acute, never pungent, curved, carinate, longer than the sepals, 10–15 mm, slightly reddish, chaffy-lepidote, entire; Fl spreading at anthesis, later erect, 18–20 mm, tubular; Ped 3–5 mm; Sep oval, 8–10 mm, narrowly marginate, base inflated, ecarinate, appressed to the petals, acute, obtuse to apiculate, orangereddish; Pet free, lamina broad, 18–20 mm, ecarinate, apex roundish, yellow to reddish; St included; Fil erect, 13 mm, orange-yellow, “joined at the base” according to the protologue but degree of connation with the common tube not described; Anth 4 mm, linear, yellow; Ov 5 mm; Sty 5 mm; Sti 7 mm. D. virgata Mez (Repert. Spec. Nov. Regni Veg. 16: 68, 1919). Type: Paraguay, Presidente Hayes (Rojas 46 [B]). — Distr: C Paraguay (Presidente Hayes); ecology not recorded. I: Smith & Downs (1974: 542). [3b] Only known from fragments; Ros unknown; L 120  2.5 cm, coriaceous, lamina attenuate to a slenderly subulate apex, subglabrous, margins densely serrate basally, laxly serrate elsewhere, Sp acicular, 1.5 mm; Inf >30 cm, simple (?), slenderly virgate, very laxly many-flowered, slender, all parts including bracts, pedicels and sepals brownish-tomentellous, glabrescent; floral Bra reflexed, ovate, broadly acute, strongly convex, 4 mm, much shorter than the sepals, entire; Fl suberect to subspreading, slender, orange when dry, subsessile; Sep broadly elliptic, 5 mm, rounded and emarginate; Pet erect, 13 mm, broadly subacute, ecarinate; St exserted; Fil high-connate above the common tube; Sty short. — [F. Krapp] D. waechteri Strehl (Bromeliaceae 42(5): 15–17, figs. 14–19, 2009). Type: Brazil, Rio Grande do Sul (Strehl 1086 [HAS]). — Distr: Brazil (C-W Rio Grande do Sul: São Francisco de Assis); on rocks. [1?] Ros with 40 living leaves, to 40 cm ∅, acaulescent; L 33–38  2–2.3 cm, flexuous, long and acute, sheath 3–3.2  2.5–2.7 cm, white, white-lepidote, serrate, L lamina arching, with

959

slightly pungent tip, light green, glabrous adaxially, white-lepidote abaxially, Sp antrorse, 3–4 mm, light green with castaneous tip, 10–15 mm apart; Inf 1.6–1.7 m, compound or rarely simple; peduncle elliptical in cross section, furfuraceous to glabrous, brittle, 1.5–1.6 m, green, covered with white powder; peduncular Bra amplexicaul, narrowing abruptly, long-acuminate from a wide base, membranous, green at the base, serrate, drying from the tip; fertile Inf part lax, reddish-white; Br 3–6, at acute angles, 30–70 cm; floral Bra triangular, acute, mucronate, 8–10  5–8 mm, base green, apical 1=2 becoming brown, drying quickly, slightly lepidote; Fl erect to subspreading, 20 mm, sessile; Sep obovate and roundish, 8–11  8–11 mm, mucronate, sometimes crenate at the apex, carinate, free in the upper 1=2, green, turning yellow basally and orange or rose towards the apex, thick-textured; Pet obovate-rounded, 16–18  5–8 mm, erect and quite recurved, thick-textured, orange-rose; St becoming visible in the throat; Fil free above the common tube, yellowish; Anth orange; Ov 6–7 mm; Sty 6 mm, orange; Fr 18–20  12–15 mm, dark brown, shiny; Se discoid, 3  5 mm, 1 mm thick, with a lateral wing. — [F. Krapp] D. walteriana Leme (Phytotaxa 67: 33–35, ills., 2012). Type: Brazil, Paraná (Kranz 13 [RB, HB]). — Distr: Brazil (Paraná); on almost horizontal granitic rock outcrops; known from a single population only. [1] Ros dense, 50-leaved, propagating by basal shoots; L 75–90  3–3.4 cm, 5 mm thick, stiffly succulent, sheath inconspicuous, L lamina narrowly triangular, attenuate and slightly canaliculate towards the pungent apex, suberect-arcuate, acuminate, green, abaxially finely veined, densely white-lepidote, margins laxly spinose, Sp narrowly triangular, predominantly antrorse-uncinate, 3–10 mm, densely white-lepidote, with castaneous tip, 15–25 mm apart; Inf 2.1 m, broadly 2 branched; peduncle erect, 80 cm, greenish-purple, subdensely to sparsely pale castaneous-lanate and densely covered with white wax; peduncular Bra narrowly triangular, acuminate and spinescent, stramineous,

960

F. Krapp and U. Eggli

canaliculate towards the apex, 60–110  20–25 mm, subdensely white-lepidote, veined, densely spinulose; fertile Inf part dense, 120 cm, rachis straight to slightly flexuous, greenishpurple, subdensely pale castaneous-lanate to glabrous and densely covered with white wax; Br 12, laxly arranged, densely 100- to 140flowered, 40–60 cm, with a 4–8 cm long sterile base, straight, greenish to greenish-purple, densely castaneous-lanate, soon glabrous and densely covered with white wax, terminal part of the main axis not distinct from the branches; primary Bra like the upper peduncular bracts, subspreading, 30–50  10–20 mm; floral Bra triangular-ovate to broadly triangular, acute, shorter than the sepals, 3–5  3–4 mm, stramineous, densely castaneous-lanate, veined; Fl subverticillate,  spreading, 16–17 mm, broadly campanulate, unscented; Ped 1 mm, green, densely castaneous-lanate; Sep orbicular, 3.5–5  4–5 mm, obtuse-emarginate, ecarinate, strongly convex, greenish, obscurely castaneouslanate, entire; Pet elliptic to oblong-elliptic, 12  6–7 mm, obtuse-emarginate, ecarinate, yellow, glabrous; St exserted at anthesis; Fil free above the 1–2 mm high common tube, 11–12 mm, yellow; Anth suboblong, 3 mm, slightly recurved at anthesis, base bilobed, acute; Ov narrowly ovoid, 6–7  3 mm, yellow; Sty 5–7 mm, yellow, Sti 1 mm, yellow, margins crenulate; Fr subglobose, acute, 14  10 mm, dark castaneous, glossy; Se subtriangular, strongly complanate, 3  2 mm. Compared with D. agudensis and D. selloa in the protologue.

stout, articulate; Sep broadly ovate, 6–8 mm, acute,  straight; Pet suberect, broadly obovate, obtuse, undulate,  orange; St included; Fil highconnate above the common tube; Anth triangular, acute, recurved towards the apex; Sty short. — [F. Krapp]

D. warmingii Mez (in Martius, Fl. Bras. 3(3): 481, 1894). Type: Brazil, Minas Gerais (Warming s.n. [C, F [photo]]). — Distr: Brazil (C Minas Gerais: Lagoa Santa); savanna, terrestrial. [3b] Only known from fragments; Ros and L unknown; Inf probably >50 cm, simple; peduncular Bra remote, broadly ovate, acuminate, entire; fertile Inf part lax, finely and fugaciously whitish-furfuraceous; floral Bra spreading or reflexed, like the peduncular bracts, nearly equalling the sepals (lower); Fl spreading or reflexed at anthesis, to 15 mm; Ped 2–3 mm,

Baker, J. G. (1889) Handbook of the Bromeliaceae. London (GB): G. Bell & Sons. https://doi.org/ 10.5962/bhl.title.51681. Bernardello, L. M. [& al. 1991], Galetto, L. & Juliani, H. R. (1991) Floral nectar, nectary structure and pollinators in some Argentinean Bromeliaceae. Ann. Bot. (Oxford), n.s. 67(4): 401–411. https://doi.org/10.1093/ oxfordjournals.aob.a088174 Braun, P. J. [& al. 2008], Esteves Pereira, E. & Scharf, U. (2008) Dyckia joanae-marcioi. Eine neue, sukkulente Bromelie aus dem nördlichen Minas Gerais, Brasilien. Bromelie 2008(1): 33–46, ills., map. Braun, P. J. (2018) Xeromorphe Bromelien aus Brasilien (1)/Xeromorphic Bromeliads from Brazil (1): Dyckia. Bromelie 2018(2): 68–77, ills.

D. weddelliana Baker (Handb. Bromel., 132, 1889). Type: Brazil, sine loco (Weddell 2584 [P, GH [photo]]). — Distr: C Brazil (Minas Gerais?, Goias?); savanna, terrestrial. I: Smith & Downs (1974: 542). [2] Ros not described; L 30–40  2–3 cm, sheath suborbicular, 4 cm, brown, glabrous, L lamina narrowly triangular, attenuate, minutely lepidote abaxially, margins laxly serrate, Sp straight, slender, 2 mm; Inf 1.2–1.5 m, simple or few-branched; peduncle stout, soon glabrous; peduncular Bra erect, the upper remote, broadly ovate, acute, entire; fertile Inf part elongate, subdensely many-flowered, ferrugineousfurfuraceous; floral Bra ovate with triangular apices, reflexed, 9 mm; Fl suberect to spreading, 11 mm; Ped 3 mm, stout; Sep elliptic, 6 mm, obtuse; Pet erect or suberect, obovate, broadly rounded,  carinate, colour not described; St included; Fil high-connate above the common tube; Anth linear, acute, strongly recurved; Sty as long as the ovary. Described on the base of material without origin other than “Central Brazil”, and insufficiently known. — [F. Krapp]

References

Dyckia BROMELIACEAE Braun, P. J. & Esteves Pereira, E. (1991) Dyckia braunii Rauh. Eine neue, weissbereifte Blattsukkulente aus Zentralbrasilien. Kakt. and. Sukk. 42(3): 58–60, ills. Braun, P. J. & Esteves Pereira, E. (2005) Succulent and xeromorphic Bromeliads of Brazil. Part 1. Dyckia marnier-lapostollei L. B. Smith. Cact. Succ. J. (US) 77(6): 284–292, ills. https://doi.org/10.2985/ 0007-9367(2005)77[284:SAXBOB]2.0.CO;2. Braun, P. J. & Esteves Pereira, E. (2006a) Succulent and xeromorphic Bromeliads of Brazil. Part 2. Orthophytum horridum, Dyckia paucispina and Bromelia estevesii. Cact. Succ. J. (US) 78(1): 31–35, ills. https://doi.org/ 10.2985/0007-9367(2006)78[31:SAXBOB]2.0.CO;2. Braun, P. J. & Esteves Pereira, E. (2006b) Succulent and xeromorphic Bromeliads of Brazil. Part 3. Encholirium maximum, Orthophytum burle-marxii and Dyckia goehringii. Cact. Succ. J. (US) 78(4): 160–164, ills. https://doi.org/10.2985/0007-9367(2006)78[160:SAX BOB]2.0.CO;2. Braun, P. J. & Esteves Pereira, E. (2009a) Dyckia edwardii P. J. Braun & al. Kakt. and. Sukk. 60(7): xxv–xxvi (index card 2009/13), ills. Braun, P. J. & Esteves Pereira, E. (2009b) Dyckia richardii P. J. Braun & Esteves. Kakt. and. Sukk. 60(7): xxvii–xxviii (index card 2009/14), ills. Büneker, H. M. [& al. 2016], Pureza Soares, K. & Coelho de Assis, L. (2016) The Dyckia sordida complex (Bromeliaceae, Pitcairnioideae) and a new species from Minas Gerais, Brazil. Phytotaxa 244(1): 57–68, key, ills. https://doi.org/10.11646/phytotaxa.244.1.4. Butcher, D. (2010) Dyckia estevesii revisited. J. Bromeliad Soc. 60(2): 82–84, ills. Dorneles, M. P. [& al. 2014], Oliveira, J. M. S. de & CantoDorow, T. S. do (2014) Dyckia racinae L. B. Sm. (Bromeliaceae): Morphological description emphasizing the reproductive structures. Iheringia, Bot. 69(2): 397–303, ills. Esteves Pereira, E. & Gouda, E. J. (2017) A new Dyckia (Bromeliaceae) from the south region of Goiás, Brazil: Dyckia piracanjubensis. J. Bromeliad Soc. 66(3): 172–181, ills. Fagundes, N. F. & Mariath, J. E. D. (2010) Morphoanatomy and ontogeny of fruit in Bromeliaceae species. Acta Bot. Brasil. 24(3): 765–779, ills. https://doi.org/ 10.1590/S0102-33062010000300020. Guarçoni, E. A. E. [& al. 2012], Sartori, M. A. & Paula, C. C. de (2012) Dyckia inflexifolia (Bromeliaceae), a new species from Brazil. Ann. Bot. Fenn. 49: 407–411, ills. Guarçoni, E. A. E. [& al. 2014], Alves Azevedo, A. & Ferreira da Costa, A. (2014) Dyckia sulcata (Bromeliaceae), a new species from Minas Gerais, Brazil, with notes on leaf anatomy. Phytotaxa 188(3): 169–175, ills. https://doi.org/10.11646/phytotaxa.188.3.6. Hmeljevski, K. V. [& al. 2011], Reis, A., Montagna, T. & Sedrez dos Reis, M. (2011) Genetic diversity, genetic drift and mixed mating system in small subpopulations of Dyckia ibiramensis, a rare endemic bromeliad from southern Brazil. Conservation Genet. 12: 761–769. https://doi.org/10.1007/s10592-011-0183-3.

961 Klein V., L. F. & Klein, J. (2014) Die Bromelien in den atlantischen Wäldern der Nord-Ost-Küste von Rio Grande do Sul/Brasilien — Teil 4/Schluss. The Bromeliads in the Atlantic forest of the northeast coast of Rio Grande do Sul/Brazil — part 4/final part. Bromelie 2014(2): 74–85, ills. Krapp, F. (2013) Phylogenie und Evolution der Gattung Dyckia (Bromeliaceae). Kassel (DE): Universität Kassel; unpublished Ph.D. thesis. Krapp, F. [& al. 2014], Pinangé, D. S. de B., BenkoIseppon, A. M., Leme, E. M. C. & Weising, K. (2014) Phylogeny and evolution of Dyckia (Bromeliaceae) inferred from chloroplast and nuclear sequences. Pl. Syst. Evol. 300(7): 1591–1614, ills., maps. https:// doi.org/10.1007/s00606-014-0985-0. Krieck, C. (2008) Ecologia reprodutiva de Dyckia encholirioides var. encholirioides (Gaud.) Mez. (Bromeliaceae) em costões oce^anicos em Florianópolis, Santa Catarina. Florianópolis (BR): Universidad Federal de Santa Catarina, Departamento de Fitotecnia; unpublished M.Sc. thesis. Lawn, G. (2011) Dyckia ‘Leopoldo Witeck’. J. Bromeliad Soc. 61(3): 136–137, ills. Leme, E. M. C. & Miranda, Z. de J. G. (2009) Studies on Dyckia from Central Brazil — part II. Two sweetly fragrant species from Goiás. J. Bromeliad Soc. 59(2): 71–79, ills. Machado, T. M. [& al. 2016], Forzza, R. C. & Stehmann, J. R. (2016) Bromeliaceae from Caparaó National Park, Minas Gerais/Espírito Santo states, Brazil, with notes on distribution and conservation. Oecol. Australis 20(2): 133–146, ills. https://doi.org/10.4257/oeco.2016.2002.10. Mendes, S. P. [& al. 2012], Costa, C. G. & Toni, K. L. G. de (2012) Androecium development in the bromeliad Dyckia pseudococcinea L. B. Sm. (Pitcairnioideae — Bromeliaceae), an endangered species endemic to Brazil: Implications for conservation. Flora (Jena) 207(8): 622–627, ills. https://doi.org/10.1016/j.flora.2012.06.016. Mez, C. (1891–1892) Bromeliaceae. In: Martius, C. F. P. von & al. (eds.): Flora Brasiliensis, 3(3): 173–286, tt. 51–62 (= fasc. 110, 1891); pp. 281 [sic]–430, tt. 63–80 (= fasc. 112, 1892). Leipzig (DE): F. Fleischer. http://biodiversitylibrary.org/page/136410. Monteiro, R. F. & Forzza, R. C. (2016) Flora das cangas da Serra dos Carajás, Pará, Brasil: Bromeliaceae. Rodriguésia 67(5): 1253–1265, ills., keys. https://doi. org/10.1590/2175-7860201667523. Paggi, G. M. [& al. 2015], Louzada, R. B., Ishii, I. H., Takahasi, A., Arruda, R. C. O. & Lorenz-Lemke, A. P. (2015) Rediscovering Dyckia excelsa (Bromeliaceae) in Mato Grosso do Sul, Brazil: Taxonomy, geographic distribution, and notes on leaf anatomy. Syst. Bot. 40(1): 129–135, ills., key. https://doi.org/10.1600/ 036364415X686422. Pinangé, D. S. B. [& al. 2017], Krapp, F., Zizka, G., Silvestro, D., Leme, E. M. C., Weising, K. & BenkoIseppon, A. M. (2017) Molecular phylogenetics, historical biogeography and character evolution in Dyckia (Bromeliaceae, Pitcairnioideae). Bot. J. Linn. Soc. 183: 39–56, ills. https://doi.org/10.1111/boj.12489.

962 Pompelli, M. F. & Guerra, M. P. (2004) Ex situ conservation of Dyckia distachya: An endangered Bromeliad from South Brazil. Crop Breed. Appl. Biotechnol. 4(3): 273–279, ills., maps. Pompelli, M. F. & Guerra, M. P. (2005) Micropropagation enables the mass propagation and conservation of Dyckia distachya Hassler. Crop Breed. Appl. Biotechnol. 5: 117–126, ills. Pompelli, M. F. [& al. 2006], Fernandes, D. & Guerra, M. P. (2006) Germination of Dyckia encholirioides (Gaudichaud) Mez var. encholirioides under saline conditions. Seed Sci. Technol. 34(3): 759–763, ills. Rauh, W. (1985) Bromelienstudien. I. Neue und wenig bekannte Arten aus Peru und anderen Ländern (XVII. Mitteilung). Trop. subtrop. Pfl.-welt 53: 1–62, ills. Rauh, W. & Gross, E. (1988) Bromelienstudien. I. Neue und wenig bekannte Arten aus Peru und anderen Ländern (XX. Mitteilung). Trop. subtrop. Pfl.-welt 65: 5–75, ills. Rauh, W. & Gross, E. (1991) Bromelienstudien. I. Neue und wenig bekannte Arten aus Peru und anderen Ländern (XXII. Mitteilung). Trop. subtrop. Pfl.-welt 79: 1–31, ills. Reitz, R. (1983) Bromeliáceas e a malária-bromélia endémica. In: Reitz, R. (ed.): Flora ilustrada catarinense, I parte as plantas, fascículo BROM. Itajaí (BR): Herbario “Barbosa Rodrigues”. Rogalski, J. M. [& al. 2009], Reis, A., Reis, M. S. dos & Hmeljevski, K. V. (2009) Biologia reprodutiva da reófita Dyckia brevifolia Baker (Bromeliaceae), no Rio Itajaí-Açu, Santa Catarina, Brazil. Revista Brasil. Bot. 32(4): 691–702, ills. https://doi.org/10.1590/ S0100-84042009000400008. Sachs, H. (2011) Auf der Suche nach/Searching for Dyckia delicata. Bromelie 2010(3): 118–121, ills. Santos-Silva, F. [& al. 2015], Antunes Mastroberti, A. & Araujo Mariath, E. de (2015) Capsule structure in three species of Dyckia (Bromeliaceae): Ontogenetic and taxonomic issues. J. Torrey Bot. Soc. 142(3): 249–257, ills. https://doi.org/10.3159/TORREY-D-14-00002.1. Saul, J. (2018) Hybriden von der Fensterbank. Hybrids from the window sill. Bromelie 2018(2): 65–67, ills. Schütz, N. [& al. 2016], Krapp, F., Wagner, N. & Weising, K. (2016) Phylogenetics of Pitcairnioideae s.s. (Bromeliaceae): Evidence from nuclear and plastid DNA sequence data. Bot. J. Linn. Soc. 181(3): 323–342, maps, ills. https://doi.org/10.1111/boj.12403. Silva, A. L. L. da [& al. 2007], Dornelles, E. B., Bisognin, D. A., Franco, E. T. H. & Horbach, M. A. (2007) Micropropagation of Dyckia agudensis Irgang & Sobral

F. Krapp and U. Eggli — An extinction threatened Bromeliad. Iheringia, Bot. 62(1–2): 39–43, ills. Siqueiro Filho, J. A. & Leme, E. M. C. (2007) Fragments of the Atlantic forest of Northeast Brazil: Biodiversity, conservation and the Bromeliads. Rio de Janeiro (BR): Andrea Jakobsson Estúdio. Smith, L. B. & Downs, R. J. (1974) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 1: Pitcairnioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/sta ble/4393694. Subils, R. (2009) Bromeliaceae, Bromeliáceas. In: Kiesling, R. (ed.): Flora de San Juan, IV: 340–363, ills., keys. Mendoza (AR): Zeta Editores. Vásquez, R. C. & Ibisch, P. L. (2012) Die Bromelien Boliviens (XI): Dyckia barthlottii — eine neue Art aus dem Chaco-Trockenwald Boliviens. The Bromeliads of Bolivia (XI): Dyckia barthlottii — a new species from the Chaco Dry Forest of Bolivia. Bromelie 2012(3): 123–127, ills. Vesprini, J. L. [& al. 2003], Galetto L. & Bernardello, G. (2003) The beneficial effect of ants on the reproductive success of Dyckia floribunda (Bromeliaceae), an extrafloral nectary plant. Canad. J. Bot. 81(1): 24–27, ills. https://doi.org/10.1139/b03-003. Voltolini, C. H. [& al. 2009], Reis, A. & Santos, M. (2009) Leaf morphoanatomy of the rheophyte Dyckia distachya Hassler (Bromeliaceae). Revista Brasil. Bioci. 7(4): 335–343, ills. Vosgueritchian, S. B. & Buzato, S. (2006) Reprodução sexuada de Dyckia tuberosa (Vell.) Beer (Bromeliaceae) e interação planta-animal. Revista Brasil. Bot. 29(3): 433–442, ills. https://doi.org/10.1590/ S0100-84042006000300010. Waldemar, C. C. & Irgang, B. E. (2003) A ocorrência do mutualismo facultativo entre Dyckia maritima Backer [sic!] (Bromeliaceae) e o cupim Cortaritermes silvestrii (Holmgren), Nasutitermitinae, em afloramentos rochosos no Parque Estadual de Itapuã, Viamão, RS. Acta Bot. Brasil. 17(1): 37–48, ills., map. https:// doi.org/10.1590/S0102-33062003000100004. Wiesbauer, M. B. (2008) Biologia reprodutiva e diversidade genética de Dyckia distachya Hassler (Bromeliaceae) como subsídio para conservação e reintrodução de populações extintas na naturaleza. Florianópolis (BR): Universidade Federal de Santa Catarina, Centro de Ciências Agrárias; unpublished thesis. Winkler, S. (1982) Die Bromeliaceae von Rio Grande do Sul (S-Brasilien). Doc. Nat. 3: 1–81, keys, 5 tt.

Dycktia BROMELIACEAE N. Schütz and F. Krapp

Dycktia D. A. Beadle (Prelim. List. Cult. Grex Bromeliad, 82, 1991). — Distr: Cultivated only. = Dyckia  Hechtia. The only known cross is D. ‘Dark Chocolate’  H. epigyna, obtained by

O. Ferris; it remained unnamed and appears to have first been mentioned in an unpublished manuscript (L. B. Smith, Manuscript of Bromeliad Hybrids and Cultivars, 1984).

N. Schütz (*) Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany e-mail: [email protected] F. Krapp Guxhagen, Germany e-mail: fl[email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_88

963

Encholirium BROMELIACEAE F. Krapp

Encholirium Martius ex Schultes & Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): lxviii, 1233, 1830). Type: Encholirium spectabile Martius ex Schultes & Schultes fil. — Pitcairnioideae — Lit: Smith & Downs (1974: 191–199, Fl. Neotropica); Forzza (2005: monograph); Forzza & Zappi (2011: partial key); Krapp & al. (2014: phylogeny). Distr: E Brazil. Etym: Gr. ‘enchos’, spear; and Gr. ‘leirion’, lily; presumably for the long slender inflorescence of some species. Incl. Encholirion hort. (nom. inval., ICN Art. 61.1). Perennial terrestrial rosette plants; Ros solitary or more frequently forming clumps; rhizome short, erect, sometimes with lateral branches; L densely rosulate, sheath wide and imbricate, L lamina green, yellowish, reddish or nigrescent, often cinereous, triangular-lanceolate, marginal Sp usually well developed, rarely margins serrate or entire; Inf terminal, tall, unbranched or rarely fewbranched; peduncular Bra numerous, lower ones foliaceous, upper ones small and often withering; Fl pedicellate, hermaphrodite, not very conspicuous; Ped usually distinct to very long in the longipedicellatum-complex; Sep green to yellowish-green, yellow, vinaceous or pinkish, convolute

F. Krapp (*) Guxhagen, Germany e-mail: fl[email protected]

or imbricate, symmetrical or sometimes asymmetrical, usually glabrous, free; Pet cream, creamgreen, green, green-yellow, yellow or slightly vinaceous, imbricate or not, symmetrical or slightly asymmetrical, free; St inserted or exserted; Fil free or connate at the base, rarely adnate to the petals; Ov completely superior or partly inferior; Sty exserted or included; Sti conduplicate-spiral with either laminar or compact lobes; Fr globose stout capsules with usually persistent floral remains; Se numerous, brown, flattened, surrounded by a continuous membranous margin, with a single rounded, truncate, long-caudate or falciform wing. A genus of 35 species, of which c. 1=3 were described subsequently to the taxonomic revision of Forzza (2005). No formal infrageneric classification has been published, but 8 species form the so called longipedicellatum-complex, characterized by long pedicels (short in E. biflorum and E. reflexum), imbricate petals and included stamens. Encholirium species are typically strongly xeromorphic plants occurring in rocky habitats. They are distributed in E Brazil, from Piauí in the N to Mato Grosso do Sul in the S. Half of all species are restricted to the Serra do Espinhaço. The diversity centre is on the Diamantina plateau and in the Serra do Cipó. Nearly all species are restricted to rock outcrops in Cerrado and Caatinga vegetations; only E. gracile and E. horridum occur on inselbergs within the Atlantic Forest, while the widespread E. spectabile is also found in the transition zone between savanna and Atlantic Forest. Compared with the typically

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_81

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narrowly endemic and rare species of Dyckia, many Encholirium species are more common and occur in larger populations. Evolution and phylogeny: A dated phylogeny based on plastid DNA sequence data shows a common origin of Encholirium plus Dyckia, and this clade separated from Deuterocohnia 5.6 mybp (Krapp & al. 2014). The monophyletic genus Dyckia is imbedded within what is currently treated as Encholirium and arose 4–4.6 mybp. This topology with Encholirium as a grade at the base of a paraphyletic Dyckia is also supported by nuclear DNA data (Krapp & al. 2014, Schütz & al. 2016) as well as by morphological characters (Forzza 2005). Up to now, no single synapomorphic morphological character has been described for Encholirium, although it is easily distinguished morphologically from Dyckia. Strictly speaking, the two genera would have to be merged into a single genus Dyckia. This merger is, however, not advocated for the time being due to insufficient sampling in the molecular studies, esp. for Encholirium, and the possibility to split up Encholirium must also be explored. Ecology: Encholirium species are commonly pollinated by hummingbirds (Forzza 2005, Leal & al. 2006: 383, Aguilar-Rodríguez & al. 2014: 159). Some species are also or exclusively pollinated by bats, e.g. E. subsecundum or E. splendidum. Queiroz & al. (2016) recorded a mixed pollination system in E. spectabile, involving primarily bats, an opossum and hummingbirds, with occasional visits by passerine birds and hawkmoths. Christianini & al. (2013) regard hummingbird pollination as ancestral for the genus. The following names are of unresolved application but are referred to this genus: Encholirium diamantinae Rauh (s.a.) (nom. inval., ICN Art. 29.1); Encholirium sabinae Rauh (s.a.) (nom. inval., ICN Art. 29.1); Encholirium sanguinolentum Hort. Paris ex C. Chevalier (1930) (nom. inval., ICN Art. 36.1c). E. agavoides Forzza & Zappi (Kew Bull. 66 (2): 282, ills. (p. 283, 285), 2011). Type: Brazil,

F. Krapp

Minas Gerais (Forzza & al. 5475 [RB, CEPEC, K, MBM, SPF, US]). — Distr: Brazil (Minas Gerais: Serra do Cipó); quartzitic sandstones; narrowly endemic. I: Zappi & Taylor (2011). Flowering plants (15–) 30–44 cm tall, forming dense mats; rhizome short; Ros 6–10 cm ∅; L suberect to patent, sheath 1.2–1.5  2.5–5.7 cm, entire, L lamina 3.2–3.7  2–3 cm, triangular, cinereous, densely lepidote, margins with patent Sp 3–4 mm long; peduncle 12–22 cm, green, erect, glabrous; peduncular Bra exceeding the internodes, 6–11 cm, erect, lanceolate to ovatelanceolate, green to stramineous, apex acute to attenuate, entire to slightly serrate at the base, glabrous, clasping the peduncle; fertile part of the Inf 6–18 cm, simple, dense, rachis hidden by the flowers, green, glabrous; floral Bra exceeding the pedicels, 8–15 ( 22)  5–7 ( 12) mm, glabrous, ovate-acuminate, completely stramineous or with green base, entire or slightly serrulate; Ped 2–3 mm, green; Sep 7–9  3–5 mm, green, ovate, acute, entire, not overlapping, symmetrical, glabrous; Pet 8–10  3–4 mm, green, elliptic, entire, not imbricate, symmetrical, glabrous, apex obtuse; St 11–14 mm, exserted; Fil free, green; Ov 3–5 mm, green; Sty 3–5 mm, exserted, green; Sti green; Fr 7–9 mm, globose, castaneous; Se  2 mm, with falciform wing. The flowers are similar to those of E. magalhaesii. E. ascendens Leme (Selbyana 30(2): 129–131, fig. 1, 2010). Type: Brazil, Minas Gerais (Leme & al. 7451 [HB, RB, SEL, VIC]). — Distr: Brazil (Minas Gerais); ecology not reported. Flowering plants 2.4–3 m tall, in clumps or solitary; L suberect-arcuate to spreading, sheath much broader than the lamina, L lamina 70  3–4 cm, narrowly triangular, apex attenuate and long-caudate, completely greyish-green or reddish towards the apex, abaxially densely whitelepidote, adaxially inconspicuously whitelepidote to glabrescent, basally flat and apically canaliculate, armed with Sp 3–10 mm long; peduncle 1.1–1.6 m, erect, rigid, glabrous; peduncular Bra foliaceous to subfoliaceous, exceeding the internodes, long attenuate-caudate, white-lepidote mainly abaxially, laxly and coarsely spinose;

Encholirium BROMELIACEAE

fertile part of the Inf 100–125 cm, compound and with 4–12 Br 27–55 cm long, without terminal raceme, densely many-flowered, erect, glabrous, rachis stout, straight; lower primary Bra subfoliaceous, long-filiform from a subtriangular base, suberect to reflexed, laxly spinose, upper primary Bra much reduced, narrowly triangular, attenuate, recurved, inconspicuously spinulose to subentire; floral Bra exceeding the sepals, 10  4 mm, long filiform-caudate from a suborbicular-triangular base, glabrous, inconspicuously crenulate to entire, recurved; Fl subspreading; Ped indistinct, 2–3 mm; Sep  7  6–7 mm, broadly ovate, apex obtuse and apiculate, margins remotely and irregularly crenulatedenticulate mainly near the apex, partially imbricate, symmetrical, glabrous; Pet 12–13  7–8 mm, colour not described, ovate, apex obtuse, entire, partially imbricate, symmetrical, glabrous; St  19 mm, exserted; Fil almost free; Ov  7 mm; Sty  5 mm, slightly shorter than the anthers; Sti subcapitate; Fr 17–19  8–10 mm; Se subcuneate. Similar to the poorly known E. belemii, but distinguished by taller stature, different floral bracts, shorter pedicels and smaller flowers. E. belemii L. B. Smith & Read (Bradea 5(27): 301–302, fig. 5, 1989). Type: Brazil, Minas Gerais (Belem 3797 [US, CEPEC, IAN, NY, UB]). — Distr: Brazil (Minas Gerais); on limestone rocks. A dubious name, only reliably known from two herbarium specimens, and without recent collections. The original description is based on very young inflorescences and is inadequate for positive identification (Forzza 2005). E. biflorum (Mez) Forzza (Bol. Inst. Bot. (São Paulo) 23(1): 16, ills. (p. 22), 2005). Type: Brazil, Minas Gerais (Glaziou 19919 [B, F [photo], P]). — Lit: Cavallari & al. (2006: genetics). Distr: Brazil (Minas Gerais); sandy and rocky soils, known from a single population only.  Dyckia biflora Mez (1894). Flowering plants 11–32 cm tall, solitary or forming small clumps; rhizome without conspicuous branches; Ros 4–11 cm ∅; L secund, sheath 0.5–1.2  1–4.2 cm, L lamina 3–9.5  0.7–1.5 cm, abaxially cinereous, adaxially green, entire, rarely

967

with Sp  1 mm long; peduncle 0.7–22 cm, castaneous to slightly vinaceous, erect, glabrous; peduncular Bra shorter than the internodes (the basal ones rarely longer), 1–3.2 cm, erect, lanceolate to ovate-lanceolate from a broadly ovate base, acute to acuminate, entire, apex lepidote, base sparsely lepidote; fertile part of the Inf 2.5–8.2 cm, simple, few-flowered, lax, rachis castaneous to slightly vinaceous, glabrous; floral Bra shorter than the flowers but exceeding the pedicels, 11–16  8–12 mm, stramineous, ovate, acute to acuminate, entire, glabrous; Fl patent to slightly reflexed; Ped 4–7 mm, glabrous; Sep 10–14  5–7 mm, green, ovate, apex obtuse, entire, not imbricate, symmetrical, glabrous; Pet 11–15  4–6 mm, colour not described, obovate, apex rounded, entire, symmetrical, imbricate, glabrous; St  8 mm, included; Fil free, slightly adnate to the petals; Ov  4 mm; Sty  1 mm, included; Fr 12–14 mm; Se 3–4 mm, with rounded wings. Belongs to the longipedicellatum-complex. The species is characterized by the very small stature and inflorescences with only 2–5 flowers. E. brachypodum L. B. Smith & Read (Bradea 5(27): 302, fig. 8, 1989). Type: Brazil, Bahia (Read & Daniels 3442 [RGB, US]). — Distr: Brazil (Bahia); granite outcrops in Caatinga or less frequently in Campo Rupestre vegetations. I: Forzza (2005: 35). Flowering plants 1.5–2.3 m tall, forming large clumps; rhizome branched; Ros 70–100 cm ∅; L erect-patent, sheath 4.2–4.7  5.4–6.5 cm, L lamina 44–82  2.3–4.2 cm, both faces cinereous, margins armed with Sp 4–10 mm; peduncle 0.9–1.5 m, green or castaneous, erect, sparsely lepidote to glabrescent; middle peduncular Bra exceeding the internodes, 8.7–17.5 cm, erect, patent or reflexed, cinereous, triangular-lanceolate to lanceolate from a broad-ovate base, apex awned, margins spinose, lepidote, upper peduncular Bra shorter than the internodes, 2.6–5.2 cm, erect to patent or reflexed, stramineous to slightly cinereous, lanceolate, acute to acuminate, entire to slightly serrate, sparsely lepidote to glabrescent; fertile part of the Inf 11–37 cm, simple, manyflowered, dense, rachis covered by the flowers, green to brown, glabrous; floral Bra equalling or

968

shorter than the flowers, 8–15  8–12 mm, castaneous to nigrescent, broadly ovate to obovate, rarely lanceolate, apex rounded to acuminate, entire or minimally serrulate, glabrous; Ped almost none; Fl patent, subsessile; Sep 7–14  6–8 mm, greenish-cream, ovate, apex rounded, entire, not imbricate, symmetrical, glabrous; Pet 13–20  6–7 mm, greenish-cream, ovate, apex rounded, entire, partially imbricate, symmetrical, glabrous; St 18–22 mm, exserted; Fil connate, adnate to the petals; Ov 12–14 mm; Sty 7–11 mm, exserted; Sti lobes laminar; Fr 22–27 mm; Se 3–4 mm, with falciform wings. Easiliy identified by the sessile flowers. E. bracteatum P. J. Braun & Esteves (Bromelie 2017(3): 105–111, ills., 2017). Type: Brazil, Bahia (Esteves Pereira 403 [UFG 18175]). — Distr: Brazil (Bahia: Wanderley); bare bambui limestone outcrops. Flowering plants to 1.85 m tall, clump-forming with short rhizomes; Ros to 50 cm ∅; L evenly spreading to somewhat laterally upcurved, stiff and succulent, to 6 mm thick, sheath 5.5  8 cm, not further described, L lamina to 35  3.4 cm, narrowly triangular with narrow tip-region, green but both faces densely silvery grey-white-lepidote, Sp retrorse, spreading or antrorse, dark green to dark brown, pungent, 6  4 mm, 0.9–2.7 cm distant; peduncle 0.95–1.5 m, green, erect, glabrous, branched but branches sometimes abortive; peduncular Bra similar to foliage leaves but increasingly smaller upwards, to 23 cm, densely grey-silvery-lepidote, uppermost bracts chestnutbrown to straw-coloured; fertile part of the Inf 48 cm, with several erect branches, very densely many-flowered, fertile parts almost cone-like before anthesis because of the imbricate floral bracts; floral Bra very obvious, broadly obovate, stiff and succulent, carinate, 23  17 mm, glossy dark brown; Fl  35  11 mm, ascending to porrect, widely open at anthesis; Ped  7 mm, cream-coloured, maculate; Sep 19  22 mm, dark brown to reddish-rose, maculate, oblong-ovate, fused for ½ of their length, glabrous; Pet  26  5 mm, pale cream-coloured changing to dark brown towards the tips, dorsally brownmaculate, oblong-ovate, obtuse, tips spreading to

F. Krapp

recurved at full anthesis; St  23 mm, exserted; Fil white-greenish, maculate; Anth  7 mm, fleshy, yellow; Ov  18  5 mm, pale green; Sty  20 mm, green to whitish; Sti connivent, dirty greenish-white; Fr size not described, pale creambrown to brown, becoming stramineous with age; Se not described. Similar to E. erectiflorum, E. fragae and E. horridum, esp. because of the inflorescence structure. — [U. Eggli]. E. bradeanum L. B. Smith (Smithsonian Misc. Collect. 126: 26, 41, fig. 2, 1955). Type: Brazil, Minas Gerais (Smith & Brade 5652 [US, RB]). — Distr: Brazil (Minas Gerais); ecology not described. Flowering plants >70 cm tall; L sheath small and inconspicuous, L lamina 24  0.4 cm, both faces covered with pale appressed scales, margins laxly armed with mostly opposite slender curved Sp 5 mm long; peduncle 8 mm in ∅, glabrous at least with age; peduncular Bra suberect, lower ones foliaceous, much exceeding the internodes, upper ones small, shorter than the internodes, entire; fertile part of the Inf 20 cm, simple, many-flowered, dense except near the base, rachis glabrous at least with age; floral Bra much exceeding the pedicels, narrowly triangular; Fl spreading; Ped 6 mm, stout, sulcate; Sep 5 mm, broadly ovate; Pet and St probably free, colour not described; Fr 9 mm, ovoid, stout, dark castaneous; Se with falciform wings. Insufficiently known and not found again in the wild. E. ctenophyllum Forzza & Zappi (Kew Bull. 66(2): 286, ills. (p. 284–285), 2011). Type: Brazil, Minas Gerais (Forzza & al. 5455 [RB, CEPEC, K, MBM, NY, SPF, US]). — Distr: Brazil (Minas Gerais: Serra do Cipó); quartzitic sandstones; narrowly endemic. Flowering plants 50–70 cm tall; rhizome short; Ros 15–28 cm ∅; L patent to recurved, sheaths 1.5–2.5  2–2.7 cm, entire, L lamina 7–16  0.4–1 cm, narrowly lanceolate, greyish to vinaceous, lepidote, armed with mostly patent Sp 6–8 mm long; peduncle 35–45 cm, green, erect, glabrous; lower peduncular Bra foliaceous,

Encholirium BROMELIACEAE

exceeding the internodes, erect, lanceolate, apex acute to attenuate, cinereous to vinaceous, strongly spinose, lepidote, upper peduncular Bra exceeding the internodes, erect, broadly ovate, apex acute, stramineous, margins serrate, glabrous; fertile part of the Inf 16–25 cm, simple, dense, rachis green, glabrous; floral Bra exceeding the pedicels, 12–32  5–10 mm, broadly ovate, apex long attenuate, completely stramineous or with green base, margins serrulate, glabrous; Ped 2–3 mm; Sep 8–12  3–5 mm, green, ovate, acute, entire, not imbricate, symmetrical, glabrous; Pet 10–13  3–5 mm, green, elliptic, crenate at the apex, entire, not imbricate, symmetrical, glabrous; St 14–16 mm, exserted; Fil free, green; Ov 3–5 mm, green; Sty 5–8 mm, green, exserted; Sti green; Fr 10–12 mm, globose, castaneous; Se  3 mm, with falciform wings. Similar to E. magalhaesii and E. subsecundum. E. diamantinum Forzza (Nordic J. Bot. 30: 157, figs. 2, 3F–H, 2012). Type: Brazil, Minas Gerais (Forzza & al. 4858 [RB, K, MBM, NY, SPF]). — Distr: Brazil (Minas Gerais: Diamantina plateau); rock outcrops; only known from 2 populations. Flowering plants 0.8–1.3 m tall, solitary or in small clumps; rhizome short; Ros 15–30 cm ∅; L reflexed, sheaths 2.5–3.2  3.8–5.2 cm, entire, L lamina 13–24  1.5–2.7 cm, lanceolate, cinereous, densely lepidote, with Sp 5–8 mm; peduncle 45–62 cm, green, erect, glabrous; lower peduncular Bra foliaceous, equalling to exceeding the internodes, 6–10 cm, erect, upper peduncular Bra shorter than the internodes, 1.7–2.6 cm, lanceolate, apex acute to attenuate, stramineous, entire, glabrous, erect and clasping the peduncle; fertile part of the Inf 28–55 cm, simple, lax, rachis green, glabrous; floral Bra exceeding the pedicels, 11–25  4–7 mm, lanceolate or ovate-lanceolate, apex acute to attenuate, completely stramineous or with green base, entire, glabrous; Fl suberect; Ped 8–10 mm; Sep 8–10  5–6 mm, with green base and castaneous apex, lanceolate, acute, entire, not imbricate, symmetrical; Pet 11–13  3–4 mm, green, narrowly elliptic, apex rounded, entire, not imbricate, symmetrical; St 14–16 mm, exserted; Fil free, green; Ov 5–8 mm, green; Sty 3–5 mm,

969

green, exserted; Sti green; Fr 10–13 mm, globose, castaneous; Se  3 mm, with falciform wings. Closely related to E. reflexum. E. disjunctum Forzza (Bol. Inst. Bot. (São Paulo) 23(1): 15–16, figs. 7, 16 (pp. 19, 39), 2005). Type: Brazil, Goiás (Forzza & al. 1570 [SPF]). — Distr: Brazil (Goiás); on sandstone. Flowering plants 47–65 cm tall, solitary or forming small clumps; rhizome long, branched; Ros 15–22 cm ∅; L erect-patent, sheath 2.8  3.7 cm, L lamina 12–17  1.2–1.7 cm, both faces green, only the abaxial-proximal region cinereous, margins with Sp 1–2 mm long; peduncle 44–47 cm, erect, glabrous; middle peduncular Bra exceeding the internodes, erect, lanceolate, acute to attenuate, margins spinose, glabrous, upper peduncular Bra shorter than the internodes and entire; fertile part of the Inf  15 cm, simple, few-flowered, lax, rachis glabrous; floral Bra shorter than the pedicels, 12–18 mm, stramineous, lanceolate, glabrous, acute to attenuate, entire; Fl (only faded flowers seen) patent to reflexed; Ped 12–36 mm, glabrous; Sep 17–19  5–7 mm, ovate, glabrous, not imbricate, symmetrical, apex obtuse, entire; Pet 10–12  16–19 mm, colour not described, obovate, glabrous, imbricate, symmetrical, apex rounded, entire; St 15–18 mm, free, included; Ov  8 mm; Sty  9 mm, included; Sti lobes compact, 1 mm; Fr 13–17 mm, dark brown, brilliant, with persistent perianth remains; Se  4 mm, with rounded wings. A member of the longipedicellatum-complex, and similar to E. pedicellatum. E. eddie-estevesii Leme & Forzza (Selbyana 23: 200, fig. 1a–f, 2002). Type: Brazil, Goiás (Esteves Pereira 346 [HB, UFG]). — Distr: Brazil (Bahia, Goiás); limestone outcrops. I: Braun (2005); Forzza (2005: 38). Flowering plants 0.9–2.2 m tall, forming large clumps; rhizome branched; Ros 60–100 cm ∅; L erect-patent, sheath 3.5–6.7  6.5–9.3 cm, L lamina 20–56  2.5–4.2 cm, both faces strongly cinereous, green to slightly reddish, armed with Sp 6–10 mm long; peduncle 45–120 cm, brownishgreen, erect, glabrous; peduncular Bra exceeding the internodes, 4–12.5 cm, erect to slightly patent,

970

base cinereous, stramineous towards the apex, triangular-lanceolate to lanceolate, acute to longattenuate, margins spinose, lepidote to glabrescent; fertile part of the Inf 22–38 cm, simple or rarely with a small branch near the base, multi-flowered, lax, rachis brown to greenish-brown, glabrous; lower floral Bra similar to the peduncular bracts, exceeding the pedicels, 17–25  7 mm, cinereous, lanceolate, acute, margins serrate, lepidote, upper floral Bra very small, shorter than the pedicels, 1–4  2–3 mm, stramineous, ovate-triangular, acute to acuminate, entire to slightly serrate, lepidote to glabrescent; Fl patent; Ped 3–4 mm, brown, glabrous; Sep 6–8  4–5 mm, yellow, abaxially with scattered pinkish blotches, ovatetriangular, acute, margins inconspicuously crenate, imbricate, symmetrical or slightly asymmetrical, glabrous; Pet 15–23  5–7 mm, yellow, abaxially with scattered pinkish blotches, elliptic, apex acute to obtuse, margins inconspicuously crenate, partially imbricate, slightly asymmetrical, glabrous; St 10–18 mm, exserted; Fil connate, not adnate to the petals; Ov 6–10 mm; Sty 5–10 mm, exserted; Sti lobes laminar; Fr 12–16 mm; Se 3–4 mm, with falciform wings. E. erectiflorum L. B. Smith (Phytologia 20: 180, t. 2: figs. 15–16, 1970). Type: Brazil, Piauí (Andrade-Lima 66–4800 [US, IPA]). — Distr: Brazil (Ceará, Piauí); on rocks in Caatinga and transitions to Cerrado vegetations. I: Forzza (2005: 44). Flowering plants 1.5–2 m tall, forming large clumps; rhizome not described; Ros 80–100 cm ∅; L erect-patent, sheath 7  5.5 cm, L lamina 50–75  2.5–3.8 cm, both faces green to slightly cinereous, sparsely lepidote, armed with Sp 6–10 mm long; peduncle 1.5 m, green, erect, glabrous; middle peduncular Bra exceeding the internodes, 20–28 cm, cinereous to stramineous, triangular-lanceolate from a broad-ovate base, apex awned, margins spinose, reflexed, lepidote, upper peduncular Bra shorter than or slightly exceeding the internodes, 4.5–7.5 cm, erect or apex reflexed, stramineous, lanceolate, apex attenuate to awned, entire, lepidote; fertile part of the Inf compound with up to 9 Br 32–72 cm long, without terminal raceme, multi-flowered, subdense, rachis green, glabrous; floral Bra shorter than

F. Krapp

the pedicels, 7–9  1 mm, green, lanceolate, apex acute to attenuate, entire, glabrous; Fl secund; Ped 8–19 mm, green, glabrous; Sep 6–8  4–6 mm, yellow, ovate-triangular, acute, entire, not imbricate, symmetrical, glabrous; Pet 18–20  5–7 mm, yellow, elliptic, apex obtuse, entire, symmetrical, not imbricate, glabrous; St 22–25 mm, exserted; Fil free, not adnate to the petals; Ov 10 mm; Sty 10 mm, exserted; Sti lobes laminar; Fr 16–21 mm; Se 3–4 mm, with falciform wings. E. fragae Forzza (Phytotaxa 227: 14, figs. 1–2 (pp. 15–16), 2015). Type: Brazil, Bahia (Fraga & al. 2735 [RB, HUEFS, US]). — Distr: Brazil (Bahia: São Desidério, Sertaneja); limestone outcrops. Flowering plants 2.2–3 m tall, solitary or forming clumps; Ros 60–100 cm ∅; L erect, sheath 4–4.7  7–7.5 cm, L lamina 56–75  3–3.4 cm, densely lepidote, greyish, armed with Sp 2–5 mm long; peduncle 1 m, greenishbrown, erect, glabrous or sparsely lepidote; peduncular Bra exceeding the internodes, 16–47 cm, cinereous, green at the base, densely lepidote, lanceolate, spinose, erect; fertile part of the Inf  1.1 m, lax, branched with Br to 63 cm, without terminal raceme, rachis green, sparsely lepidote; floral Bra longer than the pedicels, 12–20  1–2 mm, green at the base, becoming stramineous, linear-triangular, entire or inconspicuously spinose, glabrous; Fl polystichous, spreading; Ped 5–12 mm, brown; Sep 18–22  5–7 mm, vinaceous, lanceolate, long-attenuate, acute, entire, not imbricate, symmetrical; Pet 28–32  6–7 mm, vinaceous, narrowly ovate, obtuse, forming a tube with only the tips recurved, entire, imbricate, symmetrical; St 26–27 mm, included or slightly exserted; Fil free; Ov 10–12 mm; Sty 16–18 mm, exserted, vinaceous; Sti vinaceous; Fr 15–25 mm, globose, chestnutbrown; Se  3 mm, with falciform wings. The species shares the inflorescence architecture (heterothetic, i.e. without terminal raceme) with E. erectiflorum, E. ascendens, E. bracteatum and E. horridum, but is easily distinguished by the deep vinaceous corolla. — [F. Krapp & U. Eggli]. E. gracile L. B. Smith (Phytologia 16: 69, t. 1: figs. 2–4, 1968). Type: Brazil, Minas Gerais

Encholirium BROMELIACEAE

(Belem 1620 [US, CEPEC, NY, UB]). — Distr: Brazil (Espírito Santo, Minas Gerais); limestone outcrops. I: Forzza (2005: 23). Flowering plants 80–100 cm tall, solitary or forming clumps; Ros 30–40 cm ∅; L erect-patent, sheath 1.3–2.2  1.8–3.3 cm, L lamina 20–35  1.6–2.2 cm, both faces cinereous, green or reddish-green, armed with Sp 3–5 mm long; peduncle 33–72 cm, light brown, erect, glabrous; middle peduncular Bra shorter than the internodes, 3.5–5.2 cm, erect or reflexed, cinereous to slightly reddish, lanceolate, apex attenuate to awned, entire to serrate, lepidote, upper peduncular Bra shorter than the internodes, light brown, lanceolate, acute, entire, erect, glabrous; fertile part of the Inf 10–35 cm, simple or rarely with 1–3 Br at the base, few-flowered, lax, rachis yellow-green, glabrous; floral Bra shorter than the pedicels, 3–9  1–6 mm, brown, ovate-triangular, acute, entire, glabrous; Fl patent; Ped 5–22 mm, white to slightly yellowish, glabrous; Sep 7–11  5–7 mm, yellow, ovate, apex acute to obtuse, entire, not imbricate, symmetrical, glabrous; Pet 14–18  4–5 mm, yellow, elliptic to ovate, acute, entire, not imbricate, symmetrical, glabrous; St 14–20 mm, exserted; Fil free, not adnate to the petals; Ov 5–7 mm; Sty 3–5 mm, exserted; Sti lobes laminar; Fr 7–8 mm; Se 1–2 mm, with long-caudate wings. Closely related to E. horridum, but much smaller in habit. The two species share a unique seed morphology with long-caudate wings. E. heloisae (L. B. Smith) Forzza & Wanderley (Bol. Bot. Univ. São Paulo 17: 264, 1998). Type: Brazil, Minas Gerais (Smith & al. 6698 [US, R]). — Lit: Christianini & al. (2013: pollination biology). Distr: Brazil (Minas Gerais: Serra do Cipó); sandy and rocky soils in Campo Rupestre vegetation. I: Forzza (2005: 14).  Dyckia heloisae L. B. Smith (1955); incl. Encholirium sazimae Rauh (1987). Flowering plants 25–50 cm tall, solitary or in small clumps; Ros 6–18 cm ∅; inner L erectpatent, outer slightly reflexed, sheath 1.5–2.2  2.8–4.2 cm, L lamina 4.5–9.5  1–1.7 cm, both faces cinereous on young leaves, later pale green to black and only cinereous at the base, margin entire or rarely with Sp 1–2 mm long; peduncle

971

13–30 cm, green or yellowish-green, erect, glabrous; peduncular Bra mostly shorter than the internodes, 1.5–4.5 cm, green with stramineous apex, lanceolate to ovate-lanceolate, acute, attenuate, entire, lepidote to glabrescent; fertile part of the Inf 9–15 cm, simple, few-flowered, lax, rachis yellow to greenish-yellow, glabrous; floral Bra 15–20 mm, stramineous to slightly greenish, lanceolate, acute to attenuate, glabrous; Fl patent; Ped 10–22 mm, greenish to yellowish-green, glabrous; Sep yellowish-green, to 12–15  6–8 mm, green, ovate, obtuse, entire, not imbricate, symmetrical, glabrous; Pet 10–16  8–10 mm, green, obovate, apex rounded, entire to slightly eroded, overlapping, symmetrical, glabrous; St 8–12 mm, included; Fil shortly connate, not adnate to the petals; Ov 5–7 mm; Sty  2 mm, included; Sti lobes compact; Fr 9–12 mm; Se  3 mm, with truncated wings. Belongs to the longipedicellatum-complex, and grows sympatrically with E. vogelii. Flowers are pollinated by hummingbirds, a local endemic bat, and an unidentified sphingid (Christianini & al. 2013). E. horridum L. B. Smith (Contr. Gray Herb. 129: 32, t. 3: figs. 1–3, 1940). Type: Brazil, Espírito Santo (Foster & Foster 193 [GH]). — Lit: Hmeljevski & al. (2015: population biology). Distr: Brazil (Espírito Santo, Minas Gerais); inselbergs in the Atlantic Forest biome. I: Forzza (2005: 41); Miller (2009). Flowering plants 1.8–2.1 m tall, generally solitary and monocarpic; Ros 60–100 cm ∅; L strongly reflexed, sheath 3–4.2  5–6.7 cm, L lamina 60–95  3–3.5 cm, green to yellowishgreen, margins armed with Sp 6–15 mm long; peduncle 40–80 cm, brown, erect, glabrous; middle peduncular Bra exceeding the internodes, 8.7–15 cm, reflexed, green becoming castaneous, triangular-lanceolate to lanceolate from a broadovate base, apex attenuate to long-attenuate, margins spinose, glabrous, upper peduncular Bra shorter than the internodes, 3–3.4 cm, erect, light brown, lanceolate, apex acute to attenuate, entire, glabrous; fertile part of the Inf 80–130 cm, compound with 10–12 pendulous Br 44–82 cm long, without terminal raceme, multi-flowered, lax to

972

subdense, rachis brown to greenish-brown, glabrous; floral Bra shorter than the pedicels, 3–7  2–3 mm, greenish-brown, ovate-triangular, acute, entire, glabrous; Fl patent; Ped 10–15 mm, white to slightly yellowish, glabrous; Sep 6–10  5–6 mm, yellow, ovate, apex acute to obtuse, entire, not imbricate, symmetrical or slightly asymmetrical, glabrous; Pet 15–22  6–8 mm, yellow, ovate, apex obtuse, entire, not imbricate, symmetrical, glabrous; St 15–22 mm, exserted; Fil free, not adnate to the petals; Ov 5–7 mm; Sty 7–8 mm, exserted; Sti lobes laminar; Fr 10–15 mm; Se 2–3 mm, with long-caudate wings. Closely related to E. gracile but much larger in habit. The two species share a unique seed morphology with long-caudate wings. Hmeljevski & al. (2015) found that pollen dispersal distances are low (mean 45.5 m, maximum 156 m) and pollination is mostly within populations. The plants are almost always monocarpic, and clonal growth is a rare event. E. irwinii L. B. Smith (Phytologia 19: 284, t. 1: fig. 12, 1970). Type: Brazil, Minas Gerais (Irwin & al. 23573 [US, NY, UB]). — Distr: Brazil (Minas Gerais: Grão Mogol); on rocks in Campo Rupestre vegetation. I: Forzza (2005: 27). Flowering plants 80–100 cm tall, forming large clumps; Ros 20–30 cm ∅; L secund or rarely erect-patent, sheaths 3.8–4  1.7–2.5 cm, L lamina 22–35  0.7–1.1 cm, both faces dark green, red to nigrescent, adaxially slightly cinereous, margins with Sp 2–4 mm long; peduncle 43–90 cm, brownish-green, erect, glabrous; middle peduncular Bra generally exceeding the internodes, 3.2–9 cm, erect, stramineous, lanceolate from a broad-ovate base, acute, attenuate to awned, margins spinose, glabrous or sparsely lepidote, upper peduncular Bra shorter than the internodes, 1.3–2.5 cm, erect, stramineous, lanceolate, acute to acuminate, entire, glabrous; fertile part of the Inf 29–40 cm, simple, lax, few-flowered, rachis green, glabrous; floral Bra smaller or slightly exceeding the pedicels, 6–11  2–3 mm, stramineous, lanceolate, acute to attenuate, entire, glabrous; Fl patent; Ped 5–9 mm, green, glabrous; Sep 5–8  3–5 mm, green, ovate-triangular, acute to acuminate, entire,

F. Krapp

not imbricate, symmetrical, glabrous; Pet 7–9  3–4 mm, green, elliptic, acute to acuminate, entire, not imbricate, symmetrical, glabrous; St 9–13 mm, exserted; Fil free, not adnate to the petals; Ov  5 mm; Sty 5–6 mm, exserted; Sti lobes compact; Fr 13–18 mm; Se 2–4 mm, with truncated wings. Narrowly endemic to the Grão Mogol area, and the only species of Encholirium there. E. josinoi-narcisae P. J. Braun & Esteves (Bromelie 2018(1): 33, ills. (pp. 32–38), 2018). Type: Brazil, Goiás (Esteves Pereira 788 [UFG]). — Distr: Brazil (N Goiás: Mun. Monte Alegre de Goiás); bambui limestone outcrops, 570 m; only known from the type locality. Flowering plants 1.5–1.9 m tall, forming small clumps to 60 cm ∅, with short rhizome; Ros 23–47 cm ∅; L lax, spreading regularly or somewhat falcately upcurved, sheath broadly ovate, 2.5–3.8  4.2–5.1 cm, stiff, esp. towards the lamina succulent, in part cinereous-lepidote, otherwise glabrous, glossy, bright brown, L lamina 9–28 ( 34)  2.5–5.2 cm broad at the base, (pale) green, cinereous-lepidote on both faces, margins spinose, Sp predominantly latrorse, brilliant blackish to brown, 7 mm, 15 mm distant; peduncle to 110 cm, green-brown, erect, peduncular Bra triangular-lanceolate, caudate, spinose, 4.6–23 cm, green-brown, cinereous-lepidote; fertile part of the Inf  75 cm, simple, laxly flowered, glabrous; floral Bra minute; Fl semi-erect; Ped 42–55 mm, bright green to brown, sulcate, glabrous; Sep 8–11  7–8 mm, basally fused 1–4 mm, yellowishgreen dotted with tiny brown-pink spots, triangular, obtuse to emarginate, glabrous; Pet 31–38  5–7 mm, pale yellow, narrowly lanceolate, free, glabrous, dorsally with tiny brown-pink dots; St to 32 mm, just visible in the throat; Fil free, 25 mm, pale yellow; Anth 6–7 mm, yellow with whitish apex; Ov trigonous, 7–19  3–3.7 mm at the base, bright green; Sty  32 mm, green; Sti slightly exserted, 4 mm, spirally conduplicate; Fr 28–31  9–14 mm, oblong, bright dark brown; Se 3  1 mm, dark brown, surrounded with a falcate white-cream wing. Compared with E. eddie-estevesii and E. longiflorum in the protologue. — [U. Eggli].

Encholirium BROMELIACEAE

E. kranzianum Leme & Forzza (Phytotaxa 227: 14, 17, figs. 3–4 (pp. 18–19), 2015). Type: Brazil, Minas Gerais (Kranz 393 [RB]). — Distr: Brazil (Minas Gerais); quartzitic rocks and sandy soil. Flowering plants 95 cm tall, solitary or forming clumps; Ros 10–14 cm ∅; L spreading-recurved, sheath 1.5–2  2.5–2.8 cm, L lamina 6.5–8.5  0.8–1.2 cm, densely white-lepidote, sparsely towards the apex, bronze-coloured, margins densely to subdensely spinose with triangular Sp 1.5–2.5 mm long; peduncle 65–68 cm long, dark greenish to dark purple, erect, glabrous, covered by a thin layer of white wax; peduncular Bra shorter than the internodes except the lowest, stramineous, soon glabrous, narrowly subtriangularlanceolate, acute-attenuate, inconspicuously spinose, recurved; fertile part of the Inf  24 cm, lax, simple, rachis dark purple, glabrous; floral Bra shorter than the pedicels, 5–8  1–3 mm, stramineous, narrowly subtriangular-lanceolate, caudate, entire or inconspicuously spinose, glabrous; Fl polystichous, campanulate, spreading; Ped 25–35 mm, dark purple; Sep 7–8  5–6 mm, dark purple, broadly oblong-ovate, subacute to obtuse, denticulate near the apex, not imbricate, symmetrical; Pet  11  11 mm, green except the dark purple centre of the apex, broadly obovate to orbicular, apex broadly rounded, irregularly crenulate, imbricate, symmetrical; St  12 mm, exserted; Fil free; Ov  5 mm; Sty  6 mm, slightly exserted, green; Sti green; Fr and Se unknown. Closely related to E. pulchrum and E. scrutor from which it can be easily distinguished by the unique campanulate flowers and some other characters. E. longiflorum Leme (Selbyana 16: 110, fig. 1, 1995). Type: Brazil, Minas Gerais (Teixeira & Carvalho 380 [MBM, HB]). — Distr: Brazil (Bahia, Minas Gerais); limestone outcrops in Cerrado and transitions to Caatinga vegetations. I: Forzza (2005: 4, 43). Flowering plants 3–3.5 m tall; Ros > 100 cm ∅; L sheath 6.8–7.5  9.2–14 cm, L lamina 53–83  5–8.3 cm, both faces strongly cinereous, margins with Sp 3–5 mm long; peduncle

973

(fragment known only) glabrous; middle peduncular Bra exceeding the internodes, 12–30 cm, cinereous, lanceolate, apex acute to awned, margins spinose, lepidote, upper peduncular Bra exceeding the internodes, 2.8 cm, stramineous, lanceolate, apex acute to awned, entire, sparsely lepidote; fertile part of the Inf 49–62 cm, simple, multi-flowered, lax (dense according to Braun & Esteves Pereira 2018), rachis glabrous; floral Bra reduced, much shorter than the pedicels, 2–5  2–3 mm, ovate-triangular, sometimes linearlanceolate in basal flowers, apex usually acute to acuminate, rarely attenuate, margins entire or minimally serrate, glabrous; Fl patent; Ped 27–35 ( 52) mm, glabrous; Sep 8–19  3–7 mm, green, ovate-triangular, apex acute to obtuse, entire, not imbricate, symmetrical, glabrous; Pet 22–38  5–12 mm, greenish-white, elliptic, apex acute to obtuse, entire, not imbricate, symmetrical, glabrous; St 23–34 mm, exserted; Fil connate, adnate to the petals; Ov 9–12 mm; Sty 5–12 mm (25 mm according to Braun & Esteves Pereira l.c.), exserted; Sti lobes laminar; Fr 22–25 mm; Se 3–4 mm, with falciform wings. Identified by long pedicels and petals. The description was completed from the comparison in the protologue of the similar E. josinoi-narcisae (Braun & Esteves Pereira 2018). — [F. Krapp & U. Eggli]. E. luxor L. B. Smith & Read (Bradea 5(27): 229, fig. 1, 1989). Type: Brazil, Minas Gerais (Anderson & al. 9259 [US, HB, IAN, MO, NY, UB]). — Distr: Brazil (Distrito Federal, Goiás, Minas Gerais); limestone outcrops, usually in open Cerrado vegetation, rarely in dense forests. I: Forzza (2005: 29). Incl. Encholirium piresianum L. B. Smith & Read (1989). Flowering plants 1.6–2.2 m tall, generally forming large clumps; rhizome branched; Ros 80–110 cm ∅; L erect-patent, sheath 3.5–5.7  6–11 cm, L lamina 25–73  2.4–7.2 cm, both faces dark green to cinereous, margins with Sp 3–6 mm long; peduncle 70–120 cm, green, erect, glabrous; middle peduncular Bra exceeding the internodes, 16–34 cm, erect or apex reflexed, base vinaceous, apex cinereous or stramineous,

974

lanceolate to triangular-lanceolate from a broadovate base, apex attenuate and awned, spinose, lepidote, upper peduncular Bra equalling or exceeding the internodes, 2.6–9.2 cm, erect, base vinaceous, stramineous at the apex, lanceolate, acute or acuminate, entire with slighly serrate apex, sparsely lepidote; fertile part of the Inf 45–85 cm, simple, multi-flowered, lax to subdense, rachis green, glabrous; floral Bra shorter than the pedicels, 5–16  2–4 mm, stramineous, slightly greenish or vinaceous, lanceolate to ovate, acute to acuminate, entire, glabrous; Fl patent to erect-patent; Ped 11–25 mm, green or vinaceous, glabrous; Sep 7–12  4–6 mm, vinaceous or green, ovate, apex rounded or slightly acute to apiculate, entire, not imbricate, symmetrical, glabrous; Pet 19–29  5–9 mm, green to slightly vinaceous, elliptic, apex acute to obtuse, entire, valvate, not imbricate, symmetrical, glabrous; St 16–27 mm, exserted; Fil connate, not adnate to the petals; Ov 8–12 mm; Sty 9–13 mm, exserted; Sti lobes laminar; Fr 12–25 mm; Se 3–4 mm, with rounded wings. E. lymanianum E. Pereira & Martinelli (Bradea 3(32): 252, t. 2: photo 2, 1982). Type: Brazil, Mato Grosso do Sul (Martinelli 400 [RB, GUA, HB, K, US]). — Distr: Brazil (Mato Grosso, Mato Grosso do Sul); sandstone in flat parts of the Cerrado. I: Forzza (2005: 4, 35). Flowering plants 1.2–2 m tall, forming large clumps; rhizome branched; Ros 80–100 cm ∅; L erect-patent, sheath 3.5  5.5 cm, entire or slightly spinose distally, L lamina 52–65  2.2–4.5 cm, both faces green to cinereous, margins with Sp 10–12 mm long; peduncle 60–100 cm, erect, lepidote to glabrescent; middle peduncular Bra exceeding the internodes, 5.5–9.5 cm, erect, cinereous to slightly vinaceous, triangular-lanceolate to lanceolate, acute, margins spinose, lepidote, upper peduncular Bra shorter than the internodes, 2.2–3.7 cm, erect, stramineous, lanceolate, acute, entire or serrate, lepidote to glabrescent; fertile part of the Inf 24–57 cm, simple, multi-flowered, lax to subdense, rachis castaneous, densely lanate, with ferrugineous trichomes; floral Bra 5–15  4–7 mm, castaneous, triangular-lanceolate to ovatetriangular, acute to acuminate, entire, densely lanate;

F. Krapp

Fl patent, subsessile; Ped none at anthesis, short after the start of fruit development; Sep 9–12  6–8 mm, ovate, apex acute to obtuse, entire, imbricate, symmetrical, lanate; Pet 17–25  11–14 mm, ovate, apex acute to rounded, margins entire to ciliate, partially imbricate, symmetrical, lanate in the centre, glabrous towards the margins; St 23–35 mm, exserted; Fil free, not adnate to the petals; Ov 10–14 mm; Sty 7–18 mm, exserted; Sti lobes laminar; Fr  25 mm; Se 5–6 mm, with falciform wings. Disjunct from all other Encholirium species. E. magalhaesii L. B. Smith (Phytologia 13: 152, t. 7: fig. 17, 1966). Type: Brazil, Minas Gerais (Magalhães 18056 [IAN]). — Distr: Brazil (Minas Gerais: Chapada Diamantina); directly on rocks in Campo Rupestre vegetation, rarely on sandy and rocky soils. I: Forzza (2005: 27). – Fig. 1.

Fig. 1 Encholirium magalhaesii (Supthut 8843: Brazil; Minas Gerais, near Consoleo Mata). (Copyright: U. Eggli)

Encholirium BROMELIACEAE

Incl. Encholirium suzannae Rauh (1988); incl. Encholirium crassiscapum E. Gross (1997). Flowering plants 45–100 cm tall, solitary or in clumps; rhizome without conspicuous branches; Ros 8–10 cm ∅; L erect-patent, sheath 1.2–2.8  2.7–3.9 cm, L lamina 8–23  0.5–0.9 cm, both faces slightly cinereous, green or sometimes slightly reddish, margins with Sp 2–5 mm long; peduncle 28–70 cm, brownish-green, erect, glabrous; peduncular Bra shorter than the internodes, 2–3.8 cm, erect or reflexed, green at the base, stramineous towards the tip, lanceolate to ovate-lanceolate, acute to acuminate, margins serrate or entire, glabrous or apex lepidote; fertile part of the Inf 10–40 cm, simple, lax, few-flowered, rachis green, glabrous; floral Bra exceeding the pedicels, 11–21  3–6 mm, green at the base, stramineous towards the tip, ovate-lanceolate, acute to acuminate, entire or slightly serrate, glabrous; Fl patent to erect; Ped 5–8 mm, green, glabrous; Sep 6–8  3–4 mm, green or apex sometimes slightly brown, ovate, apex obtuse, entire, not imbricate, symmetrical, glabrous; Pet 7–9  2–3 mm, green, elliptic, apex obtuse, not imbricate, symmetrical, glabrous; St 14–18 mm, exserted; Fil free, not adnate to the petals; Ov 5–6 mm; Sty 4–5 mm, exserted; Sti lobes compact; Fr 7–11 mm; Se  3 mm, with rounded wings. Similar to E. subsecundum, from which it differs by the smaller habit and smaller flower parts. E. maximum Forzza & Leme (Selbyana 23: 202, fig. 1 g–n, 2002). Type: Brazil, Bahia (Forzza & al. 1234 [SPF, CEPEC, HB, K, MBM, NY, SP, US]). — Distr: Brazil (Bahia); directly on rocks; only known from the type locality. I: Forzza (2005: 38); Braun & Esteves Pereira (2006). Flowering plants 1.5–3.8 m tall, generally forming large clumps; rhizome without conspicuous branches; Ros 70–120 cm ∅; L erect-patent, sheath 3.2–4  4.5–7.2 cm, L lamina 30–65  1.8–2.5 cm, both faces cinereous, rarely reddish, margins with Sp 5–9 mm long; peduncle 60–130 cm, green, erect, glabrous; middle peduncular Bra far exceeding the internodes, 12.5–23 cm, reflexed, cinereous, triangular-lanceolate to lanceolate, attenuate to long-attenuate, base broad-ovate, margins spinose, densely lepidote, upper

975

peduncular Bra shorter than the internodes, 2.2–4.5 cm, erect to slightly reflexed, castaneous, lanceolate, apex acute, entire to slightly serrate, glabrous; fertile part of the Inf 60–150 cm, simple, multi-flowered, lax to subdense, rachis bright green, glabrous; floral Bra shorter than the flowers, 13–22  13–17 mm, brown, broadly ovate to obovate, apex acuminate, margins inconspicuously serrate, glabrous; Fl patent, subsessile; Sep 9–13  9–13 mm, yellow, greenish-yellow or apex slightly brown, obovate, apex rounded to retuse, entire or minutely crenate, imbricate, symmetrical or inconspicuously asymmetrical, glabrous; Pet 21–25  11–15 mm, vivid yellow, ovate, apex rounded, entire, partially imbricate, symmetrical, glabrous; St 30–32 mm, exserted; Fil free, not adnate to the petals; hypanthium reduced; Ov 20–22 mm; Sty 7–11 mm, exserted; Sti lobes laminar; Fr 27–33 mm; Se 3–4 mm, with rounded wings. The largest species of the genus. E. pedicellatum (Mez) Rauh (Trop. subtrop. Pfl.-welt 60: 15, ills. (pp. 14–16), 1987). Type: Brazil, Minas Gerais (Schwacke 8413 [B, RB]). — Lit: Cavallari & al. (2006: genetics). Distr: Brazil (Minas Gerais: Diamantina plateau); Campo Rupestre vegetation, sandy and rocky soils; known from a single population only. I: Forzza (2005: 4, 17). – Figs. 2 and 3.  Dyckia pedicellata Mez (1896). Flowering plants 65–70 cm tall, solitary or in clumps; rhizome without conspicuous branches; Ros 9–16 cm ∅; L patent to reflexed, sheaths 1.5–1.8  2.5–3.5 cm, L lamina 6.5–10  1.8–2.5 cm, both faces strongly cinereous, margins with Sp 3–5 mm long; peduncle 27–40 cm, green, erect, glabrous; peduncular Bra shorter than the internodes, 3.5 cm, erect, base stramineous, apex cinereous, lanceolate, acute to acuminate, entire to slightly serrate, glabrous but the upper ones with lepidote apex; fertile part of the Inf 22–25 cm, simple, few-flowered, lax, rachis green, glabrous; floral Bra shorter than the pedicels, 9–18  2–3 mm, stramineous, lanceolate, acute to attenuate, entire, glabrous; Fl patent to slightly reflexed; Ped 22–55 mm, green, glabrous; Sep 11–14  5–8 mm, green,

976

F. Krapp

Fig. 2 Encholirium pedicellatum (Rauh 56195: Brazil; Minas Gerais, near Diamantina). (Copyright: U. Eggli)

not adnate to the petals; Ov  11 mm, Sty 4–6 mm, included; Sti lobes compact; Fr  14 mm; Se  3 mm, with rounded wings. Belonging to the longipedicellatum-complex.

Fig. 3 Encholirium pedicellatum (same as previous illustration). (Copyright: U. Eggli)

ovate, apex obtuse, entire, not imbricate, symmetrical, glabrous; Pet 14–17  12 mm, green, obovate, apex rounded or obtuse, entire, imbricate, symmetrical, glabrous; St  12 mm, included; Fil connate,

E. pierre-braunii Esteves (Bromelie 2017(2): 83–87, ills., 2017). Type: Brazil, Bahia (EstevesPereira 489 [UFG 50.535]). — Distr: Brazil (W Bahia: Serra do Ramalho); exposed karstic bambuí limestone, 740 m, only known from the type locality. Flowering plants to 2.9 m tall, solitary or offseting and group-forming; Ros to 90 cm ∅, dense; L very succulent and stiff, sheath 5.5  8.5 cm, broadly concave, dark brown, glossy, glabrous, L lamina long attenuate, sometimes sinuously curved, to 50  3.8 cm, upper face dark brownish-ruby to brownish-green to almost black, with large silver-grey to whitish areas around the spine bases, lower face dull greyish, finely veined, Sp laxly disposed, variously oriented but often retrorse, rigid, 9 mm, basally 3–17 mm wide, 2.5 cm distant, densely silvery-white-lepidote; peduncle 1.1–2 m, glabrous, erect, pale green; peduncular Bra similar to rosette leaves bud rapidly becoming smaller, to 27 cm, longer than the internodes; fertile part of the Inf to 85 cm, simple, laxly many-flowered; lower floral Bra triangular, long-attenuate, 11 mm, uppermost only 2 mm or rudimentary; Fl horizontally patent; Ped to 27 mm, slender, pale green; Sep 10  6 mm, erect, pale green with minute reddish-brown

Encholirium BROMELIACEAE

spots, ovate-triangular, glabrous; Pet 35  7 mm, somewhat flaring, pale green with minute reddishbrown spots giving the flowers an overall brownish-dull appearance, triangular-lanceolate, slightly succulent, glabrous; St exserted; Fil  30 mm, cream-coloured; Anth  8 mm, dark green; Ov conical, 23  4.6 mm ∅, dark green, glossy; Sty  8 mm, bright green basally, creamy with reddish-brown streaks above; Sti somewhat irregularly laminar, 7 mm, cream-yellowish, with reddish-brown spots, glabrous; Fr size not described but probably 25 mm, pale cream to chestnut-brown, paler with age; Se incl. the wing 4.3  4 mm, seed proper medium brown, papery wing pale creamy-ochre. — [U. Eggli]. E. pulchrum Forzza & al. (Nordic J. Bot. 30: 153, figs. 1, 3A–C, 2012). Type: Brazil, Minas Gerais (Ribeiro & Paula 48 [RB]). — Distr: Brazil (Minas Gerais: Diamantina plateau); flat granitic rock outcrops; known from a single site only. Flowering plants 70–78 cm tall, solitary or in small clumps; rhizome short; Ros  40 cm ∅; L spreading to slightly reflexed, sheath not described, L lamina 20–25  2.5 cm, narrowly triangular to lanceolate, green to reddish, glabrous except the inconspicuously white-lepidote adaxial margins, abaxially lustrous, margins with Sp 3–8 mm long; peduncle 30–35 cm, green, erect, glabrous; lower peduncular Bra foliaceous, equalling or exceeding the internodes, 8–10 cm, erect, upper peduncle Bra shorter than the internodes, 3–4.5 cm, erect and amplexicaul, completely stramineous or with green base, lanceolate, apex acute to attenuate, entire, glabrous; fertile part of the Inf 37–45 cm, simple, lax, erect, rachis green, glabrous; floral Bra shorter than the pedicels, 8–22  1–6 mm, completely stramineous, lanceolate to ovate or filiform, apex acute to attenuate, entire to slightly serrulate, glabrous; Fl suberect to spreading; Ped 25–57 mm, green; Sep 6–10  4–6 mm, green, ovate to broadly oblong-elliptic, apex rounded or crenate, entire or remotely and irregularly denticulate near the apex, imbricate, symmetrical; Pet 9.5–12  8–10 mm, green, oblong to orbicular, apex rounded to obtuseemarginate, entire, imbricate, symmetrical; St 10–11 mm, included; Fil free, green; Ov 7–8 mm,

977

green; Sty 2–4 mm, green, included; Sti green; Fr and Se not described. Belonging to the longipedicellatum-complex, and similar to E. pedicellatum. E. reflexum Forzza & Wanderley (Novon 11 (1): 40–42, Fig. 1, 2001). Type: Brazil, Minas Gerais (Forzza & al. 800 [SPF, BHCB, US]). — Distr: Brazil (Minas Gerais); Campo Rupestre vegetation, directly on rocks. I: Forzza (2005: 22). Flowering plants 60–100 cm tall, solitary or in clumps; rhizome without conspicuous branches; Ros 15–30 cm ∅; L reflexed, sheath 4.2–6  1.8–2.5 cm, L lamina 13–24  1.5–2.7 cm, both faces cinereous, rarely reddish, margins with Sp 4–7 mm long; peduncle 25–62 cm, green, erect, glabrous; middle peduncular Bra shorter than or equalling the internodes, 6–10 cm, glabrous and becoming stramineous at the base, apex cinereous and lepidote, lanceolate from a long-ovate base, attenuate, margins spinose, apex erect or slightly reflexed, upper peduncular Br shorter than the internodes, 1.7–2.6 cm, lanceolate, stramineous, attenuate and acute, entire, erect, glabrous; fertile part of the Inf 22–33 cm, simple, few-flowered, lax, rachis green, glabrous; floral Bra shorter than the flowers, 11–15  4–5 mm, stramineous, lanceolate, acute to attenuate, entire, glabrous; Fl patent; Ped 3–4 mm, green, glabrous; Sep 5–7  5–7 mm, green, obovate, apex rounded, sometimes inconspicuously apiculate, entire, imbricate, symmetrical, glabrous; Pet 7–9  7–9 mm, green, obovate, apex rounded, entire, symmetrical, imbricate, glabrous; St  10 mm, included or slightly exserted; Fil connate, not adnate to the petals; Ov 5–6 mm; Sty  4 mm, included; Sti lobes compact; Fr 11–18 mm; Se  3 mm, with rounded wings. Belonging to the longipedicellatum-complex. E. scrutor (L. B. Smith) Rauh (Trop. subtrop. Pfl.-welt 60: 94, 1987). Type: Brazil, Minas Gerais (Cordeiro & Simonis 4048 [SPF, U]). — Distr: Brazil (Minas Gerais: around Diamantina); Campo Rupestre vegetation, sandy and rocky soils. I: Forzza (2005: 4, 14).

978

 Dyckia scrutor L. B. Smith (1987); incl. Encholirium carmineoviridiflorum Rauh (1987); incl. Encholirium inerme Rauh (1987). Flowering plants 19–56 cm tall, solitary or in clumps; rhizome without conspicuous branches; Ros (4-) 8–15 cm ∅; L secund to slightly secund, rarely erect, sheath 1.2–1.7  1.9–2.5 cm, L lamina 2–11  0.6–1.3 cm, both faces green to cinereous, sometimes black-vinaceous when flowering, margins entire or rarely spinose with Sp  1 mm long; peduncle 12–28 cm, rose to slightly vinaceous, delicate, glabrous or sparsely lepidote; peduncular Bra shorter than the internodes, 0.7–1.6 cm, erect, stramineous or slightly pinkish, lanceolate to ovate-lanceolate, long-attenuate, acute, entire, basal ones with densely lepidote apex, upper ones glabrous; fertile part of the Inf 3–16 cm, simple, few-flowered, lax, rachis pink, glabrous; floral Bra shorter than the pedicels, 4–11  2–4 mm, pinkish, ovate-triangular, acute to acuminate, entire, glabrous; Fl patent to slightly reflexed; Ped 15–51 mm, pink, glabrous; Sep 6–15  4–9 mm, carmine to pinkish near the base, dark green above, ovate, obtuse, entire, not imbricate, symmetrical, glabrous; Pet 8–13  6–9 mm, cream-green at the base, (dark) green towards the apex, obovate, apex rounded, entire, imbricate, symmetrical, glabrous; St 6–10 mm, included; Fil connate, adnate to sepals and petals; Ov 6–7 mm; Sty  1 mm, included; Sti lobes compact; Fr 10–13 mm; Se 4–5 mm, with roundish wings. Belonging to the longipedicellatum-complex. E. spectabile Martius ex Schultes & Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): lxviii, 1233, 1830). Type: Brazil, Bahia (Martius 2483 [M]). — Lit: Siqueiro Filho & Leme (2007: 318–320, with ills., 320-323, as E. pernambucanum); Queiroz & al. (2016); Gonçalves-Oliveira & al. (2017: population genetics); Distr: Brazil (Bahia, Ceará, Distrito Federal, Paraiba, Piauí, Pernambuco, Rio Grande do Norte, Sergipe); rocky habitats (inselbergs) in Caatinga vegetation. I: Forzza (2005: 32).  Dyckia spectabilis (Martius ex Schultes & Schultes fil.) Baker (1889); incl. Puya saxatilis Martius (1828) (nom. inval., ICN Art. 38.1a); incl. Encholirium densiflorum Ule (1908); incl.

F. Krapp

Encholirium rupestre Ule (1908); incl. Encholirium hoehneanum L. B. Smith (1943); incl. Encholirium lutzii L. B. Smith (1958); incl. Encholirium patens L. B. Smith (1972); incl. Encholirium bahianum L. B. Smith & Read (1989); incl. Encholirium harleyi L. B. Smith & Read (1989); incl. Encholirium paraibae L. B. Smith & Read (1989); incl. Encholirium pernambucanum L. B. Smith & Read (1989). Flowering plants 1.2–2.5 m tall, forming large clumps; rhizome branched; Ros 50–110 cm ∅; L erect-patent, sheath 2.8–4.7  3.8–6.7 cm, L lamina 25–84  2.2–4.7 cm, both faces very variable in colour, light green, green-red, greenyellow, yellow or yellow-red, margins with Sp 6–12 mm long; peduncle 80–150 cm, green to castaneous, erect, glabrous; middle peduncular Bra exceeding the internodes, 12–25 cm, erect or apically reflexed, base vinaceous, towards the apex cinereous or stramineous, triangular-lanceolate to lanceolate from a broad-ovate base, apex awned, spinose, lepidote, upper peduncular Bra smaller, equalling or exceeding the internodes, 2.4–9.5 cm, vinaceous base with stramineous apex to completely stramineous, lanceolate, acute to acuminate, entire to slightly serrate, erect, usually glabrous or sparsely lepidote; fertile part of the Inf 35–88 cm, simple or rarely with partially undeveloped branches at the base, multi-flowered, lax, subdense to dense, rachis green or greenishbrown, glabrous; lower floral Bra equalling or exceeding the pedicels but not the flowers, 5–22  1–6 mm, castaneous to nigrescent, in young inflorescences slightly greenish with vinaceous apex, filiform, lanceolate, triangular, ovatelanceolate, acute, acuminate to attenuate, entire, glabrous; Fl erect-patent; Ped 3–15 mm, green to brown, glabrous; Sep 7–12  3–8 mm, green, yellowish-green or green with vinaceous apex, ovate, apex acute to obtuse or rarely slightly retuse, margins entire to slightly crenulate, not imbricate, symmetrical, glabrous; Pet 9–22  4–10 mm, green or cream-green, apex vinaceous in buds, elliptic, apex obtuse, entire, not imbricate, symmetrical, glabrous; St 15–32 mm, exserted; Fil free; Ov 8–13 mm; Sty 12–15 mm, exserted; Sti lobes laminar; Fr 12–27 mm; Se 2–4 mm, with falciform wings.

Encholirium BROMELIACEAE

By far the most widespread and morphologically most variable species of the genus. Many formerly distinct species were synonymized by Forzza (2005). According to Braun & EstevesPereira (2017), the species also occurs in Alagoas and Maranhão. Nascimento & al. (2012) report that the leaf bases (grated, sun-dried, sieved) were used to produce a bitter-tasting flour for couscous preparation during times of famine in Pernambuco. Queiroz & al. (2016) found a mixed pollination system: The flowers open in mid-afternoon and last for 20 hours. They are effectively visited by 3 species of bats and the opossum Didelphis albiventris during the night, and 3 species of hummingbirds during the day; hawkmoths and passerine birds are recorded as sporadic further visitors. Hummingbirds were also recorded as flower visitors by Las-Casas & al. (2012). The population genetic study of Gonçalves-Oliveira & al. (2017) indicates that gene flow via bat pollination is higher than that through seed dispersal. Each inselberg population has a unique genetic composition, but two population groups (one from the SE Caatinga, the other from the remaining range in the SW Caatinga, the N Espinhaço Range, and the Borborema Plateau) have been found, indicating the possible presence of a cryptic species. — [F. Krapp & U. Eggli]. E. splendidum Forzza (Phytotaxa 227: 17, 23, figs. 5–7 (pp. 20–22), 2015). Type: Brazil, Bahia (Forzza & al. 4078 [RB, B, HUEFS, K, NY, SPF, US]). — Distr: Brazil (Bahia); limestone outcrops. Flowering plants 1.4–1.6 m tall, propagating by long stolons; Ros 20–40 cm ∅; L spreading to slightly reflexed, sheath 3.5–4.5  4.2–5.7 cm, L lamina 28–33  1.8–2.2 cm, glabrous, inconspicuously lepidote, green to reddish, armed with broadly triangular Sp 3–5 mm long; peduncle 70–95 cm, green, erect, glabrous; lower peduncular Bra exceeding the internodes, 18–32 cm, upper peduncular Bra exceeding or shorter than the internodes, 5.5–9.5 cm, green at the base or completely stramineous, glabrous or inconspicuously white-lepidote, lanceolate, attenuate, spinose, erect to spreading; fertile part of the Inf 48–62 cm, lax, simple, rachis green, glabrous; floral Bra

979

much shorter than the pedicels, 2–3  1–2 mm, stramineous or green, lanceolate to ovate, acute, entire, glabrous; Fl polystichous, spreading; Ped 2.2–4.2 mm, green; Sep 7–11  4–6 mm, green, sometimes with purplish-vinaceous spots, ovatelanceolate, apex narrowly rounded, entire, not imbricate, symmetrical; Pet 30–35  5–7 mm, green, sometimes with purplish-vinaceous spots, oblong-lanceolate, apex narrowly rounded, curled, entire, not imbricate, symmetrical; St 28–32 mm, exceeding the petals; Fil free; Ov 10–12 mm; Sty 7–9 mm, exserted, pale green; Sti pale green; Fr 15–25 mm, globose, chestnut-brown; Se 3–4 mm, with falciform wings. Closely related to E. longiflorum but much smaller. Bat pollination was observed for this species. E. subsecundum (Baker) Mez (in A. & C. de Candolle, Monogr. Phan. 9: 540, 1896). Type: Brazil, Minas Gerais (St. Hilaire E-496 [P]). — Lit: Sazima & al. (1989: pollination ecology, as E. glaziovii); Cavallari & al. (2006: genetics). Distr: Brazil (Minas Gerais); on rocks.  Dyckia subsecunda Baker (1889); incl. Encholirium glaziovii Mez (1894). Flowering plants 1.5–2 m tall, usually in large clumps; rhizome branched; Ros 60–100 cm ∅; L erect-patent, sometimes slightly arching, sheath 4–6.3  4.2–5.8 cm, L lamina 46–73  1.8–4 cm, both faces green or cinereous, margins with Sp 5–15 mm long; peduncle 70–150 cm, green or brown, erect or recurved, glabrous; peduncular Bra generally exceeding the internodes, middle ones 12–35 cm, erect to reflexed, stramineous, lanceolate from a broad-oval base, apex awned, margins spinose, glabrous, upper peduncular Bra 4.3–5.5 cm, erect or apex reflexed, vinaceous or brown when young, later stramineous, lanceolate, acute-attenuate, serrate, glabrous; fertile part of the Inf 15–40 cm, simple, multi-flowered, dense or rarely lax, rachis green to slightly grey, glabrous; floral Bra exceeding the flowers, 20–35  8–16 mm, stramineous or base slightly greenish, ovate-lanceolate, acute, attenuate, entire or serrate, glabrous; Fl erect; Ped 5–10 mm, green, glabrous; Sep 12–18  5–7 mm, green, triangular-lanceolate, acute, entire, not imbricate,

980

symmetrical, glabrous; Pet 15–22  2–4 mm, green, elliptic, apex obtuse, not imbricate, symmetrical, glabrous; St 15–25 mm, exserted; Fil free; Ov 5–7 mm; Sty 7–10 mm, exserted; Sti lobes laminar; Fr 15–25 mm; Se  5 mm, falciform. Usually growing as solitary clumps, not in large populations as most other species. E. viride P. J. Braun & Esteves (Bromelie 2017 (1): 5, ills. (pp. 6–10), 2017). Type: Brazil, Minas Gerais (Esteves-Pereira 472 [UFG]). — Distr: Brazil (Minas Gerais: Matias Cardoso); on limestone in transition of Campo Cerrado/Caatinga vegetation, 500 m. Flowering plants 1.8–2.7 m tall, offsetting with thick short rhizomes and forming groups; Ros 35 cm tall, 75–95 cm ∅; L mostly archingspreading, firm, succulent, sheath 8  9 cm, almost globosely curved and amplexicaul, dark brown, both faces glabrous and glossy, margins with a few minute teeth apically, L lamina 35–80  6.5–11 cm, long triangular-attenuate, often contorted, palish green, with pungent brown tip, both faces veined, densely grey-lepidote, margins with antrorse Sp 6–11 mm long, with broad green deltoid base, 21 mm apart; Inf erect, simple; peduncle to 1.05 m, first remaining short for a long time and rapidly elongating just before anthesis, glabrous; peduncular Bra erect to contorted, the largest to 76 cm, coriaceous, margins slightly spinose; fertile Inf part 0.85–1.34 cm, rather densely many-flowered, green, glabrous; floral Bra minute, triangular, succulent, green; Fl spreading; Ped 13 mm, pale green; Sep 8  5 mm, triangular, obtuse, succulent, green, glabrous; Pet 17  7 mm, linear to oblongovate, obtuse, bright green, with or without pinkish-violet speckles, glabrous; St  17 mm, exserted; Fil free, white; Anth 5 mm, sagittate, green; Ov 5  5 mm, 3-sulcate, glossy (dark) green; Sty  15 mm, bright green; Sti 3 mm, green-yellow, somewhat spirally contorted; Fr brown changing to straw-coloured with age, size not described; Se irregularly shaped, 4.3  4 mm, cream-coloured with bright brown testa, papery. Compared with the widespread and variable E. spectabile, but with smaller rosettes, distinctly

F. Krapp

green and broader leaves and conspicuously green flowers. — [U. Eggli]. E. viridicentrum Leme & O. B. C. Ribeiro (Phytotaxa 177(2): 86–89, ills., 2014). Type: Brazil, Minas Gerais (Ribeiro & Leme 282 [RB, HB]). — Distr: Brazil (Minas Gerais); quartzitic outcrops in Campo Rupestre vegetation. Flowering plants 25–30 cm tall; Ros 12–15 cm ∅; L spreading, recurved, sheath 1.4  1–1.8 cm, L lamina 5.5–5.7  0.5–0.8 cm, white-lepidote except the glabrous adaxial apex, dark purple with green or yellowish-green base, armed with narrowly triangular Sp 1.5–2 mm long; peduncle 14–16 cm, green, erect, glabrous; peduncular Bra longer than the internodes, stramineous, lower ones leaf-like, upper ones broadly ovate, caudate, serrulate, glabrous; fertile part of the Inf 6.5–9 cm, dense, simple, rachis green, glabrous; floral Bra  equalling the flowers, 12–15  4–6 mm, green, soon stramineous, ovate-lanceolate, acuminate, entire, glabrous; Fl suberect; Ped 2 mm, green, glabrous; Sep 7–8  3–4 mm, green, narrowly ovate, acute, apiculate, entire, not imbricate, symmetrical, glabrous; Pet 9  2–2.5 mm, green, narrowly ovate-lanceolate, apex narrow and emarginate, entire, not imbricate, symmetrical, glabrous; St 12–13 mm, exserted; Fil free or nearly so; Ov 3–4 mm; Sty 4–5 mm, exserted; Sti green; Fr and Se unknown. Closely related to E. ctenophyllum, but much smaller. E. vogelii Rauh (Trop. subtrop. Pfl.-welt 60: 95–98, ills., 1987). Type: Brazil, Minas Gerais (Vogel & Sazima 209 [HEID, UEC]). — Lit: Christianini & al. (2013: pollination ecology). Distr: Brazil (Minas Gerais: Serra do Cipó); sandy or rocky soils, between rocks. I: Forzza (2005: 17). Flowering plants 1.4–2.5 m tall, forming large clumps; rhizome without conspicuous branches; Ros 80–100 cm ∅; inner L erect, outer L patent, sometimes secund, sheath 4.8–7.4  7.2–9.3 cm, L lamina 34–65  3–4 cm, both faces green or slightly reddish, margins with Sp 5–10 mm long; peduncle 70–80 cm, green, erect, glabrous; peduncular Bra exceeding the internodes, 12–35 cm, erect or with reflexed apex, green but

Encholirium BROMELIACEAE

withering stramineous, triangular-lanceolate from a broad-ovate base, attenuate, spinose, glabrescent; fertile part of the Inf 30–50 cm, simple, manyflowered, lax, apex sometimes somewhat arching, rachis green, glabrous; floral Bra exceeding the pedicels but not the flowers, 15–70  20–40 mm, castaneous or slightly greenish, lanceolate, acute to long-attenuate, entire, lanuginose with white trichomes, simple; Fl patent; Ped 23–34 mm, green, glabrous; Sep 8–15  5–7 mm, green, ovate, apex obtuse, margins densely whitelanuginose, slightly eroded, not imbricate, symmetrical, glabrous or sparsely lanuginose; Pet 9–15  6–9 mm, green, obovate, apex rounded, with slightly eroded margins, sparsely whitelanuginose, imbricate, symmetrical, glabrous; St 9–13 mm, included; Fil connate, not adnate to the petals; Ov  10 mm; Sty  1 mm, included; Sti lobes compact; Fr 12–15 mm; Se  5 mm, falciform. Belonging to the longipedicellatum-complex, and occurring sympatrically with E. heloisae. Pollination is exclusively by hummingbirds (Christianini & al. 2013).

References Aguilar-Rodríguez, P. A. [& al. 2014], Krömer, T. & MacSwiney G., M. C. (2014) The secrets of nightblooming bromeliads and bats. J. Bromeliad Soc. 64(3): 156–165, ills. Braun, P. J. (2005) Reise zum Fundort von Encholirium eddie-estevesii (Bromeliaceae) in Zentral-Brasilien. Kakt. and. Sukk. 56(2): 50–53, ills. Braun, P. J. & Esteves Pereira, E. (2006) Succulent and xeromorphic Bromeliads of Brazil. Part 3. Encholirium maximum, Orthophytum burle-marxii and Dyckia goehringii. Cact. Succ. J. (US) 78(4): 160–164, ills. https://doi.org/10.2985/0007-9367(2006)78[160:SAX BOB]2.0.CO;2 Braun, P. J. & Esteves Pereira, E. (2017) Ein [sic!] neue felsbewohnende Art der Gattung Encholirium aus dem äussersten Norden von Minas Gerais, Brasilien/A new rupicolous species of Encholirium from the uppermost north of Minas Gerais, Brazil: Encholirium viride. Bromelie 2017(1): 4–10, ills. Braun, P. J. & Esteves Pereira, E. (2018) Eine neue, isoliert wachsende Encholirium Art aus dem Norden von Goiás (Zentral-Brasilien)/A new isolated growing species of Encholirium from northern Goiás (central Brazil): Encholirium josinoi-narcisae. Bromelie 2018(1): 32–38, ills.

981 Cavallari, M. M. [& al. 2006], Forzza, R. C., Veasey, E. A., Zucchi, M. I. & Oliveira, G. C. X. (2006) Genetic variation in three endangered species of Encholirium (Bromeliaceae) from Cadeia do Espinhaço, Brazil, detected using RAPD markers. Biodivers. & Conservation 15: 4357–4373. https://doi.org/10.1007/s10531005-3741-5 Christianini, A. V. [& al. 2013], Forzza, R. C. & Buzato, S. (2013) Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads. Pl. Biol. (Stuttgart) 15(2): 360–368, ills. https://doi. org/10.1111/j.1438-8677.2012.00649.x Forzza, R. C. (2005) Revisão taxonômica de Encholirium Mart. ex Schult. & Schult. f. (Pitcairnioideae — Bromeliaceae). Bol. Bot. Univ. São Paulo 23(1): 1–49, ills., keys. https://doi.org/10.11606/issn.23169052.v23i1p1-49 Forzza, R. C. & Zappi, D. C. (2011) Side by side: Two remarkable new species of Encholirium Mart. ex Schult. & Schult. f. (Bromeliaceae) found in the Cadeia do Espinhaço, Minas Gerais, Brazil. Kew Bull. 66(2): 281–287, ills., key. https://doi.org/10.1007/s12225011-9283-y Gonçalves-Oliveira, R. C. [& al. 2017], Wöhrmann, T., Benko-Iseppon, A. M., Krapp, F., Alves, M., Wanderley, M. G. L. & Weising, K. (2017) Population genetic structure of the rock outcrop species Encholirium spectabile (Bromeliaceae): The role of pollination vs. seed dispersal and evolutionary implications. Amer. J. Bot. 104(6): 868–878, map. https:// doi.org/10.3732/ajb.1600410 Hmeljevski, K. V. [& al. 2015], Reis, M. S. dos & Forzza, R. C. (2015) Patterns of gene flow in Encholirium horridum L. B. Sm., a monocarpic species of Bromeliaceae from Brazil. J. Heredity 106(1): 93–101. https://doi.org/10.1093/jhered/esu067 Krapp, F. [& al. 2014], Pinangé, D. S. de B., BenkoIseppon, A. M., Leme, E. M. C. & Weising, K. (2014) Phylogeny and evolution of Dyckia (Bromeliaceae) inferred from chloroplast and nuclear sequences. Pl. Syst. Evol. 300(7): 1591–1614, ills., maps. https:// doi.org/10.1007/s00606-014-0985-0 Las-Casas, F. M. G. [& al. 2012], Azevedo Júnior, S. M. & Dias Filho, M. M. (2012) The community of hummingbirds (Aves: Trochilidae) and the assemblage of flowers in a Caatinga vegetation. Brazil. J. Biol. 72(1): 51–58. Leal, F. C. [& al. 2006], Lopes, A. V. & Machado, I. C. (2006) Polinização por beija-flores em uma area de caatinga no Municipio de Floresta, Pernambuco, Nordeste do Brasil. Revista Brasil. Bot. 29(3): 379–389. https://doi.org/ 10.1590/S0100-84042006000300005 Miller, L. J. (2009) There’s more than just cacti in eastern Brazil: Some bromeliads we encountered. Cact. Succ. J. (US) 81(3): 122–125, ills. https://doi.org/10.2985/ 015.081.0305 Nascimento, V. T. do [& al. 2012], Vasconcelos, M. A. da S., Maciel, M. I. S. & Albuquerque, U. P. (2012) Famine foods of Brazil’s seasonal dry forests: Ethnobotanical and nutritional aspects. Econ. Bot. 66(1): 22–34, ills. https://doi.org/10.1007/s12231-012-9187-2

982 Queiroz, J. A. [& al. 2016], Quirino, Z. G. M., Lopes, A. V. & Machado, I. C. (2016) Vertebrate mixed pollination system in Encholirium spectabile: A bromeliad pollinated by bats, opossum and hummingbirds in a tropical dry forest. J. Arid Environm. 125: 21–30, ills. https://doi.org/10.1016/j.jaridenv.2015.09. 015 Sazima, I. [& al. 1989], Vogel, S. & Sazima, M. (1989) Bat pollination of Encholirium glaziovii, a terrestrial bromeliad. Pl. Syst. Evol. 168(3–4): 167–179, ills. https:// doi.org/10.1007/BF00936097 Schütz, N. [& al. 2016], Krapp, F., Wagner, N. & Weising, K. (2016) Phylogenetics of Pitcairnioideae s.s. (Bromeliaceae): Evidence from nuclear and plastid

F. Krapp DNA sequence data. Bot. J. Linn. Soc. 181(3): 323–342, maps, ills. https://doi.org/10.1111/boj.12403 Siqueiro Filho, J. A. & Leme, E. M. C. (2007) Fragments of the Atlantic forest of Northeast Brazil: Biodiversity, conservation and the Bromeliads. Rio de Janeiro (BR): Andrea Jakobsson Estúdio. Smith, L. B. & Downs, R. J. (1974) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 1: Pitcairnioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/4393694 Zappi, D. C. & Taylor, N. P. (2011) A new species of Pilosocereus subgenus Gounellea, P. frewenii, from SE Brazil. Bradleya 29: 131–136, ills. https://doi.org/ 10.25223/brad.n29.2011.a16

Enchotia BROMELIACEAE U. Eggli

Enchotia Lemiaux (J. Bromeliad Soc. 61(3): 140, 2011). — Distr: Cultivated only. = Encholirium  Hechtia. The only known hybrid (‘Ruby’) is a cross between H. rosea (male)

 E. horridum (female), and was obtained by a Florida-based nursery. The leaves of the hybrid are described as spineless, which is surprising in view of the leaf morphology of both parental species.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_89

983

Fascicularia BROMELIACEAE U. Eggli

Fascicularia Mez (in Martius, Fl. Bras. 3(3): 627, 1891). Type: Bromelia bicolor Ruiz & Pavón. — Bromelioideae — Lit: Zizka & al. (1999: monograph). Distr: C-S Chile (Región V to Región X: 34–42
240 S, and probably even further S to 45
170 S); neophyte in W Europe. Etym: Lat., clustered, bundled; not explained, and presumably for the densely rosulate leaves. Incl. Hechtia Rivière (1871) (nom. illeg., Art. 53.1). Type: not typified. Perennial terrestrial or less often epiphytic rosette plants, stem none or short and stout; Ros solitary or in small groups; L numerous, densely arranged, ascending-spreading to spreading, L sheath broadly ovate to triangular-ovate, 3–9  1–4.5 cm, membranous and white at the base, coriaceous in the upper ½ and with spinose-serrate margin,  lepidote, glabrescent and veined towards the base, L lamina narrowly linear, stiff, attenuate into the pungent tip, (17–) 22–113 (133)  (0.6–) 0.8–1.8 (2.2) cm, upper face channelled or flat, lepidote to glabrate, lower face  densely lepidote, margin coarsely spinose, Sp 1–2.5 mm, antrorse or near the base spreading to retrorse, innermost L at flowering time bright red and

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

surrounding the inflorescence, with gradually reduced lamina; Inf terminal, simple, condensed and head-like,  sessile in the rosette centre, 3.5–9  3–9.5 cm; outer Bra ovate to oblong, as long or longer than the flowers, serrate, lamina short or absent; floral Bra to 7 cm, shorter than the flowers, ovate to oblong to spatulate, membranous, apically serrate, apical densely lepidote on the outer face; Fl numerous, densely arranged, sessile or shortly pedicellate, with a conspicuous epiygnous tube, (2.2–) 3.3–6.5 (7.5)  0.6–0.9 (1.2) cm, hermaphrodite, actinomorphic; Sep free, carinate, 10–24  3.5–8 mm, coriaceous, retuse to praemorse, with a short inconspicuous mucro, outer face lepidote-tomentose; Pet free, obtuse,  fleshy, 15–26  4–8 mm, deep to light blue (or occasionally almost white), with 2 small scales at the base; St included; Fil free, broadened at the base; Ov inferior, dorsally compressed, obconical; Sty elongate, included; Sti not contorted; Fr dry berries, to 7.4 cm (incl. perianth remains); Se ovoid to fusiform, 2.2–3  1–1.2 mm, dark brownish, rugose, without appendages. Similar to Ochagavia (see there for differences), and difficult to distinguish when not in flower. Traditionally regarded as having five species (Smith & Downs 1979: 1711–1715), the recent monograph by Zizka & al. (1999) concluded that only a single variable species (divided in two subspecies) can be recognized. Fascicularia

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_82

985

986

is amongst the most distinctly water-storing Bromelioideae (Horres & Zizka 1995). The leaves have a strongly developed water-storage tissue that makes up as much as 60% of leaf crosssectional area. Fascicularia groups with Ochagavia and Greigia in recent molecular phylogenies, and this group is amongst the earliest diverging genera of Bromelioideae, together with Deinacanthon (Schulte & al. 2005, Schulte & Zizka 2008, Schulte & al. 2009, Givnish & al. 2011). Silvestro & al. (2014) and Evans & al. (2015) found Fasicularia to be embedded in Ochagavia. The genus is not widely cultivated, but is fairly winter-hardy in the mild climate of S England. Roguenant & al. (2016: with ills.) report it as subspontaneous neophyte from S Ireland, S England and W France (Bretagne and several islands). F. bicolor (Ruiz & Pavón) Mez (in A. & C. de Candolle, Monogr. Phan. 9: 9, 1896). Type [neo]: Chile, Región VII Maule (Ochsenius s.n. [BR]). — Distr: As for the genus.  Bromelia bicolor Ruiz & Pavón (1802)  Billbergia bicolor (Ruiz & Pavón) Schultes fil. (1830)  Rhodostachys bicolor (Ruiz & Pavón) Bentham & Hooker fil. (1883). Description as for the genus. F. bicolor ssp. bicolor — Distr: C to S Chile (Región V to Región X [N Chiloé]); mostly on rocks near the coast; neophyte in W Europe Fig. 1 Fascicularia bicolor ssp. bicolor. (Copyright: E. J. Gouda)

U. Eggli

(Ireland, England, W France). I: Rauh (1990: 120: fig. 92, as F. pitcairniifolia); Nelson & Zizka (1997: 233); Roguenant & al. (2016: 568). – Fig. 1. Incl. Bromelia joinvillei E. Morren (1876)  Rhodostachys joinvillei (E. Morren) Bentham (1883); incl. Rhodostachys micrantha Philippi (1895)  Fascicularia micrantha (Philippi) Mez (1934); incl. Fascicularia parviflora Mez (1896). L succulent esp. towards the base, lamina (1–) 1.2–1.8 (2.2) cm wide, upper face flat or slightly convex, margins not recurved; floral Bra (3.2–) 4–5.8 (7) cm; Fl 3.7–6.5 (7.5) cm. F. bicolor ssp. canaliculata E. C. Nelson & Zizka (New Plantsman 4(4): 238, 1997). Type: Chile, Región X Valdivia (Sparre 3546 [S]). — Distr: S Chile (Región VII to Región X [Chiloé]); usually epiphytic in temperate evergreen forests. I: Roguenant & al. (2016: 569). Incl. Bromelia albobracteata Steudel (1857) (nom. inval., Art. 32.1c)  Rhodostachys albobracteata (Steudel) Baker (1889) (nom. inval., Art. 32.1c); incl. Hechtia joinvillei Rivière (1871); incl. Pourretia joinvillei hort. ex Chantin ex Rivière (1871); incl. Rhodostachys pitcairniifolia var. kirchhoffiana Wittmack (1890)  Fascicularia kirchhoffiana (Wittmack) Mez (1919). L not markedly succulent towards the base, lamina (0.6–) 0.8–1.1 (1.4) cm wide, upper face channelled, margins distinctly recurved; floral Bra (2.2–) 3–3.8 cm; Fl (2.2–) 3.3–4.4 cm.

Fascicularia BROMELIACEAE

References Evans, T. M. [& al. 2015], Jabaily, R. S., Gelli de Faria, A. P., Oliveira F. de Sousa, L. de, Wendt, T. & Brown, G. K. (2015) Phylogenetic relationships in Bromeliaceae subfamily Bromelioideae based on chloroplast DNA sequence data. Syst. Bot. 40(1): 116–128. https://doi. org/10.1600/036364415X686413. Givnish, T. J. [& al. 2011], Barfuss, M. H. J., Ee, B. van, Riina, R., Schulte, K., Horres, R., Gonsiska, P. A., Jabaily, R. S., Crayn, D. M., Smith, J. A. C., Winter, K., Brown, G. K., Evans, T. M., Holst, B. K., Luther, H., Till, W., Zizka, G., Berry, P. E. & Sytsma, K. J. (2011) Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: Insights from an eight-locus plastid phylogeny. Amer. J. Bot. 98(5): 872–895, ills., maps. https://doi.org/10.3732/ajb.1000059. Horres, R. & Zizka, G. (1995) Untersuchungen zur Blattsukkulenz bei Bromeliaceae. Beitr. Biol. Pfl. 69(1): 43–76, ills. Nelson, E. C. & Zizka, G. (1997) Fascicularia (Bromeliaceae): Which species are cultivated and naturalized in northwestern Europe? New Plantsman 4(4): 232–239, ills. Rauh, W. (1990) Bromelien. Ed. 3. Stuttgart (DE): Verlag Eugen Ulmer. Roguenant, A. [& al. 2016], Lecoufle, M. & Raynal-Roques, A. (2016) Les Broméliacées. Approche panoramique d’une grand famille “américaine”. Paris (FR): Belin.

987 Schulte, K. & Zizka, G. (2008) Multi locus plastid phylogeny of Bromelioideae (Bromeliaceae) and the taxonomic utility of petal appendages and pollen characters. Candollea 63(2): 209–225. http://www.ville-ge.ch/cjb/ publications_candollea_632.php Schulte, K. [& al. 2005], Horres, R. & Zizka, G. (2005) Molecular phylogeny of Bromelioideae and its implications on biogeography and the evolution of CAM in the family. Senckenberg. Biol. 85(1): 113–125, maps. Schulte, K. [& al. 2009], Barfuss, M. H. & Zizka, G. (2009) Phylogeny of Bromelioideae (Bromeliaceae) inferred from nuclear and plastid DNA loci reveals the evolution of the tank habit within the subfamily. Molec. Phylogen. Evol. 51(2): 327–339. https://doi.org/ 10.1016/j.ympev.2009.02.003. Silvestro, D. [& al. 2014], Zizka, G. & Schulte, K. (2014) Disentangling the effects of key innovations on the diversification of Bromelioideae (Bromeliaceae). Evolution 68(1): 163–175. https://doi.org/10.1111/ evo.12236. Smith, L. B. & Downs, R. J. (1979) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 3: Bromelioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/sta ble/i399940 Zizka, G. [& al. 1999], Horres, R., Nelson, E. C. & Weising, K. (1999) Revision of the genus Fascicularia Mez (Bromeliaceae). Bot. J. Linn. Soc. 129(4): 315–332, map, key. https://doi.org/10.1111/j.10958339.1999.tb00507.x.

Fosterella BROMELIACEAE U. Eggli

Fosterella L. B. Smith (Phytologia 7: 171, t. 1, 1960). Type: Pitcairnia micrantha Lindley. — Pitcairnioideae — Lit: Smith & Downs (1974: 199–209, Fl. Neotropica); Rex & al. (2007: molecular phylogeny); Ibisch & al. (2008: key to species); Rex & al. (2009: molecular phylogeny); Peters (2009: monograph); Wagner & al. (2013: evolution); Lima Silva & al. (2016: cytology). Distr: N, C & S America (S Mexico, El Salvador, Guatemala, Brazil, Peruvian Andes to Bolivia, Paraguay and N Argentina), 100–2750 m. Etym: For Mulford Bateman Foster (1888–1978), US-American horticulturist and plant collector, and co-founder and first president of the US-American Bromeliad Society; and Lat. diminutive suffix, for the small flowers. Perennial terrestrial or saxicolous rosette plants, mesophytic to xerophytic, acaulescent or shortly caulescent, offsetting; L 5–30, flat or  ascending, sheath broadly ovate to triangular, entire or slightly serrate, glabrous or upper face slightly lepidote, somewhat succulent, lamina linear, narrowly triangular or narrowly to broadly lanceolate or oblanceolate, rarely basally much narrowed and petiolate, 10–100  1–8 cm, thin or basal part somewhat succulent, entire or

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

serrate near the base, rarely somewhat undulate, upper face glabrous or sparsely lepidote towards the base, lower face  densely covered with peltate or stellate hairs; Inf pedunculate, indeterminate, racemose to paniculate with up to 30 branches with up to 3.-order branching, axis green or reddish, glabrous, villous or cobwebby, rarely glaucous; peduncular Bra  appressed, narrowly triangular, the lower often leaf-like, shorter than the internodes and sometimes imbricate, entire or slightly serrate, green, reddish or straw-coloured; floral Bra broadly ovate to triangular, acute, shorter or longer than the pedicels, lower face glabrous or variously hairy to lepidote; Fl spreading or secund, erect or pendent, inconspicuous, mostly whitish, hermaphrodite, actinomorphic, sessile or with Ped to 15 mm; Sep 3, free, convolute, much shorter than the petals, to 9 mm, glabrous or hairy, rarely glaucous; Pet 3, free, 3–24 mm, straight, recurved during anthesis and straight afterwards, or revolute during anthesis and afterwards, without appendage; St in 2 whorls, exserted, the outer free, the inner adnate to the petals; Anth basifixed, linear, coiled at anthesis; Ov superior, deeply grooved; Sty short to long; Sti complex simple-erect or conduplicate-spiral; Fr basically septicidal capsules; Se numerous, filiform or fusiform, 2–4 mm, brown with white membranous appendage, bicaudate. — Cytology: 2n = 50, polyploids (to 2n = 150) comparatively common (Peters 2009: 20, summarizing earlier studies by several authors).

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_83

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A genus of 31 species (Peters 2009), traditionally placed in Pitcairnioideae, where it is placed as sister to a clade that includes Encholirium, Dyckia and Deuterocohnia in the molecular phylogenies of Givnish & al. (2007), Rex & al. (2009) and Schütz & al. (2016). Rex & al. (2009) found a basal dichotomy in 2 major clades, and a total of 6 major lineages, which are only informally named (clade 1: albicans group, rusbyi group, micrantha group, weberbaueri group; clade 2: penduliflora group, weddelliana group (Peters 2009). The genus originated in the late miocene ( 9.6 mybp) during the last rapid Andean uplift, and the ancestral habitat has been reconstructed as seasonally deciduous tropical forest and azonal lowland sites (Wagner & al. 2013). The same authors found that the disjunct occurrence of 2 species in S Mexico, C America and Amazonia is due to 2 recent long-distance dispersal events. Speciation is proposed to have been mostly allopatric. The diversification rate is low, and Andean uplift did not enhance speciation rate. Many of the species are narrow rare endemics esp. in the arid to semihumid habitats of the NE Andean slopes of Bolivia. Wagner & al. (2015) found that at least some species are partially self-fertile, and that hybrids are easily formed amongst the majority of the 4 taxa studied, indicating that reproduction barriers are weak or absent. A possible important role for hybridization in the evolution of the genus was also suggested by Lima Silva & al. (2016) on the base of chromosomal features. The same authors found that polyploidization events could be comparatively recent, at least in some taxa. Succulence is present in the vast majority of the species, but usually confined to the midvein-areas of the basal-most parts of the lamina. In some taxa from arid or semi-arid places, succulence is more pronounced, and these are covered below. The account is based entirely on the data presented in the monograph by Peters (2009), where illustrations of all taxa can also be found. F. albicans (Grisebach) L. B. Smith (Phytologia 7: 171, 1960). Type: Argentina, Salta (Lorentz & Hieronymus 502 [GOET, B, CORD]). — Lit: Peters (2009: 27–34, with ills.). Distr: S & W Bolivia (La Paz, Cochabamba, Santa Cruz, Chuquisaca,

U. Eggli

Tarija), N Argentina (Jujuy, Salta, Tucumán); clearings in Yungas vegetation, 550–2650 m. – Fig. 1.  Cottendorfia albicans Grisebach (1879)  Lindmania albicans (Grisebach) Mez (1896); incl. Fosterella fuentesii Ibisch & al. (2002). Subcaulescent, colony-forming, flowering to 100 ( 180) cm tall; L 20–30, irregularly erect to spreading, sheath 5–7 cm wide, entire, glabrous, lamina narrowly oblanceolate, acuminate, basally narrowing, 40–100  2–3 ( 4) cm, succulent, serrate towards the base, upper face sparsely lepidote, lower face completely covered by interwoven peltate hairs; Inf 70–180 cm, erect, paniculate with 1.- and 2.-order branches, peduncle 50–120 cm, densely cobwebby; peduncular Bra 4–12 cm, longer than the internodes, entire; primary Bra 2–3 cm, longer than the sterile branch base; Br 8–20, inclined, arcuate, to 18 cm, to 50-flowered, with 2–5 cm long secondary branches; floral Bra 4–5 mm; Fl spreading, suberect, subsessile; Sep 3–4 mm, green, sparsely cobwebby; Pet 6–8 mm, white, recoiled like a watchspring. The species is both morphologically and ecologically variable and similar to the more mesophytic F. caulescens (not treated here). F. cotacajensis M. Kessler & al. (Revista Soc. Boliv. Bot. 2(2): 111, 1999). Type: Bolivia, Cochabamba (Kessler 9620 [LPB, SEL]). — Lit: Peters (2009: 56–60, with ills.). Distr: C-W Bolivia (La Paz, Cochabamba); inter-Andean dry semi-deciduous forests, sunny dry rocky slopes, 1150–2200 m. Caulescent, flowering to 100 cm tall; stems to 30 cm, sparsely offsetting; L 15–25, densely spreading-arching at the top parts of the stems, sheath to 3 cm wide, entire, glabrous, lamina lanceolate, 15–45  to 2 cm, succulent, serrate towards the base, upper face sparsely lepidote, lower face thickly covered with interwoven peltate trichomes; Inf to 100 cm, erect, paniculate with 1.- and 2.-order branches, peduncle 20–50 cm, glabrous, glaucous; peduncular Bra 3–5 cm, longer than the internodes, slightly serrate; primary Bra 1–3 cm, as long as or longer than the sterile branch base; Br 10–15, ascending, straight, to 30 cm, to 25-flowered, with 2–3 cm long secondary branches; floral Bra 4–7 mm; Fl secund, pendulous; Ped 3–4 mm; Sep 5 mm, green, glabrous; Pet 8 mm, white, recoiled like a watchspring.

Fosterella BROMELIACEAE

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Fig. 1 Fosterella albicans. (Copyright: E. J. Gouda)

Closely related to the morphologically similar F. weddelliana from lower and less arid habitats, but differing in narrower leaves and longer floral bracts and sepals. F. penduliflora (C. H. Wright) L. B. Smith (Phytologia 7: 172, 1960). Type: Peru (Forget s.n. [K]). — Lit: Peters (2009, 105–113, with ills.). Distr: Peru?, Bolivia (La Paz, Cochabamba, Santa Cruz, Chuquisaca, Tarija), N Argentina (Jujuy, Salta), Yungas vegetation and deciduous to evergreen inter-Andean dry forests and montane Chaco vegetation, 200–2650 m. I: Rauh (1990, 272, fig. 341). – Fig. 2.  Catopsis penduliflora C. H. Wright (1910)  Lindmania penduliflora (C. H. Wright) Stapf (1924); incl. Fosterella chiquitana Ibisch & al. (1999); incl. Fosterella latifolia Ibisch & al. (1999). Acaulescent, flowering to 120 cm tall; Ros open, flat, sparsely offsetting; L 8–15, sheath 2–5 cm wide, entire, glabrous, lamina lanceolate, acuminate, narrowed towards the base, 20–40  2.5–8 cm, succulent, entire, upper face glabrous, lower face often flushed reddish, sparsely lepidote; Inf to 120 cm,

erect, paniculate, with 1.- and 2.-order branches, peduncle 25–60 ( 85) cm, glabrous, often glaucous; peduncular Bra 1–3 cm, as long as the internodes, often reddish, sparsely lepidote; primary Bra 0.5–1.5 cm, shorter than the sterile branch base; Br 10–15, inclined, arcuate, 8–22 cm, to 35-flowered, with 3–5 cm long secondary branches; floral Bra 2–4 ( 10) mm, entire, often reddish; Fl secund, pendulous; Ped 2–6 mm; Sep 2–3 mm, green or often reddish, glabrous; Pet 7–10 mm, white, recurved. The most common species, both in the field and in cultivation. The type was reported from Peru, but apart from a second collection from Peru, also without locality, the taxon is only known from Bolivia and N Argentina, and its occurrence in Peru is doubtful. F. rexiae Ibisch & al. (Selbyana 23(2): 213, 2002). Type: Bolivia, La Paz (Vásquez & Gerlach 3673 [LPB]). — Lit: Peters (2009: 119–122, with ills.). Distr: W Bolivia (La Paz: near Caranavi); evergreen Yungas vegetation, 800–1300 m; known from the type locality only. Caulescent, flowering to 90 cm tall; Ros open, semiglobose, sparsely offsetting; L 10–15, sheath

992

Fig. 2 Fosterella penduliflora. (Copyright: E. J. Gouda)

2.5 cm wide, entire, glabrous, lamina linear to narrowly lanceolate, acuminate, narrowed towards the base, 20–30  1.7 cm, succulent, serrate towards the base, upper face towards the base and lower face entirely with dense appressed peltate trichomes; Inf to 90 cm, erect, paniculate, with 1.and 2.-order branches, peduncle to 55 cm, slightly cobwebby becoming glabrescent; peduncular Bra 5 cm, as long or longer than the internodes, entire, lower face slightly cobwebby; primary Bra 1.5 cm, longer than the sterile branch base, slightly cobwebby; Br 15–18, inclined, arcuate, 8–10 cm, to 20-flowered, with 3–5 cm long secondary branches; floral Bra 2 mm, entire, slightly cobwebby; Fl spreading, erect, sessile; Sep 3 mm, green to reddish, sparsely long-hairy, glabrescent; Pet 5 mm, white, recoiled like a watchspring. Closely related to the similar F. albicans, but with flatter rosette, less tomentose leaves, and less densely flowered inflorescences. F. rusbyi (Mez) L. B. Smith (Phytologia 7: 172, 1960). Type: Bolivia, La Paz (Bang 2571 [B, BM, F, G, GH, K, MO, NY, US]). — Lit: Peters (2009: 132–138, with ills.). Distr: W

U. Eggli

Bolivia (La Paz, Cochabamba); evergreen to deciduous Yungas and inter-Andean dry forests, terrestrial or saxicolous, 700–1900 m. I: Rauh (1990: 272, fig. 342). – Fig. 3.  Lindmania rusbyi Mez (1901); incl. Fosterella elata H. Luther (1981). Acaulescent, flowering to 200 cm tall; Ros open, flat, sparsely offsetting; L 10–15, sheath 5–6 cm wide, entire, glabrous, lamina linear to narrowly lanceolate, acuminate, narrowed towards the base, 20–45  1–6 cm, succulent, serrate towards the base, undulate, upper face lepidote towards the base, lower face densely white-hairy with peltate trichomes; Inf to 200 cm, erect, paniculate, with 1.- and 2.-order branches, peduncle 30–120 cm, glabrous; peduncular Bra 4–10 cm, longer than the internodes, entire, lower face cobwebby; primary Bra 0.6–1 ( 2.5) cm, shorter than or as long as the sterile branch base, lepidote, glabrescent; Br 10–15, inclined, arcuate, 8–15 cm, to 15-flowered, with 5–8 cm long secondary branches; floral Bra 2–3 mm, entire, glabrous; Fl secund, pendulous; Ped 1.5–2 mm; Sep 2–3 mm, green, glabrous; Pet 7 mm, greenish, cream or rose-coloured, recoiled like a watchspring. Similar and closely related to F. vasquezii, but with broader, serrate and undulate leaves, larger inflorescences and longer petals. F. vasquezii E. Gross & Ibisch (J. Bromeliad Soc. 47(5): 212, ills. (pp. 214–215, 1997). Type: Bolivia, Santa Cruz (Ibisch & al. 93.0652 [HEID, LPB]). — Lit: Peters (2009: 148–151, with ills.). Distr: E Bolivia (Santa Cruz); transition between humid and Chiquitano dry forests, 200–850 m. Acaulescent, flowering to 50 cm tall; Ros open, flat, sparsely offsetting; L 10–15, sheath 5 cm wide, obscurely serrate, glabrous, lamina lanceolate, 25–45  1.5–3.5 cm, succulent, obscurely serrate towards the base, undulate, upper face glabrous, lower face with a dense thick layer of peltate hairs; Inf to 50 cm, erect, paniculate, with 1.- and 2.-order branches, peduncle 20–30 cm, glabrous; peduncular Bra 3 cm, longer than the internodes, entire, lower face with interwoven hairs; primary Bra 1 cm, shorter than the sterile branch bases, glabrous; Br 10–20, ascending, straight, 10–15 cm, to 10- to 25-flowered, with 3–5 cm long secondary branches; floral Bra 2–3 mm, entire, glabrous; Fl secund,

Fosterella BROMELIACEAE

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Fig. 3 Fosterella rusbyi. (Copyright: E. J. Gouda)

pendulous; Ped 2–3 mm; Sep 2 mm, green, glabrous; Pet 5 mm, white, recoiled like a watchspring. Closely related to F. rusbyi (see there for differences). F. weddelliana (Brongniart ex Baker) L. B. Smith (Phytologia 7: 172, 1960). Type: Bolivia, La Paz (Weddell 4233 [P, B]). — Lit: Peters (2009: 162–167, with ills.). Distr: W-C Bolivia (La Paz), evergreen to deciduous Yungas and inter-Andean dry forests, terrestrial or saxicolous, 750–2750 m.  Cottendorfia weddelliana Brongniart ex Baker (1889)  Lindmania weddelliana (Brongniart ex Baker) Mez (1896); incl. Fosterella nowickii Ibisch & al. (2002). Caulescent, flowering to 150 cm tall; Ros open, very sparsely offsetting; L 8–15, irregularly spreading to arching, sheath 5–7 cm wide, serrate, glabrous, lamina narrowly lanceolate, acuminate to acute, 15–50  2–3.5 cm, succulent, serrate towards the base, upper face lepidote towards the base, lower face with a thick layer of interwoven peltate hairs; Inf to 150 cm, erect, paniculate, with 1.- and 2.order branches, peduncle 30–80 cm, green to reddish, glabrous, glaucous, peduncular Bra 5–8 cm, longer than the internodes, serrate, with interwoven hairs, primary Bra 2.5–5 cm, as long as or longer than the sterile branch base, entire to obscurely serrate, hairy, glabrescent; Br 8–25, ascending to

inclined, 10–15 cm, to 30-flowered, with up to 5 cm long secondary branches; floral Bra 2–3 mm, entire, glabrous; Fl secund, pendulous; Ped 2–4 mm; Sep 2–3 mm, green to reddish, glabrous; Pet 4–5 mm, white to greenish or reddish, recoiled like a watchspring; Fr 4–5  2–3 mm. Originally incorrectly described as acaulescent. Closely related to F. cotacajensis (see there for differences). F. windischii L. B. Smith & Read (Bradea 6(15): 137, 1992). Type: Brazil, Mato Grosso (Windisch 2044 [US, HB]). — Lit: Peters (2009: 168–171, with ills.). Distr: E Bolivia (Santa Cruz), adjacent Brazil (Mato Grosso); dry rock slopes of Caparús table mountain in evergreen humid lowland forest, 300–550 m. Acaulescent, flowering to 60 cm tall; Ros open, sparsely offsetting; L 6–12, arching-spreading, sheath to 2.5 cm wide, entire, glabrous, lamina narrowly lanceolate, acuminate, narrowed towards the base, 30–50  1.5–2.5 cm, succulent towards the base, entire or obscurely serrate towards the base, upper face glabrous, lower face with a thick layer of interwoven hairs; Inf to 60 cm, erect, compound-racemose, peduncle to 30 cm, glabrous; peduncular Bra 2 cm, longer than the internodes, entire, lower face with interwoven hairs; primary Bra 0.5 cm, shorter than the sterile branch base,

994

lepidote; Br 6–15, ascending, straight, 6–10 cm, to 20-flowered; floral Bra 1–2 mm, entire, glabrous; Fl secund, pendulous; Ped 2–3 mm; Sep 2–3 mm, green, glabrous; Pet 4–5 mm, white, recoiled like a watchspring. According to molecular data part of a group that includes F. vasquezii and F. yuvinkae. F. yuvinkae Ibisch & al. (Selbyana 23(2): 216, 2002). Type: Bolivia, Santa Cruz (Reichle P-SR1 [LPB]). — Lit: Peters (2009: 172–176, with ills.). Distr: Bolivia (Santa Cruz); Chiquitano dry deciduous forest, steep rock slopes or sandy ground, 400–1200 m. Acaulescent, flowering to 100 cm tall, sparsely offsetting; Ros open, flat; L 15–20, sheath 2–3 cm wide, entire, villous, lamina linear to narrowly oblanceolate, acuminate, narrowing towards the base, 20–35  1.5–3 cm, succulent towards the base, entire, upper face glabrous, lower face densely tomentose with stellate hairs; Inf to 60 cm, erect, paniculate, with 1.- and 2.-order branches, peduncle 30–50 cm, glabrous, glaucous; peduncular Bra 2–4 cm, longer than the internodes, entire, lower face long-hairy; primary Bra 0.5–1 cm, shorter than the sterile branch base, long-hairy, glabrescent; Br to 15, inclined, arcuate, 8–20 cm, to 20-flowered, with 5–7 cm long secondary branches; floral Bra 2 mm, entire, glabrous; Fl secund, pendulous; Ped 2–4 mm; Sep 3–4 mm, green, glabrous; Pet 7–9 mm, only 1.5 mm wide, white, recurved. Similar to F. penduliflora but with the leaves densely tomentose on the lower face. The species is unique in the genus because of the very narrow petals. Molecular data shows a close relationship to F. vasquezii and F. windischii, which all occur on the Brazilian Shield.

References Givnish, T. J. [& al. 2007], Millan, K. C., Berry, P. E. & Sytsma, K. J. (2007) Phylogeny, adaptive radiation, and

U. Eggli historical biogeography of Bromeliaceae inferred from ndhF sequence data. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution: Poales. Aliso 23: 3–26. https://doi.org/10.5642/aliso.2007 2301.04 Ibisch, P. L. [& al. 2008], Read, R. & Peters, J. (2008) Key to the species of the genus Fosterella (Bromeliaceae). Selbyana 29(2): 195–198. http://www.jstor.org/stable/ 41760333 Lima Silva, H. M. de [& al. 2016], Vasconcelos, E. V., Benko-Iseppon, A. M., Wagner, N., Weising, K. & Brasileiro-Vidal, A. C. (2016) Chromosomal features of Fosterella species (Bromeliaceae, Pitcarnioideae). Bot. J. Linn. Soc. 181(3): 532–541, ills. https://doi.org/ 10.1111/boj.12390 Peters, J. (2009) Revision of the genus Fosterella (Bromeliaceae). Kassel (DE): Universität Kassel, Fachbereich Naturwissenschaften, unpublished Ph.D. thesis. http://nbn-resolving.de/urn:nbn:de:hebis:34-2009 081429441 Rauh, W. (1990) Bromelien. Ed. 3. Stuttgart (DE): Verlag Eugen Ulmer. Rex, M. [& al. 2007], Patzold, K., Schulte, K., Zizka, G., Vásquez, R., Ibisch, P. L. & Weising, K. (2007) AFLP analysis of genetic relationships in the genus Fosterella L. B. Smith (Pitcairnioideae, Bromeliaceae). Genome 50(1): 90–105. https://doi.org/10.1139/g06-141 Rex, M. [& al. 2009], Schulte, K., Zizka, G., Peters, J., Vásquez, R., Ibisch, P. L. & Weising, K. (2009) Phylogenetic analysis of Fosterella L. B. Sm. (Pitcairnioideae, Bromeliaceae) based on four chloroplast DNA regions. Molec. Phylogen. Evol. 51(3): 472–485. https://doi.org/ 10.1016/j.ympev.2009.01.001 Schütz, N. [& al. 2016], Krapp, F., Wagner, N. & Weising, K. (2016) Phylogenetics of Pitcairnioideae s.s. (Bromeliaceae): Evidence from nuclear and plastid DNA sequence data. Bot. J. Linn. Soc. 181(3): 323–342, maps, ills. https://doi.org/10.1111/boj.12403 Smith, L. B. & Downs, R. J. (1974) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 1: Pitcairnioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/4393694 Wagner, N. D. [& al. 2015], Wöhrmann, T., Öder, V., Burmeister, A. & Weising, K. (2015) Reproduction biology and chloroplast inheritance in Bromeliaceae: A case study in Fosterella (Pitcairnioideae). Pl. Syst. Evol. 301 (9): 2231–2246, map, ills. https://doi.org/10.1007/ s00606-015-1226-x Wagner, N. D. [& al. 2013], Silvestro, D., Brie, D., Ibisch, P. L., Zizka, G., Weising, K. & Schulte, K. (2013) Spatiotemporal evolution of Fosterella (Bromeliaceae) in the Central Andean biodiversity hotspot. J. Biogeogr. 40(5): 869–880. https://doi.org/10.1111/jbi.12052

Hechcohnia BROMELIACEAE N. Schütz and F. Krapp

Hechcohnia G. H. Anderson ex J. Grant (Selbyana 19: 117, 1998). — Distr: Cultivated only. = Hechtia  Deuterocohnia. The only known cross (D. schreiteri (female)  H. glomerata) remained unnamed (Anderson 1986: 103). The plants and the leaf spines are more delicate than in either parent, and

the plant colour varies from green to light rose.

References Anderson, G. H. (1986) Hybrids by design and by chance. J. Bromeliad Soc. 36(3): 99–103, 109, ills.

N. Schütz (*) Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany e-mail: [email protected] F. Krapp Guxhagen, Germany e-mail: fl[email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_90

995

Hechtia BROMELIACEAE A. Espejo-Serna, A. R. López-Ferrari, and N. Martínez-Correa

Hechtia Klotzsch (Allg. Gartenzeitung 3: 401, 1835). Type: Hechtia stenopetala Klotzsch. — Hechtioideae — Lit: Mez (1896: synopsis); Mez (1935: 351–360, synopsis); Smith & Downs (1974: 577–604, Fl. Neotropica); McVaugh (1989: Fl. Novo-Galiciana); Utley & Burt-Utley (1994: Fl. Mesoamericana); Espejo-Serna & al. (2005: Fl. Veracruz); Espejo-Serna & al. (2010a: Fl. Bajio); Ramírez Morillo & al. (2014a: growth patterns). Distr: S USA (Texas), Mexico (widespread) and N C America (Guatemala, Honduras, El Salvador, Nicaragua, Belize). Etym: For Julius Gottfried Conrad Hecht (1771–1837), German botanist and counselor to the King of Prussia. Incl. Bakeria André (1889) (nom. illeg., Art. 53.1). Type: Bakeria tillandsioides André. Incl. Bakerantha L. B. Smith (1934). Type: Bakeria tillandsioides André. Incl. Niveophyllum Matuda (1965). Type: Niveophyllum caeruleum Matuda. Perennial xerophytic rosette plants, dioecious, polycarpic or monocarpic, caespitose or rarely solitary, terrestrial and frequently saxicolous;

A. Espejo-Serna (*) · A. R. López-Ferrari · N. MartínezCorrea Herbario Metropolitano, Depto. Biología, División de Ciencias Biológicas y de la Salud, Universidad Autónoma Metropolitana Iztapalapa, Ciudad de México, Mexico e-mail: [email protected]; [email protected]; [email protected]

Ros stemless or sometimes shortly to distinctly caulescent (H. mooreana, H. myriantha), rarely rhizomatous (H. caulescens) or more rarely stoloniferous (H. pretiosa); Ros small to medium or rarely very large (H. myriantha), dense; L longlived, succulent and xeromorphic, narrowly triangular to linear, lepidote throughout or only abaxially, tip pungent, rarely soft (H. lundelliorum, H. purpusii, H. tillandsioides); L margins strongly spiny or more rarely minutely serrate (H. lundelliorum, H. purpusii, H. tillandsioides); Sp uncinate, retrorse, antrorse and/or divaricate; Inf erect or ascending to rarely pendent, paniculate, rarely racemose, 0.5–4 m (= overall size, i.e. peduncle and floriferous part), 1–3 branched, terminal or seemingly lateral by reduction of the floriferous rosette; lower peduncular Bra similar to innermost rosette leaves, diminishing in size upwards; peduncle long to short, cylindrical to complanate; Inf branches long racemose to capitate; Fl actinomorphic, unisexual (and plants dioecious), sessile to long pedicellate; Ped frequently articulated with the rhachis; Per campanulate, very rarely tubular (H. pretiosa), usually white, greenish or yellowish, rarely rose, lilac, red or reddish; Sep 3, free, sepaloid; Pet 3, free or rarely basally connate (H. pretiosa), petaloid, naked; male Fl with St longer or shorter than the petals, equal to subequal, Fil white, rarely greenish, filiform or linear to triangular and flattened, Anth dorsifixed, oblong, Ov vestigial; female Fl with Ov superior to inferior, rarely inferior (H. epigyna, H. malvernii), ovoid to long ovoid or ellipsoid, glabrous to lepidote, 3-locular, ovules numerous, Sty absent or rarely present

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_84

997

998

(H. guatemalensis), Sti 3-branched erect to recurved branches, Fil of the staminodes flattened, triangular, antheroids present or sometimes absent; Fr dehiscent septicidal and loculicidal capsules, ellipsoid to ovoid, sometimes triquetrous, sepals, petals and stigma branches persistent at the apex; Se numerous, fusiform, circumalate and bicaudate. A genus with 69 species, 63 (92.5%) of them endemic to Mexico (Espejo-Serna 2012), and entirely endemic to the region termed “Megamexico 3” by Rzedowski (1991). The genus is still not well known, and not less than 17 species (28% of the genus) have been described in the past 7 years, and several more remain to be described. No subgeneric classification has been established, and no phylogenetic analysis is available at this time, even though some subgroups like the H. podantha, H. glomerata, H. tillandsioides, and H. zamudioi complexes, can be circumscribed. Dioecy, as exhibited by Hechtia, is unique in the family with the exception of andromonoecy in Cryptanthus and a few isolated taxa of Aechmea, Androlepis and Catopsis, and gynomonoecy in a few isolated taxa of Dyckia. There is no sexual dimorphism in the vegetative characters, but the male and female flowers can appear quite different (e.g. H. confusa). Flowering is always terminal, but the flowering rosette is pushed into a seemingly lateral position in some species, and appears completely lateral when the flowering rosette is prematurely overtopped by the offset it produces before starting to flower. Truly lateral flowering, as discussed by Ramírez Morillo & al. (2014a), does not occur in the genus. All taxa exhibit  succulent leaves, varying from rigid (H. podantha, H. roseana) to soft and flexible (H. tillandsioides), and the genus is thus covered in its entirety below. Hechtia species inhabit xeric vegetation from arid shrublands to tropical deciduous and semideciduous forests, with few taxa from humid places such as oak forests or canyons and river sides. The following names are of unresolved application but are referred to this genus: Bromelia antiacantha Bertoloni (1824)  Agallostachys antiacantha (Bertoloni) Beer (1856); Hechtia

A. Espejo-Serna et al.

cordylinoides Baker (1881); Hechtia longifolia hort. ex Baker (1889); Hechtia striata Lemaire (1846); Hechtia suaveolens E. Morren ex Mez (1896). H. aquamarina I. Ramírez & Jiménez (Phytotaxa 2012(33): 1, 2012). Type: Mexico, Puebla (Ramírez & Carnevali 1689 [CICY, B, G, K, MEXU, MO, SEL, UAMIZ, XAL, WU]). — Lit: Espejo-Serna & al. (2007: as Hechtia sp. 3); Ramírez Morillo & Jimenez-Nah (2012: as H. pueblensis). Distr: C Mexico (SE Puebla, NW Oaxaca); xerophytic shrublands, >900 m.  Hechtia pueblensis I. Ramírez & Jiménez (2012) (nom. illeg., Art. 53.1). Caespitose, flowering 80 cm tall; Ros 35–60 cm ∅; L 12–16, sheaths pale brown, transversely oblong, 3–3.5  3.5–5 cm, entire, glabrous, L lamina glaucous, 22–30  2–2.5 cm, densely white-lepidote abaxially and white-lepidote adaxially, Sp antrorse, 4–5 mm, 1–1.5 cm apart; male Inf 2 branched, with 55 primary Br 10–23 cm long, secondary Br 2, basal, present only on the basal primary branches; floral Bra brownish, ovate, 2.2–2.4  1.1–1.3 mm, acuminate, shorter than the sepals; Ped 1.2–1.4 mm; Fl fragrant; Sep green, ovate oblong, 2.1–2.2  1.8–2 mm, acute; Pet pale green, elliptic, 4–5  2.3–2.5 mm; St subequal, longer than the petals; Fil 4.3–5 mm; Anth green, 1.8–2 mm, mucronate; female Inf 1 branched, with 15 primary Br 18–20 cm long; floral Bra brownish, widely triangular, 1–1.4  1.4 mm, minutely serrate, shorter than the sepals; Ped 15–17  6–7 mm ∅; Fl fragrant; Sep pale green, triangular, 2.2  2 mm, obtuse; Pet green, long triangular, 3.7–4.5  2 mm; Ov pale rose, ellipsoid, 7  2.8 mm; Fr light brown, 6–9  3.5–5.8 mm; Se brown, 2.8–5.7  0.87–1.2 mm. H. bracteata Mez (in A. & C. de Candolle, Monogr. Phan. 9: 550, 1896). Type: Mexico, Veracruz (Müller 813 [W?, B, NY]). — Lit: Espejo-Serna & al. (2005); Burt-Utley & Utley (2011). Distr: C Mexico (Puebla, Veracruz); shrubland and tropical deciduous forests on calcareous soils, 1880–2050 m. Caespitose, flowering 2–3 m tall; Ros globose, to 70 cm high and 50 cm ∅; L numerous, sheaths

Hechtia BROMELIACEAE

pale yellow to light brown, widely ovate to square or transversely elliptic, 6–7.5  6.6–12 cm, serrate, densely lepidote at the apical portion on both faces, L lamina light green, 37.6–60  1–3.5 cm, glabrous adaxially, white-lepidote abaxially, Sp dark brown to black, uncinate, antrorse, 3–4 mm, 1.5–2 cm apart; Inf floccose-tomentose, 2 branched; male Inf with 21 primary Br 2.5–8 cm long, secondary Br 2, basal; floral Bra light brown, ovate to widely ovate, 4–8  2.4 mm, acuminate, erose, glabrous, longer than the sepals; Fl polystichous, divaricate, 3.6  1.9–3.1 mm, densely arranged; Ped 2–2.5 mm, lepidote; Sep brown, ovate to broadly ovate, 2.5–4  1.2–2 mm, entire, glabrous, acute; Pet shortly connate at the base, yellowish, oblong to elliptic, 3.5–5  1.6–2 mm, entire, rounded; St equal, shorter than the petals; Fil 3–3.1 mm; Anth yellow, 1–2 mm; female Inf with 60 or more primary, short to capituliform primary Br 2–3.5 cm long; secondary Br 2, basal; floral Bra brown, ovate, 4.5–6.9  1.1–4.2 mm, acute, entire, glabrous, longer than the sepals; Ped 2.4–4.3 mm, lepidote; Sep brownish, ovate, 3–4.5  1.4–2 mm, entire, acute; Pet yellowish, triangular, 3.9–4.2  1.4–1.6 mm, entire, acute; Ov green, ovoid, 8.5  6.2 mm ∅; Fr brownish, 7–9.9  4.4–6.9 mm ∅; Se 2.5–3 mm. Easily distinguished on account of the caespitose growth and the tall inflorescences, as well as by the long and broad floral bracts that cover the flowers. — The type number was originally erroneously given as “Müller 803”. H. carlsoniae Burt-Utley & Utley (Syst. Bot. 13(2): 276–279, ills., 1988). Type: Mexico, Guerrero (Burt-Utley & Utley 7492 [MEXU, BH, CAS, F, MICH, MO, NY, US]). — Lit: PulidoEsparza & al. (2004). Distr: S Mexico (Guerrero: Taxco, Iguala, Buenavista de Cuéllar); tropical deciduous forest, crevices or cliffs, 800–1050 m. Caespitose, flowering to 1.4 m tall; L 10–15 per rosette, frequently falcate, sheaths broadly ovate or elliptic to reniform, 2.3–4.8  1.9–6.4 cm, margin finely spiny, glabrous and lustrous on both faces, becoming densely lepidote for their length, L lamina 17–62  1–2.1 cm, appressed cinereous-lepidote on both faces, Sp antrorse,

999

red-brown, thin, 1.5–2.5 mm, 6–10 mm apart; male Inf 1 branched, with 12–13 primary Br 7.5–20 cm long; floral Bra narrowly ovate, ovateelliptic or ovate-triangular, 2–4.5  0.8–2 mm, attenuate-acuminate, entire to distally erose; Fl divaricate, sometimes appearing verticillate; Ped 0.2–1.8 mm; Sep white-greenish with apical purple little dots, ovate to ovate-triangular, 2.5–3.3  1.5–2 mm, glabrous, acute, margin distally erose; Pet white with apical purple little dots, elliptic to ovate-elliptic, 4.2–5  2–3 mm, rounded to acute, margin distally erose; St longer than the petals; Fil 4.5–5.5 mm; Anth green, 1–1.6 mm; female Inf simple or 1 branched, with 2–7 primary Br 4–17.5 cm long; floral Bra narrowly triangular to oblong, 2–8  0.8–3 mm, attenuate-acuminate to apiculate, margin erose, shorter or longer than the pedicels; Ped 2.5–4 mm; Sep green, purple at the apex, narrowly ovate-triangular, 2.5–4  2–2.5 mm, acuminate or rarely apiculate, margin distally erose, glabrous; Pet white, ovate-triangular, 4.2–7.5  2.5–3.5 mm, acuminate; Ov purple, ovoid to ellipsoid, 7–8 mm  4 mm ∅; Fr nutant, 9.5–16.5  4.5–8 mm, triquetrous, glabrous; Se dark brown, 7–8  1.5 mm. H. caudata L. B. Smith (Phytologia 8: 5, t. 1, figs. 1–5, 1961). Type: Mexico, Oaxaca (Foster & Van Hyning 2999 [US, US]). — Distr: Mexico (Oaxaca: Santiago Lachiguiri, Tehuantepec); xerophytic shrubland, saxicolous, 300–400 m; known only from the type material. Growth habit unknown, flowering to 1.3 m tall; L 25, sheaths brown to dark brown, lustrous, suborbicular, 4–7  3–8 cm, slightly serrulate, lepidote distally, L lamina reddish adaxially, 50–80  3–5.5 cm, glabrous adaxially, appressed cinereous-lepidote abaxially, Sp antrorse, 2–4 mm, 5–15 mm apart; Inf seemingly lateral, 1 branched, glabrous; peduncular Bra ovate with long linear, caudate, subentire, lepidote lamina, the upper much reduced and without lamina; lower primary Bra like the peduncular bracts, caudate; male Inf with 30 primary Br 3.5–8 cm long, slightly compressed at the base; floral Bra stramineous, ovate, 2.6–4 mm, acuminate, as long as or slightly longer than the pedicels; Fl divaricate to ascending; Ped slender, 1–1.3 mm; Sep white, elliptic to

1000

ovate, 2.5–3  1.7 mm, obtuse; Pet white, elliptic to ovate, 4.3–4.6  2.3–2.5 mm, obtuse; St longer than the petals; Fil 3.9 mm; Anth green, 1.6 mm; female Inf with 40 primary Br 4–10 cm long; floral Bra stramineous, ovate, 2 mm, acuminate; Fl divaricate, laxly disposed; Sep triangular, 1.3–1.4  1.3–1.4 mm, acute; Pet white, narrowly triangular, 4.5–4.7  1.9–2.3 mm, acuminate; Ov ovoid; Fr light brown, 6.4–11  4 mm ∅; Se light brown, 3–4 mm. H. caulescens López-Ferrari & al. (Novon 19 (2): 197–200, ills., 2009). Type: Mexico, Puebla (Martínez & al. 58 [UAMIZ, IEB, MEXU]). — Distr: C Mexico (Morelos, Puebla, Oaxaca); arid scrub and tropical deciduous forests, on the ground or on rocks, 1200–1600 m. Caespitose, flowering 0.8–1.9 m tall; stems cylindrical, long rhizomatous, prostrate or decumbent, 3–3.7 cm ∅; Ros regular, 40 cm ∅; L numerous, sheaths light brown, depressed ovate, 3.2–4.7  4.9–6 cm, lustrous and white-lepidote at the apex on both faces, L lamina green to dark green, 29–40  1.8–3 cm, densely white-lepidote on both faces, Sp brown, curved, antrorse or some retrorse, 1.9–4 mm, 1.5–1.9 cm apart; Inf 2 branched; peduncle 4–12 mm ∅; peduncular Bra foliaceous, entire, glabrous, becoming gradually shorter upwards; male Inf with 18–20 ascending to erect primary Br 10–22 cm long; secondary Br ascending, 4.5–7 cm long; floral Bra light brown, triangular, 2.2–2.5  1.3–1.7 mm, entire, acuminate; Fl ascending to divaricate, sessile or subsessile, densely disposed; Sep light brown to brown basally, ovate-triangular, 2.1–2.2  1.6–1.8 mm; Pet white, elliptic, 3.5–4  2–2.5 mm; St subequal, longer than the petals; Fil 2.8–3 mm; Anth white to whitish-yellow, 1 mm; female Inf with 20–30 ascending primary Br 0.6–2.3 cm long; secondary Br 2.5–9 cm long; floral Bra light brown, triangular, 2  1.8 mm, slightly erose, acuminate; Fl ascending to appressed; Ped 1.3–1.7 mm; Sep greenishbrown, triangular, 2.7–3.7  1.3–1.7 mm; Pet white, narrowly triangular, 3.4–3.8  1.1–1.4 mm; Ov green, ovoid to long ovoid, 3.3–3.9  1.2–1.5 mm, glabrous; Fr 8.5–10  3.2–4.3 mm; Se 2.8–4 mm.

A. Espejo-Serna et al.

H. chichinautzensis Martínez-Correa & al. (Syst. Bot. 35(4): 746, ills. (pp. 747–748), 2010). Type: Mexico, Morelos (Martínez-Correa & al. 37 [UAMIZ, CICY, IEB, MEXU]). — Distr: C Mexico (Morelos: Cuernavaca, Tepoztlán), rosetterich scrub on igneous rocks of lava flows, on rocks, 2000–2500 m. Caespitose, flowering 0.6–1.7 m tall; L numerous, sheaths yellow to brown, ovate to square, glabrous basally and sparsely lepidote at the apex on both faces, 2.5–4.3  2–6.4 cm, L lamina green, 10–36  1–4 cm, glabrous adaxially, sparsely lepidote abaxially, Sp divaricate to antrorse, 2–6.3 mm, 0.6–1.4 cm apart; Inf lepidote, 2 branched; peduncle 4.5–15 mm ∅; peduncular Bra foliaceous, lepidote, serrate, 4–12.3 cm, gradually becoming shorter upwards; male Inf with up to 57 ascending primary Br 0.6–5.4 cm long; Fl fragrant, laxly arranged, 3.2–4.7  2–3 mm ∅; Ped 1.6–2.4 mm; floral Bra brown, triangular, entire, glabrous, 1.5–3.5  0.4–0.7 mm; Sep brown, ovate, 2–3.1  1.3–1.7 mm, entire, acute; Pet green with a longitudinal brown line abaxially, oblong, 3–4.4  1.8–2.4 mm, rounded; St equal, longer than the petals; Fil 3.2–4.2 mm; Anth brown to black, 0.8–1.7 mm; female Inf with up to 35 ascending primary Br 2.2–5.5 cm long; floral Bra brown, triangular, entire, glabrous, 1.5–2.4  0.7–1.2 mm; Fl fragrant, laxly arranged, 3.8–6.8  2–3.5 mm ∅; Ped 1.5–4.4 mm; Sep brown, triangular, 1.6–2.8  0.9–1.8 mm, entire, acute; Pet green with a brown longitudinal line abaxially, triangular, 2.2–4.4  1.3–2 mm, entire, acute; Ov ovoid, 4–7  2–2.4 mm ∅; Fr 10–14  3–7 mm; Se 2.4–5.5 mm. H. colossa Martínez-Correa & al. (Syst. Bot. 35(4): 746–747, ills. (pp. 752–753), 2010). Type: Mexico, Oaxaca (Espejo & al. 6872 [UAMIZ]). — Distr: S Mexico (Puebla, Oaxaca); tropical deciduous forest, limestone slopes, 1400–1900 m. Caespitose, flowering 1.9–2 m tall; L numerous, sheaths yellow adaxially, brown abaxially, widely ovate to square, 7.5  9.7–11.3 cm, glabrous basally and lepidote at the apex on both faces, serrulate, L lamina light green, 45–55  5–6.3 cm, glabrous adaxially, sparsely lepidote abaxially, Sp divaricate to antrorse, 4.3–5.5 mm,

Hechtia BROMELIACEAE

1.3–2 cm apart; Inf 2 branched; peduncle 20 mm ∅; peduncular Bra foliaceous, 14 cm, lepidote, spiny, the apical ones narrowly triangular, lepidote, entire; male Inf 1.25–1.94 m, with up to 60 ascending primary Br 9.8–12 cm long; floral Bra brown, triangular, 4.9–5.7  0.9 mm, entire, glabrous, attenuate; Fl divaricate, densely disposed; Ped 2.1–3 mm, glabrous; Sep brown, triangular, 3.2–3.8  1.7–2.1 mm, entire, acute; Pet green, elliptic to oblong, 5.8–6.2  2.6 mm, rounded, glabrous, entire; St equal, longer than the petals; Fil 4.5–4.6 mm; Anth yellow, 1.1–1.2 mm; female Inf with 47 ascending primary Br 17.5 cm long, secondary Br ascending, 7.5–9.5 cm; floral Bra brown, triangular, 4.8  1.7 mm, entire, glabrous; Fl divaricate to ascending, densely disposed; Ped 5 mm, glabrous; Sep brown, ovate, 2.5  1.5 mm; Pet green, triangular, 4.8  2.1 mm, glabrous, entire; Fr 15  5.3 mm ∅; Se 5.4 mm. H. complanata Burt-Utley (Phytoneuron 2012 (69): 6–10, ills., 2012). Type: Mexico, Oaxaca (Utley & Utley 8823 [MEXU, BM, CAS, GH, MICH, MO, US, USF]). — Distr: S Mexico (Oaxaca); thorn scrub and tropical deciduous forest, on the ground or on rocks on talus slopes. Solitary, flowering to 2.3 m tall; Ros 1–1.2 m ∅, forming long rhizomes to 30 cm, 4 cm ∅; L numerous, reflexed in the apical portion, sometimes the tips strongly curled, sheaths stramineous to castaneous, semiorbicular to transversely elliptic, 6–7.5  9.5–16 cm, margins finely spinulose, floccose at the apex, glabrous and lustrous on both faces, becoming densely lepidote distally, L lamina green, occasionally pink-tinged, 45–75  5–6 cm, very densely white-lepidote on both faces, most conspicuous on the lower , glabrescent adaxially at the apical portion, Sp castaneous to dark brown, antrorse and retrorse, or rarely divaricate, 3.5–7.5 mm, 2.7–5.3 cm apart; Inf finely lepidote to glabrous; peduncle 12–30 mm ∅; peduncular Bra rose-greenish, linear to narrowly triangular, 31–33 cm, becoming progressively reduced distally and exceeding the internodes, pungent; male Inf 2 branched, with 1–14 divaricate to ascending primary Br (8–) 11.5–32 cm long; floral Bra shorter than to

1001

equalling the sepals, ovate to narrowly triangular, 0.9–2  0.3–0.8 mm, finely spinulose-serrulate, attenuate-acuminate, lepidote, acute; Fl divaricate, sometimes appearing verticillate; Ped articulated, stout, conical, 0.3–1.5 mm, lepidote; Sep ovate to deltoid, 1.2–1.6  1–1.6 mm, apiculate, occasionally finely serrulate-spinulose or erose, lepidote; Pet pale yellow, ovate to elliptic, 2.5–4.5  1.3–2.2 mm, apiculate, glabrous; female Inf 1 or infrequently 2 branched, with up to 34 divaricate to ascending primary Br (8.5–) 13–23.5 cm long; floral Bra triangular, 1–1.5  0.4–0.7 mm, apiculate, finely serrulate-spinulose, lepidote to occasionally glabrous; Fl divaricate, laxly disposed, sometimes appearing verticillate; Ped articulated, stout, conical, 0.5–1 mm, lepidote to occasionally glabrous; Sep triangular, 0.8–1.2  0.8–1.2 mm, entire to irregularly finely serrulate, lepidote to occasionally glabrous; Pet green, triangular, 2.1–2.9  1–1.8 mm, entire, apiculate to acute, glabrous; Ov lepidote; Fr dark brown but drying olive-green, 7.5–8.5  3–4 mm ∅, finely lepidote; Se not known. H. confusa L. B. Smith (Contr. Gray Herb. 117: 22, t. 1, figs. 71–72, 1937). Type: Mexico, Puebla (Pringle 7479 [GH, VT]). — Lit: Smith (1938); Espejo-Serna & al. (2007); Burt-Utley & Utley (2011). Distr: C Mexico (Puebla, Oaxaca); rosette-rich scrub and thorn scrub, limestone hills and slopes, and on calcareous bluffs on rocks, 1900–2600 m. – Figs. 1 and 2. Caespitose, flowering to 1.3 m tall; Ros 25  20 cm ∅; L numerous, sheaths yellow to brownish, ovate, 2–6 (7–)  1.3–6 ( 9) cm, serrate, glabrous basally, lepidote on both faces, L lamina dark green, recurved, 9–35 ( 50)  0.35–0.9 cm, pungent, glabrous adaxially, densely cinereous-lepidote abaxially, Sp retrorse or divaricate, some antrorse, 1.5–8 mm, 0.5–1.7 cm apart; Inf 2 branched; basal peduncular Bra foliaceous, caudate, sparsely lepidote, serrate, 3.7–12.8 cm, the apical ones triangular, erose; male Inf with up 38 primary Br; floral Bra brownish, widely ovate, 3.8–7.3  1.7–4.5 ( 6) mm, erose, lepidote, acute; Fl sessile, 3–6.9  2.1–3.1 mm ∅; Sep brownish, ovate, 2.4–4.6  1.4–3.2 mm, entire, rounded; Pet green-yellowish, oblong to

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A. Espejo-Serna et al.

Fig. 1 Hechtia confusa (male flowers). (Copyright: A. Espejo-Serna)

6.6 ( 9.5)  2.4 mm ∅; Sep brownish, ovate, 2.7–5.1  1.2–3.1 mm, acute; Pet greenyellowish, triangular 3.5–5.1  1.2–3 mm, entire, rounded; Ov ovoid, 6–7  2.1–2.5 mm ∅; Fr brownish, 6.5–10  3–8 mm ∅; Se 2.2–3.6 mm.

Fig. 2 Hechtia confusa (female flowers). (Copyright: A. Espejo-Serna)

elliptic, 2.5–5.7  1.6–4 mm, entire, rounded; St equal, longer than the petals, triangular, flattened; Fil 2.4–4.4 mm; Anth yellow, 0.8–2.2 ( 2.8) mm; female Inf with up 38 primary Br; floral Bra brownish, widely ovate, 4.3–7.9  1.5–4.8 mm, erose, lepidote, acute; Fl sessile, densely disposed,

H. conzattiana L. B. Smith (Contr. Gray Herb. 117: 19, t. 1, fig. 56, 1937). Type: Mexico, Oaxaca (Conzatti & Gómez 3501 [US, MEXU]). — Distr: S Mexico (Oaxaca, Puebla); oak forest, 1300 m. Flowering to 1 m tall; L numerous, sheaths brown, ovate, 4 cm long, L lamina 15–40  1–1.3 cm, sparsely pale-lepidote adaxially, soon becoming glabrous, prominently nerved and lepidote abaxially, largely acuminate; Sp divaricate to barely antrorse, 2 mm, 0.8–1 cm apart; Inf glabrous; peduncular Bra red with prominent hyaline margins, linear from a triangular-ovate base, lepidote, becoming progressively reduced distally; male Inf 1 branched with 35 primary divaricate Br 4–11 cm long; floral Bra brownish, elliptic, 2–2.5 mm, erose, longer than the pedicels; Fl divaricate, densely disposed; Ped 1.5 mm; Sep pale brown, blackish punctate, broadly elliptic, 2 mm, entire to faintly erose, obtuse; Pet white, elliptic, 2–2.3  2.5–4 mm, obtuse; St longer than the petals; Fil 3–3.2 mm; Anth yellow, 1.5 mm; female Inf, Fl and Fr unknown. H. deceptrix I. Ramírez & Hornung (Phytotaxa 221(2): 158, Figs. 1, 2, 3a, 3e, 2015).

Hechtia BROMELIACEAE

Type: Mexico, Hidalgo (López-Ferrari & al. 3309 [UAMIZ, IEB]). — Distr: Mexico (Hidalgo: Actopan, Atotonilco El Grande, Cardonal); xerophilous scrub on calcareous cliffs, 1700–1800 m. Caespitose, flowering to 2.5 m tall; Ros 50–60 cm ∅; L 50–80, sheaths light brown, broadly ovate, 4–6  4–5 cm, margins erose, lustrous and glabrous adaxially, densely whitelepidote abaxially, L lamina green, narrowly triangular, 26–50  1–3.5 cm, densely white-lepidote abaxially, glabrous adaxially, Sp light green or occasionally purple, antrorse, 1.5–2.5 mm, 0.4–1 cm apart; male Inf 1 branched, with 30 Br, occasionally the lower ones with a pair of secondary basal branches; peduncular Bra foliaceous, sheaths light brown, triangular, 2.5–4  3–4 cm, margins entire to erose, lamina green, narrowly triangular, 4–6  0.6–1.2 cm, margins spinose, glabrous adaxially, densely white-lepidote abaxially; Br ascending, 4–17 cm; floral Bra green with light brown apex, ovate-elliptic, 6–7  2.6–3 mm, margins erose, glabrous; Fl polystichous; Ped 4–5 mm; Sep green with light brown apex, triangular, 3.5–3.8  1.8–2 mm, entire, glabrous; Pet light green, widely elliptic, 4.2–4.5  3.3–3.5 mm, rounded; St unequal; Fil 1.9–3.5 mm; Anth green, 0.9–1.5 mm; female Inf 1 branched, with 7–16 Br, occasionally the lower ones with a pair of secondary basal branches; peduncular Bra foliaceous, sheaths green, triangular, 2.5–4  3–4 cm, margins entire to erose, lamina green, narrowly triangular, 7–9  0.6–1 cm, margins spinose, glabrous adaxially, densely white-lepidote abaxially; Br ascending, 5–14 cm; floral Bra green, ovate-elliptic, 8  4–5 mm, margins erose, glabrous; Fl polystichous; Ped 1 m  7 cm, glabrous above, appressedly cinereous-lepidote abaxially, Sp antrorse, 6 mm; Inf seemingly lateral, 1 branched; peduncular Bra broadly triangular, exposing almost the whole peduncle but the linear mostly entire laminae equalling or exceeding the internodes; male Inf with 15 slender primary Br 6.5–25 cm long; floral Bra narrowly triangular to ovate, 4.2–4.4  1.7 mm, acuminate, longer than the pedicels; Fl ascending, some appearing verticillate; Ped slender, 1–1.3 mm; Sep white, ovate-triangular, 2.3–2.4  1.7–2 mm, glabrous, acute; Pet white tinged with red at the base adaxially, ovateelliptic, 4.8–5  3.2–3.3 mm, glabrous, acute; St shorter than the petals; Fil 3.5–3.6 mm; Anth yellow, 1.6 mm; female Inf 1 branched; floral Bra broadly ovate, 4 mm, acute, much exceeding the stout obconic pedicels; Fl divaricate; Sep

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hyaline with a thickened brown base, ovateoblong, 3 mm, obtuse; Pet white, 4 mm; Fr brown, 8 mm, acute, sharply trigonous, smooth, lustrous. H. fragilis Burt-Utley & Utley (Brittonia 39 (1): 40–42, ills., 1987). Type: Mexico, Oaxaca (Burt-Utley & Utley 6980 [MEXU, MO, US]). — Distr: Mexico (Oaxaca, Puebla); tropical deciduous forest and thorn scrub, on the ground or on limestone cliffs, 600–1200 ( 1800?) m. Caespitose in small groups, flowering 40–100 cm tall; L 20–25, brittle, sheaths yellow to light brown, transversely oblong, 1.6–4  3.8–5.6 cm, margin finely spiny, L lamina light green with red spots, 10–21  1–3 cm, glabrous adaxially, appressedly cinereous-lepidote abaxially, Sp retrorse to divaricate, green to red-brown, 1.3–4 mm; Inf 1 branched, sparsely tomentose; male Inf with 40 primary Br; floral Bra ovatetriangular, 2–3.1  0.7–1 mm, sparsely tomentose-lepidote, acuminate, occasionally apiculate, finely erose; Ped 1.5–2.3 mm; Sep green tinged purple, ovate to ovate-elliptic, 1.3–2.5  1.9–2.1 mm, glabrous, acute to obtuse; Pet greenish-white tinged purple, broadly elliptic to oblong, 4.5–5  2.9–3.1 mm, glabrous; St as long as the petals; Fil 3–5 mm; Anth green, 1.5–2 mm; female Inf with 20 primary Br; floral Bra narrowly ovate to ovate-triangular, 1.6–4.5  1–2 mm, acuminate, margin distally erose, sparingly tomentose to lepidote; Ped 1.5–2.5 mm; Sep brownish-green, ovate-triangular to triangular, 2–2.9  1.3–1.6 mm, glabrous to sparingly tomentose, acute, margin hyaline, entire, sometimes distally erose; Pet green tinged purple, ovate to ovate-elliptic, 3.6–4.7  2.3–2.5 mm, glabrous, acuminate; Ov wholly superior, green, purplish at the apex, ovoid, 4.5–5.1  3.4–3.6 mm; Fr dark brown, 5.5–8  3.5–5 mm; Se brown, 5.5–6 mm. H. galeottii Mez (Repert. Spec. Nov. Regni Veg. 16: 71–72, 1919). Type [lecto]: Mexico, Oaxaca (Pringle 6703 [B, B, BR, ENCB, G, GH, HBG, M, MEXU, P, UC, VT, WU, Z]). — Lit: Espejo-Serna & al. (2013). Distr: S Mexico (Oaxaca: Asunción Nochixtlán, San Jerónimo Sosola); oak forest and transitions to tropical

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deciduous forests, terrestrial on calcareous hills, 1500–2000 m. Solitary; flowering to 2 m tall; L numerous, sheaths light brown, oblong to suborbicular, 5.2–6  5–5.5 cm, glabrous and lustrous basally, becoming densely white-lepidote distally on both faces, L lamina green, 30–40  3.5–4.5 cm, glabrous adaxially, finely appressedly white-lepidote abaxially, Sp stout, dark to light brown or green, curved, antrorse, with a conspicuous triangular base, 4 mm, 2–4 cm apart; Inf very sparsely floccoselepidote, glabrescent; peduncle dark brown, cylindrical, 2 cm ∅ basally; peduncular Bra light brown, linear-triangular to triangular, pungent; male Inf 2 branched, with numerous divaricate to ascending primary Br 16–35 cm long, with 2 short (2.5–13 cm) secondary branches at the base; floral Bra light brown, ovate-triangular, 1 mm, sparsely lepidote, acute, erose, shorter than the sepals; Fl ascending, sometimes appearing verticillate, subsessile or very shortly pedicellate; Ped stout, conical, 1.5 mm; Sep broadly ovate, 1.5  1.4–1.6 mm, obtuse; Pet densely black-punctate, elliptic to ovate-elliptic, 2.5–2.8  1.9–2 mm, rounded; St longer than the petals; Fil 2.5–2.8 mm; Anth greenish, 1 mm; female Inf 1 branched, with numerous divaricate to ascending primary Br 12.5–28 cm long; floral Bra light brown, dark brown towards the base, ovate-triangular, 1.5 mm, lepidote, erose, shorter than the sepals; Fl ascending, densely to laxly disposed, sometimes appearing verticillate, subsessile; Sep ovate to triangular, 1.8–2  1.4–1.5 mm, acute; Pet narrowly triangular to deltoid, 3.3–3.4  1.4–1.5 mm, acute; Ov ellipsoid, 2.5  1 mm, glabrous; Fr dark brown, 7.5–8 mm, acute; Se reddish-brown, 3.5 mm. See Espejo-Serna & al. (2013) for details of the typification and the type locality. H. glauca Burt-Utley & Utley (Brittonia 45: 220–222, fig. 1, 1993). Type: Mexico, Michoacán (Burt-Utley & Utley 8387 [MEXU, B, CAS, DUKE, IEB, MICH, MO, NY, US]). — Distr: Mexico (Michoacán: Arteaga); on exposed rocky cliffs, 300–900 m.

A. Espejo-Serna et al.

Usually solitary or in groups of few rosettes, flowering 0.9–1.5 m tall; Ros strongly spreading, to 80 cm ∅; L frequently falcate, sheaths stramineous to light brown, often deep pink-red distally, transversely elliptic to reniform, 4.5–6.7  6.5–11 cm, margins crenulate to weakly spinulose, glabrous and lustrous on both faces, L lamina blue-green, occasionally suffused with pink, wax-covered, strongly recurved distally, 33.5–70  5.8–9.3 cm, glabrous on both faces, Sp red-brown, antrorse, 0.8–3 mm, 0.7–3 cm apart; Inf glabrous; peduncular Bra foliaceous, becoming progressively reduced distally; male Inf weakly 2 branched; floral Bra broadly ovate, 1.2–2  1.5–1.7 mm, apiculate to attenuate-acuminate, very finely denticulate to erose, longer than the pedicels; Ped stout, 0.3–1.2 mm, articulated; Sep light green or green suffused with pink, ovate to broadly ovate, 1.6–2.2  1.5–2.3 mm, glabrous, rounded to acute or acuminate, entire or rarely somewhat erose; Pet white to light lavender-pink, subspatulate, 4.5–5  1.8–2.2 mm, rounded; female Inf 1 or rarely 2 branched; floral Bra ovate, 1–2.3  1–1.3 mm, acuminate to apiculate, erose distally, shorter than the pedicels; Ped 2–4.5 mm, articulated; Sep light green or green suffused with pink, triangular to ovate-triangular, 1.7–2.5  1.4–2 mm, apiculate to acute, entire or somewhat erose; Pet white to faintly or deeply suffused with dark lavender-pink, triangular, 3.5–5  1.3–2.1 mm, acuminate to apiculate; Ov largely superior, light green to light green suffused with pink; Fr brown, (8–) 9–12  3.5–5 mm, glabrous, smooth. H. glomerata Zuccarini (Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 3: 240–242, t. 6, 1840). Type: Mexico, Hidalgo? (Karwinsky s.n. [[lecto — icono]: l.c., t. 6]). — Lit: BurtUtley & Utley (2011). Distr: Mexico (Nuevo León, Tamaulipas, San Luis Potosí, Aguascalientes, Hidalgo, Querétaro, Guanjuato, México, Chiapas), Guatemala (Guatemala, Huehuetenango, Quiché); thorn scrub vegetation and tropical deciduous forest, generally on limestone hills or cliffs, on the ground or on rocks, 200–2100 m. Incl. Dasylirion pitcairniifolium Karwinsky & Zuccarini ex Zuccarini (1838) (nom. inval., ICN

Hechtia BROMELIACEAE

Art. 38.3); incl. Yucca pitcairniifolia Karwinsky ex Zuccarini (1838) (nom. inval., ICN Art. 38.3); incl. Hechtia ghiesbreghtiana Lemaire (1862) (nom. inval., ICN Art. 38.3); incl. Hechtia ghiesbreghtii Lemaire (1863); incl. Hechtia argentea K. Koch ex Baker (1884)  Dyckia argentea (K. Koch ex Baker) G. Nicholson (1886); incl. Hechtia capituligera Mez (1896); incl. Hechtia gamopetala Mez (1896); incl. Hechtia morreniana Mez (1896); incl. Hechtia lepidophylla I. Ramírez (2008). Solitary or caespitose, flowering 1.4–2.5 m tall; Ros globose, 20–90 cm ∅; L numerous, 50–100 per rosette, sheaths white to yellowish with a dark brown to black spot, oblong to depressed-ovate, 3–6  3–8 cm, glabrous, lustrous, entire or sometimes minutely denticulate towards the apical portion, L lamina green to light or greyish, 20–85  1–3.5 cm, densely cinerous to white-lepidote or glabrous and lustrous adaxially, densely white-lepidote abaxially, Sp light brown to brown or reddish, antrorse to divaricate, 4–7 mm, 0.7–2 cm apart; Inf seemingly lateral, 1–2 branched, rarely simple, white-lepidote to white-tomentulose, soon glabrous; peduncle cylindrical, flattened at the base, 1–3 cm ∅, white-lepidote to glabrescent; peduncular Bra brown to light brown, triangular to narrowly triangular, 3–5.5  0.9–1.5 cm, entire to minutely denticulate, white-lepidote to glabrescent on both faces, acute to acuminate, the basal ones imbricate and longer than the internodes, the upper shorter than the internodes; male Inf with 13–35 divaricate to ascending, glomerate to elongated primary Br 1–15 ( 23) cm long; primary Bra brown to light brown, triangular, 8–12  3–4 mm, acute to acuminate, margins spiny, becoming progressively reduced distally; secondary Br, when present, glomerate, 0.6–2.2 cm; secondary Bra ovate-triangular, 7–8  3–4 mm, apiculate; floral Bra brownish, widely triangular-ovate, 4–7  5–7 mm, fimbriate, lepidote abaxially, acute to apiculate; Fl ascending, sessile or shortly pedicellate; Sep white to rose-coloured, ovate, widely elliptic to oblong, 3.5–4.5  3 mm, densely whitelepidote, entire, rounded; Pet white, obovate to ovate or elliptic, 4.5–6  3–3.5 mm, rounded; St

1007

unequal, shorter than the petals; Fil 2.5–4 mm; Anth green, 1–1.5 mm; female Inf with 15–39 divaricate to ascending, glomerate to elongated primary Br 1–15 cm long; primary Bra brown to light brown, triangular, becoming progressively reduced distally; secondary Br, when present, glomerate, 1.5 cm; secondary Bra ovatetriangular, 7–8  3–4 mm, apiculate; floral Bra brownish, triangular-ovate to triangular, 4.5–6.5  3–5 mm, brownish, densely white-lepidote, fimbriate, acute to apiculate, longer than the sepals; Fl ascending, sessile to shortly pedicellate; Ped stout, 1 mm; Sep greenish or brownish to reddish, ovate to triangular-ovate, 4.2–5.8  3–4 mm, sparsely to densely white-lepidote, margin hyaline, entire to erose, rounded to acute; Pet white to rose-coloured, ovate to oblong, 4.5–7  3–4 mm, rounded to apiculate; Ov green, ellipsoid to oblong, 4.5–7.5  3–3.5 mm, densely whitelepidote to light brown-floccose-lepidote; Fr light to dark brown, 6.5–10  4–5 mm, densely whitelepidote to glabrescent; Se brown to reddish, 4.5–5 mm. Smith & Downs (1974) cite a Karwinski specimen in the Munich herbarium as type, but this specimen was only prepared 1853 from cultivated descendants of the original collection, i.e. 13 years after the publication of the name. According to the nomenclatural rules, this material cannot be the type. Therefore, the detailed figure (plate 6) in the protologue of H. glomerata is here designated as lectotype of the name. H. glomerata is a highly variable species with a wide distribution in Mexico and Guatemala. The populations show variation in some characters, particularly in the size of the plants and inflorescences, in the pubescence of the leaves, and in the form of the floral structures. H. argentea (accepted as separate species by Siekkinen (2015)), H. ghiesbreghtii and H. lepidophylla are here listed as synonyms since we consider them to represent only forms of this variation. More detailed studies at the level of populations are needed for a better understanding of the H. glomerata complex, however. H. guatemalensis Mez (Repert. Spec. Nov. Regni Veg. 3: 14, 1906). Type: Guatemala,

1008

Guatemala (Pittier 137 [US, B]). — Distr: Guatemala, Honduras, El Salvador, Nicaragua, Belize; tropical deciduous forest, thorn scrub and pine-oak forest, on the ground or on rocks, 200–1500 m. Caespitose, flowering to 2 m tall; Ros 60–100 cm ∅; L numerous, sheaths light brown, widely ovate to suborbicular, 3.5–4  4.5–6 cm, glabrous, lustrous, margins weakly spinulose, L lamina 30–80  2–5.3 cm, lepidote to glabrous adaxially, densely white-lepidote abaxially, long caudate, Sp red-brown, antrorse, 3–4.5 mm, 0.7–1.7 cm apart; Inf glabrous; peduncle pale brown, cylindrical, 1–2 cm ∅, basally slender, glabrous; peduncular Bra light brown, linear to linear-triangular, margins spiny; male Inf 2 branched, laxly flowered; primary Bra light brown, triangular, 1.7–3 cm  4–7 mm, margins minutely spiny; primary Br slender, 23–30 cm; secondary Bra narrowly triangular, 5–7 mm; secondary Br 6–16 cm; floral Bra narrowly ovate, 1.3–1.5 mm long, scarious, acuminate, equalling the pedicels; Fl polystichous, reflexed; Ped slender, 1.5 mm; Sep triangular, 2–2.2  2 mm, obtuse; Pet white, elliptic, 4–4.5  2.4–2.7 mm, obtuse; St longer than the petals; Fil 4.6–4.9 mm; Anth green, 2.3–2.6 mm; female Inf generally 2 branched, rarely 1 or 3 branched, laxly flowered; primary Bra light brown, triangular, 0.8–1.4 cm  4–7 mm, entire; primary Br slender, 23–41 cm; secondary Bra narrowly triangular, 3–4 mm; secondary Br 7.5–15 cm; floral Bra triangular, 1.5 mm, acuminate; Fl polystichous, reflexed; Ped 1 mm; Sep white, triangular, 1.8–3  1.8–2 mm, acute; Pet white, elliptic, 3.5–3.7  2 mm, acuminate; Ov almost wholly inferior, green, ellipsoid, 3–3.3 mm  2.5–2.6 mm, glabrous; Sty 1 mm; Fr light brown, 5–8 mm, glabrous, prominently and irregularly veined; Se red-brown. H. hernandez-sandovalii I. Ramírez & al. (Phytotaxa 112(2): 34–38, ills., 2013). Type: Mexico, Tamaulipas (Hernández & Martínez 1797 [MEXU, QMEX, TEX-LL, UAT]). — Distr: Mexico (SW Tamaulipas: Miquihuana); Agave-Hechtia rosette-rich shrubland, on limestone hills or rocky exposed areas, 1550–2200 m.

A. Espejo-Serna et al.

Caespitose, flowering 1–1.8 m tall; Ros to 35 cm ∅; L 25–35, sheaths white-yellowish adaxially, brownish abaxially, sometimes with a darker brown area on the apical zone, transversely oblong, 2.5–3  (2.6–) 3.7–4.8 cm, glabrous on both faces, finely dentate at the apex, L lamina green, 21–39  1.3–2.9 cm, acuminate, glabrous to sparsely white-lepidote adaxially, white-lepidote abaxially, Sp reddish, generally retrorse, 4–7 mm, (0.8–) 1.6–4.4 ( 5.1) cm apart; Inf seemingly lateral; male Inf 1 branched, sparsely white-lepidote but sometimes glabrous, with 29 primary capitate Br; peduncular Bra brownish, triangular to triangular-ovate, 2.3–6.2  1–2.7 cm, acute and acuminate, sparsely lepidote, finely and laxly denticulate at the base, entire towards the apex; primary Bra brownish, triangular, 1–4  0.6–1.2 cm, acute, sparsely lepidote, entire; Fl densely clustered, covered by a thick brownish cotton-like indumentum, 6.5–7.5  3–3.2 mm, sessile; floral Bra brownish, long-ovate, 6.3–8.2  4–4.4 mm, acute, erose, densely white lepidote, longer than the petals; Sep connate for their length, brownish, oblong, 4.4–5.3  2.3–2.5 mm, acute, erose, densely white-lepidote; Pet white, obovate, 5.5–6  2.8–3 mm, rounded, entire, densely white-lepidote; St adnate to the petals and pistillode base, barely longer than the petals; Fil 2.9–3.5 mm; Anth 2.2 mm; female Inf 1 branched, sparsely white-lepidote, with 11–13 capitate primary Br; peduncular Bra brownish, 1.7–6.3  0.7–2.6 cm, acute, sparsely lepidote, finely and laxly denticulate at the base, entire towards the apex; Fl densely clustered, covered with a thick, white, cotton-like indumentum, 7–7.5  4–4.6 mm, sessile; floral Bra brownish, transversely oblong to broadly elliptic, 8.7–9.4  5.8–6 mm, acute, margin erose, densely white-lepidote; Sep connate for of their length, brownish, carinate, ovate, 6.2–6.5  4–4.2 mm, acute, margin erose, densely white-lepidote; Pet free, white, ovate to long ovate, 4–4.9  3.8–4.4 mm, acute, entire, densely white-lepidote; Ov white to greenish, ovoid, 6  3 mm; Fr brown, 7.2–8  4–5.2 mm; Se brownish, 3.3–3.9  1.4 mm. H. hintoniana Burt-Utley & al. (Phytoneuron 2011(59): 2, 2011). Type: Mexico, México

Hechtia BROMELIACEAE

(Matuda & al. 31124 [MEXU, CAS, MEXU, MO, NY, US]). — Distr: Mexico (México, Michoacán); tropical deciduous forest, 750–900 m. Habit unknown, flowering to 2 m tall; L presumably numerous, sheaths narrow, 3.5–5.5 cm wide, marginally spiny and floccose, distally lepidote on both faces, L lamina straight to somewhat falcate, 36.5–93  0.6–2.2 ( 3.5) cm, lepidote on both faces, Sp red-brown to pale yellow, generally antrorse, 1.8–2.5 mm, 0.7–1.5 cm apart; Inf presumably terminal; peduncle 1.2–1.4 cm ∅, glabrous to glabrescent; upper peduncular Bra chartaceous, shorter than the internodes, rarely equal to or exceeding them, lanceolate-ovate with appressed linear-triangular lamina 1.5–3.9 cm long; male Inf lepidote, 2 branched, with numerous primary Br 8–16 cm long and (2–) 3–6 secondary Br; primary Bra ascending to subascending, (4.5–) 7.8–22 cm; floral Bra shorter than to exceeding the pedicels, chartaceous, ovatetriangular to oblong or suborbicular, 0.7–1.5  0.4–1 mm, externally lepidote, marginally finely serrulate to erose, spinulose or apiculate to praemorse; Fl divaricate to weakly ascending, subsessile to shortly pedicellate; Ped weakly articulated, stout, lepidote; Sep ovate-triangular to oblong, 0.7–1.3  0.7–1.3 mm, margin entire to erose distally, lepidote to glabrous, apically praemorse; Pet oblong-elliptic to ovate-elliptic, 2–3  1.4–2 mm, entire, glabrous, apically denticulate to praemorse; St equal, Fil basally briefly adnate to the petals; female Inf 1 branched (or if 2 branched then with 1–2 short secondary branches); primary Br ascending to subascending, (6.5–) 15–31 cm, lepidote, densely manyflowered throughout or flowers disposed in verticills; floral Bra shorter than to exceeding the pedicels, chartaceous, narrowly ovatetriangular, 1.3–2.2  0.8–1.5 mm, lepidote, entire to serrulate or erose, attenuate-acuminate to praemorse; Fl divaricate to ascending; Ped articulated, stout, 0.8–1.5 mm, lepidote; Sep triangular to ovate-triangular, 1.7–2.1  1–1.8 mm, glabrous to weakly lepidote, entire and hyaline at the margins, rounded to acute or praemorse; Pet narrowly ovatetriangular, 3.1–4  1.2–1.9 mm, glabrous, rounded; Ov glabrous; Fr 8–10  3–4.5 ( 5.5) mm, glabrous, shiny and somewhat ridged, reticulate.

1009

H. huamelulaensis I. Ramírez & Carnevali (Phytotaxa 178(2): 119–121, ills., 2014). Type: Mexico, Oaxaca (Ramírez & Carnevali 1675c [CICY]). — Distr: Mexico (Oaxaca); xerophytic vegetation, 60 m. Caespitose in small groups, flowering to 1.5 m tall; Ros 50–60 cm ∅; L 30–40, sheaths yellowish-green, transversely oblong, 7–8  10–12 cm, glabrous on both faces, L lamina dull green, narrowly triangular, 30–35  6–8 cm, glabrous adaxially, white-lepidote abaxially, longacuminate, Sp red, antrorse, 3–5 mm, 1–1.5 cm apart; male Inf 1 branched with 45–55 branches; peduncular Bra foliaceous, sheaths triangular, 2–3  4–7 mm, lamina narrowly triangular, 2–10  0.5–0.8 cm, margins entire to finely serrate, sparsely white-lepidote at the apex; Br divaricate, 2.5–13.5 cm; floral Bra broadly ovate, 2.3–2.6  2 mm, margins erose, acuminate, glabrous; Fl polystichous; Ped 1 mm; Sep ovate to triangular-ovate, 2.5–3.5  1.3–2 mm, acute; Pet elliptic, 4.5–6  2.5–3 mm, rounded; St subequal; Fil 4–4.7 mm; Anth 1.6–2 mm; female Inf 1 branched with 45–100 branches; peduncular Bra foliaceous, sheaths broadly triangular, 3–5  5–9 mm, lamina narrowly triangular, 2–12  0.7–1 cm, margins finely serrate, sparsely whitelepidote; Br divaricate, 1.5–13 cm; floral Bra basally green, apically brown, oblong to triangular, 2.4–4.7  1.5–3 mm, margins erose, acuminate, glabrous; Fl polystichous; Ped 1.5–3 mm; Sep purple, triangular, 1.3–2.6  2–2.5 mm, margins entire to erose, acute; Pet white with a red stripe, ovate, 4–5.2  2–2.7 mm, acute; Ov light green, purple at the apex, ovoid, 3.2–3.5 mm; Fr narrowly ellipsoid, 9–12.5 mm; Se 6–8.5 mm. H. iltisii Burt-Utley & Utley (Brittonia 45: 222–225, fig. 2, 1993). Type: Mexico, Jalisco (Burt-Utley & Utley 7703 [MEXU, DUKE, MICH, MO, NY, US, WIS]). — Distr: Mexico (Jalisco: La Huerta); thorn-scrub vegetation, on rock outcrops, 250–300 m. Caespitose, flowering 0.93–2.1 m tall; Ros to 70 cm ∅; L numerous, straight to occasionally falcate, sheaths stramineous, transversely elliptic to reniform, 3.5–5.6  5.5–9.5 cm, glabrous becoming lepidote distally on both faces,

1010

somewhat inflated in older leaves, spinulose, L lamina 42.5–62.5 cm, glabrous adaxially, lepidote abaxially, Sp antrorse, occasionally retrorse, 1.5–4.5 mm, 0.5–3 cm apart; male Inf 1 branched; primary Br subascending; floral Bra pink, broadly ovate, 5–8  4–6.5 mm, acuminate, lepidote, finely spinulose-serrate, longer than the pedicels; Fl pedicellate or occasionally subsessile; Ped stout, 0.5–3 mm, lepidote; Sep pink but drying dark maroon, ovate to ovate-triangular, 4.5–8  3.5–4.7 mm, lepidote, rounded to acute; Pet green, ovate, 5.5–9  3–4.5 mm, glabrous abaxially, somewhat cucullate apically; St shorter than the petals; female Inf 1 or very rarely 2 branched; primary Br subascending; floral Bra pink but drying maroon, ovate to broadly ovate, 5–11  3–6.5 mm, attenuate-acuminate, lepidote, finely spinulose-serrate to erose, shorter or longer than the pedicels; Fl pedicellate; Ped stout, triquetrous, 2–6 mm, lepidote; Sep pink, triangular, 5–7  2.8–3.8 mm, lepidote, attenuate-acuminate, very finely erose to serrulate distally; Pet green, triangular, 7–10  2.5–4 mm, glabrous abaxially; Fr olive-brown, (10–) 15–21  6–9 ( 11) mm, glabrous, smooth. In the protologue the presence of abnormal branches and flowers that occasionally had four sepals and petals, as well as ovaries with four locules have been mentioned. Staminate flowers on normal inflorescences occasionally had five sepals and petals and up to 10 stamens. Similar variations in the number of sepals and petals were observed in other species such as H. glomerata, H. montana (described for the synonymous H. gayorum) or H. tillandsioides. H. isthmusiana Burt-Utley (Phytoneuron 2012(69): 10–13, ills., 2012). Type: Mexico, Oaxaca (Utley & Utley 8491 [MEXU, CAS, GH, MO, NY, US, USF, XAL]). — Distr: Mexico (SE Oaxaca: Ciudad Ixtepec, Asunción Ixtaltepec, Tehuantepec); thorn scrub and tropical deciduous forests, on the ground or on rocks on low rocky hillsides and slopes, 0–300 m. I: Ramírez Morillo & al. (2014b). Caespitose, caulescent, flowering 0.6–1 m tall; Ros 25–50 cm ∅; L several to many, sheaths pale

A. Espejo-Serna et al.

to dark castaneous, 2–4  (2.8–) 4–8 cm, glabrous and lustrous on both faces, becoming lepidote distally, margins finely spinulose and floccose, L lamina bright green to pale dusty rose, straight to subfalcate, 12–32  1.5–3 cm, appressedly cinereous-lepidote on both faces, Sp generally antrorse, (0.5–) 1.8–2.5 mm, (0.6–) 0.8–1.8 cm apart; Inf glabrous, typically 1 branched; male Inf with 14 ascending primary Br (1.8–) 2.5–7.5 cm long; floral Bra chartaceous, ovate to oblong, 2–3.5 ( 5)  1–2.5 ( 3) mm, glabrous, attenuate-acuminate to apiculate, margin finely denticulate to serrulate or erose distally; Fl divaricate to ascending; Ped articulated, conical, 1.2–3.5 mm; Sep white to cream, ovate to ovatetriangular, 1.5–2.7  1–1.8 mm, apically praemorse, margin hyaline; Pet white to cream but often tipped rose, elliptic to ovate-elliptic, 3.5–4.5  1.9–3.3 mm, glabrous; female Inf with 19 ascending primary Br (0.1–) 1.5–3.5 ( 4.5) cm long; floral Bra chartaceous, ovate-triangular to oblong, 2–3.5  1.2–2 mm, apiculate to acuminate, margin erose to very finely serrulate, hyaline; Fl divaricate; Ped articulated, 1.5–2.5 mm; Sep white or greenish to cream, triangular, 1.8–2.5  1.2–1.5 mm, glabrous, acuminate to apiculate, praemorse; Pet white or greenish to cream but often tipped rose, triangular, 4–5  1.5–2.5 mm, glabrous, margin hyaline; Fr dark castaneous, (6–) 7.5–11  3–4.5 mm ∅. H. ixtlanensis Burt-Utley (Phytoneuron 2012 (69): 1–5, ills., 2012). Type: Mexico, Oaxaca (Utley & Utley 7961 [MEXU, B, C, CAS, GH, M, MICH, MO, NY, US, USF, XAL]). — Distr: Mexico (Oaxaca); pine-oak forest, on moist steep slopes, 1650–1900 m. Forming clumps, flowering 1.5–2.7 m tall; Ros to 80 cm ∅; L very numerous, sheaths stramineous but oldest bases brown to castaneous distally, semiorbicular to transversely elliptic, 5–8.5  8–12 cm, margin finely spiny and distally floccose, glabrous and lustrous on both faces, becoming densely lepidote distally, L lamina straight to subfalcate, 32–56  2–3.5 ( 4.5) cm, appressedly cinereous-lepidote but occasionally glabrescent on both faces, Sp red-brown, antrorse

Hechtia BROMELIACEAE

to retrorse, (1.5–) 3–6 ( 8) mm, 1.5–4.5 cm apart; Inf lepidote; peduncular bracts foliaceous, pungent, becoming progressively reduced distally; male Inf 2 branched or rarely 1, with numerous divaricate primary Br 8–25 cm long, and with (0–) 5–9 ( 11) divaricate to ascending secondary Br 0.5–9.5 cm long; floral Bra ovate-triangular to broadly ovate, 1.2–3  0.8–1.8 mm, lepidote, finely spinulose-serrulate; Fl very numerous, divaricate; Ped articulated, conical, stout, 0.7–1.8 mm, glabrous to sparingly lepidote; Sep ovate-triangular to triangular, 1–1.6  0.8–1.8 mm, glabrous to sparingly lepidote, apiculate, entire to erose or very finely serrulate; Pet cream, ovate to elliptic, (2–) 2.5–3.2  1.5–2 mm, glabrous, rounded to praemorse; female Inf 1 or rarely 2 branched, with up to 50 primary Br 6–22 cm long, and with 0–2 secondary Br 3–6 cm long; floral Bra ovate-triangular, 1.3–2.5  1–1.6 mm, glabrous to sparingly lepidote, proximally irregularly and finely serrulatespinulose, apiculate; Fl very numerous, sometimes verticillate, ascending; Ped stout, 0.5–1.5 mm, glabrous to lepidote; Sep ovate-triangular, 1–1.5  0.7–1.5 mm, glabrous to lepidote, apiculate, erose distally; Pet greenish-yellow, triangular, (1.5–) 2–3  1–1.5 mm, glabrous, rounded to acute; Fr greenish-brown, 6.5–8  2.8–4 mm, smooth but sparingly finely ridged, glossy. H. jaliscana L. B. Smith (Phytologia 10: 482, t. 1, fig. 10, 1964). Type: Mexico, Jalisco (McVaugh 18530 [MICH]). — Distr: Mexico (Jalisco: Amatitán, San Cristóbal de la Barranca, Zapopan); steep moist ravines with tropical deciduous forest, on rocks, 750–1300 m. Caespitose, flowering to 1.8 m tall; L numerous, sheaths elliptic, 5–6 cm, glabrous basally, serrate distally, L lamina 50–80  2.7–3 cm, glabrous adaxially, finely appressedly cinereouslepidote abaxially, margin undulate, Sp divaricate to antrorse, 3 mm, 1.7–3 cm apart; Inf 1 branched, sparsely pale-lepidote, becoming glabrous; male Inf with 10–15 primary Br; basal peduncular Bra linear, to 15 cm, margins spiny; floral Bra ovate, 6–9 mm, acuminate, erose; Fl ascending to divaricate, sessile to shortly

1011

pedicellate; Ped 2.5 mm; Sep whitish, ovatedeltoid, 4–5  4 mm, glabrous, acute; Pet white, oblong-elliptic, 6–7  3.5–4 mm, acute to rounded; St equalling the petals; Fil 5.5–6 mm; Anth yellow, 1.5–2 mm; female Inf with 20 primary Br 7–20 cm long; primary Bra lanceolate, caudate-acuminate, 2–4 cm, serrate; floral Bra green towards the base and brown towards the apex, lanceolate-ovate, 9 mm, acuminate, minutely denticulate, about equalling the sepals; Fl divaricate, laxly and subverticillately disposed; Ped stout, cylindrical, 4 mm, subalate, articulate; Sep brown with prominent scarious margin, ovate, 3.5–4 mm, attenuate, irregularly erose, acute or obtuse; Pet white, triangular-ovate, 3.5–4  3 mm, attenuate, acute; Fr brown, 8–11  4–5 mm, sparsely lepidote, glabrescent and lustrous when mature. H. laevis L. B. Smith (Phytologia 10: 482–483, t. 1, figs. 11–12, 1964). Type: Mexico, Colima (McVaugh 15528 [MICH, MICH, US]). — Distr: Mexico (Colima); open rocky slopes with tropical deciduous forests, also reported from chalky soils, on the ground or on rocks, 400–1000 m. I: Ramírez-Morillo & al. (2016: 263). Caespitose, flowering 1–2 m tall; L numerous, densely rosulate, sheaths stramineous to light brown, transversally oblong to oblong, 4–4.5  4–7 cm, glabrous, barely lepidote distally, lustrous, margins serrate, L lamina rigid, 20–60  2.5–3 cm, pale appressed-lepidote on both faces, becoming glabrous adaxially, Sp reddish, antrorse, 2.5–5.5 mm, 1.4–2 cm apart; Inf 1 branched, sparsely white-lepidote when young, glabrescent with age; peduncle straight, 4–10 mm ∅; peduncular Bra erect, entire or laxly spiny, the lower linear from a small triangular base, the upper narrowly triangular; male Inf with 30–45 divaricate to ascending primary Br 3–25 cm long; floral Bra minute, narrowly triangular, densely whitelepidote; Fl divaricate, laxly disposed; Ped filiform, 1.5–3 mm, white-lepidote; Sep white with the median nerve purple, oblong-ovate, 1–1.3  1 mm, erose, densely white-lepidote; Pet white to yellowish, widely ovate to suborbicular, 2–2.4  2 mm, obtuse; St subequal,

1012

shorter to as long as the petals; Fil 1.4 mm; Anth 1 mm; female Inf with 20–40 divaricate to ascending primary Br 5.5–20 cm long; floral Bra minute, 2  0.3–0.5 mm, shorter than the pedicels; Fl divaricate, laxly disposed; Ped slenderly cylindrical, 2.5–3.5 mm, white-lepidote, smooth; Sep purplish, triangular, 2.2–2.4  1.2–1.5 mm, sparsely white-lepidote, acute; Pet white with the median nerve purplish, narrowly triangular, 3.5–4.4  1.8–2 mm, acuminate; Ov purple, ellipsoid, 3.4  2.2 mm; Fr brown, 8 mm, deeply sulcate between the carpels, smooth, covered at first with appressed white scales; Se brown, 5–6 mm. Material from Jalisco identified as this species was recently described as H. santanae. H. lanata L. B. Smith (Phytologia 8: 5, t. 1, figs. 6–9, 1961). Type [neo]: Mexico, Oaxaca (Ceja & al. 1928 [UAMIZ, IEB]). — Lit: Espejo-Serna & al. (2012). Distr: Mexico (Oaxaca: Santiago Lachiguiri), thorn scrub and tropical deciduous forest, rocky slopes and cliffs, on rocks, 750–1300 m. I: Kemble (2016). – Fig. 3. Solitary, caulescent, flowering to 1.2 m tall; Ros 30–40 cm ∅, forming long rhizomes to 30 cm and 4 cm ∅; L numerous, reflexed in the apical part, sometimes the tips strongly curled, Fig. 3 Hechtia lanata (near Santiago Lachiguiri). (Copyright: A. EspejoSerna)

A. Espejo-Serna et al.

sheaths stramineous to castaneous, suborbicular to broadly ovate, 4–6  4–8.5 cm, margins finely spinulose, floccose at the apex, glabrous and lustrous on both faces becoming densely lepidote distally, L lamina green, occasionally pink-tinged, 12–110  3–8 cm, very densely white-lepidote on both faces, most conspicuous on the lower , glabrescent adaxially in the apical part, Sp green to dark brown, antrorse and retrorse, or rarely divaricate, 3–8 mm, 4–7.5 cm apart; Inf densely lepidote, glabrescent with age; peduncle cylindrical, 4–10 mm ∅; peduncular Bra rose-greenish, linear to narrowly triangular, from a deltoid base, 3–18 cm, becoming progressively reduced distally and exceeding the internodes; male Inf 1 or rarely 2 branched, with 35–50 divaricate to ascending primary Br 2.5–17 cm long; floral Bra shorter than to equalling the sepals, ovate to triangular, 2–2.3  0.3–0.8 mm, finely spinuloseserrulate, attenuate-acuminate, lepidote, acute; Fl divaricate, sometimes appearing verticillate; Ped articulated, stout, conical, 0.5–2 mm, lepidote; Sep green to light brown apically, triangular, 1.5–2  1–1.6 mm, apiculate, occasionally finely serrulate-spinulose or erose esp. distally, lepidote, acute; Pet white, elliptic, 3–4.5  2–2.5 mm, entire, rounded to acute, apiculate, glabrous; St longer than the petals; Fil 4.5 mm; Anth yellow,

Hechtia BROMELIACEAE

1 mm; female Inf 1 or infrequently 2 branched, with up to 34 divaricate to ascending primary Br (1.5–) 4–9 cm long; floral Bra narrowly triangular, 1–1.5  0.4–0.7 mm, apiculate, entire to finely serrulate-spinulose, lepidote to occasionally glabrous; Fl divaricate, laxly disposed, sometimes appearing verticillate; Ped articulated, stout, conical, 1.5–2.5 mm, lepidote to occasionally glabrous; Sep green, apically brown, triangular, 2–2.5  2–2.3 mm, entire to irregularly finely serrulate, lepidote to occasionally glabrous, acute; Pet white, with a broad green central line, ovate, 3.1–3.5  3.1 mm, entire, apiculate to acute, glabrous; Ov green, ovoid, 4.5–5  1–2 mm; Fr dark brown, drying olivegreen, 8–10  3–4 mm, lustrous, glabrous to finely lepidote; Se reddish-brown, 2.8–3 mm  0.5 mm. H. laxissima L. B. Smith (Contr. US Nation. Herb. 29: 521–522, fig. 77, 1954). Type: Mexico, Michoacán (Moore & al. 5770 [US]). — Distr: Mexico (Michoacán: Múgica, Gabriel Zamora); on cliffs, 400 m. Caespitose, flowering >1.7 m tall; L numerous, sheaths not described, L lamina 40  4 cm, glabrous adaxially, finely veined and covered with pale appressed scales abaxially, Sp pale brown, 4 mm; Inf seemingly lateral, glabrous; peduncle flattened at the base, 1 cm ∅; peduncular Bra stramineous, triangular, acuminate, thin; male Inf unknown; female Inf 1 branched, with 15 divaricate or ascending, very slender primary Br 20 cm long; floral Bra greenish-rose, ovate, acute, mostly about as long as the pedicels; Fl divaricate, very laxly disposed; Ped slenderly cylindrical, 2–4 mm; Sep dark rose with a greenish tinge, very broadly ovate, 3 mm, acute; Pet green with rose tips, ovate, 5 mm, acute; Ov glabrous; Fr unknown. H. liebmannii Mez (Bot. Jahrb. Syst. 30 (Beiblatt 67): 6, 1901). Type: Mexico, Puebla (Liebmann s.n. [C, B]). — Distr: Mexico (Puebla, Veracruz); thorn scrub on lava flows, on the ground or on volcanic rocks, 2400–2500 m. Incl. Hechtia perotensis I. Ramírez & Martínez-Correa (2007).

1013

Caespitose to densely caespitose, flowering 0.55–2 m tall; Ros compact, globose, to 50  40 cm, sometimes slightly asymmetrical; L numerous, sheaths yellow to brownish, lustrous, widely ovate to subquadrate, 3.5–5.5  3.5–8.5 cm, white-lepidote apically, L lamina light green, 15.7–30.5  1–1.8 cm, glabrescent adaxially, minutely white-lepidote abaxially, Sp brown to dark brown, antrorse to divaricate, 5–6 mm, 1.5–2 cm apart; Inf 2–3 branched; peduncle cylindrical, 1–3 cm ∅; peduncular Bra brown, triangular to narrowly triangular, 3–20  1–3.5 cm, acute to acuminate, densely imbricate, sheaths entire, lamina erose to spiny, densely lepidote at the base abaxially, glabrescent adaxially; male Inf with numerous, ascending, glomerate to shortly elongate primary Br 2–3.5 cm long with 2 basal shorter secondary Br; primary Bra brown, ovate to narrowly ovate, 2.5–7  0.8–1 cm, lepidote abaxially, erose, acuminate; floral Bra green, lanceolate to oblong, 4 mm; Fl divaricate, laxly disposed; Ped 2–3 mm; Sep green, ovate to narrowly ovate, 4.5–5  2–2.5 mm, acute to rounded; Pet green to yellowish, elliptic to oblong, 6–7  3–4 mm, rounded; St as long as or shorter than the petals; Fil 5 mm; Anth yellow, curved, 1.2–1.5 mm; female Inf with numerous ascending glomerate to elongate primary Br 1.5–8 cm long; primary Bra widely triangular, 2–2.5  1–1.3 cm, acuminate to acute; floral Bra yellowish, triangular, 3–5 mm; Fl densely disposed, ascending; Ped obconical, 2–4 mm; Sep brownish, widely triangular, 4–5  2.4–2.7 mm, acute and apiculate; Pet green, triangular, 6  3 mm, acute; Ov green, ovoid; Fr brown, 9–10  3.5–7 mm, lustrous; Se light brown, 2.9–3.5 mm. H. lundelliorum L. B. Smith (in N. L. Britton & al. (eds.), North Amer. Fl. 19: 97–98, 1938). Type: Mexico, San Luis Potosí (Lundell & Lundell 7265 [MICH]). — Lit: López-Ferrari & al. (2011). Distr: Mexico (San Luis Potosí, Querétaro, Hidalgo, Puebla); tropical deciduous and semideciduous forest, limestone ledges and cliffs, on rocks, 150–1250 m. Forming large pendulous colonies, flowering 0.8–1.8 m tall; Ros asymmetrical in mature plants, 30–120 cm long; L numerous, very long and

1014

pendent, sheaths white to yellowish, broadly oblong to oblong, 5–6  3–5 cm, minutely serrate esp. towards the apical portion, white-lepidote on both faces, L lamina light green adaxially, densely white-lepidote abaxially, long-attenuate, 60–110  1–2 cm, margins minutely serrate; Inf erect to arched and/or pendulous, 3 branched; peduncle brown, cylindrical, 0.8–2 cm ∅, glabrous; peduncular Bra brown to light brown or green, foliaceous, linear to long triangular, 5–70  1 cm, entire to minutely denticulate, densely lepidote abaxially, glabrous to glabrescent adaxially, attenuate; male Inf with numerous ascending stalked primary Br 25–40 cm long; primary Bra light brown, linear to triangular, 2–25 cm  4–6 mm, long-attenuate to acuminate; secondary Br numerous, ascending to appressed, 6–13 cm long, stalked; secondary Bra triangular, 2  1 mm, deciduous; tertiary Br numerous; Fl numerous on each branch, polystichous; Ped filiform, 1.5–2.5 mm; floral Bra light brown, triangular, 2  2.5 mm, rounded; Sep brown to light brown suffused with white, slightly rose towards the margins, ovate to triangular, 1.2  1.4 mm, glabrous, entire, rounded; Pet white, elliptic, 3.5–4.2  2.1–2.4 mm, rounded; St subequal, shorter than the petals; Fil 2–3 mm; Anth yellow, 1.4 mm; female Inf with numerous ascending stalked primary Br 25–35 cm long; primary Bra brown to light brown, linear to triangular, 3–12 cm  4.5–5 mm, acuminate; secondary Br numerous, ascending to appressed, 14–17 cm long; secondary Bra triangular, 2  1 mm, deciduous; tertiary Br numerous; Fl numerous on each branch, polystichous; Ped 1.4–1.7 mm; floral Bra light brown, hyaline, ovate to triangular, 1  1 mm, rounded; Sep light brown suffused with white, ovate to triangular, 1.5  1 mm, acute; Pet white, oblong-elliptic, 3.5  1.7 mm, rounded; Ov green, ovoid, 3  2 mm, glabrous; Fr brown to dark brown, slightly trigonous, 5–7  2–3 mm, glabrous, reflexed; Se brown to reddish-brown, 1.3 mm. H. lyman-smithii Burt-Utley & Utley (Brittonia 39(1): 37–40, ills., 1987). Type: Mexico, Oaxaca (Burt-Utley & Utley 6982 [MEXU, MO, NY, US]). — Distr: Mexico (Puebla:

A. Espejo-Serna et al.

Caltepec; Oaxaca: San Antonio Nanahuatipan, Santa María Ixcatlán, Santa María Tecomavaca); thorn scrub and tropical deciduous forest, steep limestone slopes and cliffs, on rocks, 600–1150 m. – Fig. 4. Densely caespitose, flowering to 80 cm tall; Ros 20–27 cm ∅; L few, sheaths reniform, 1.8–2.8  3.5–4.5 cm, margins inconspicuously spiny, glabrous and lustrous, becoming lepidote distally, L lamina light green, often suffused with purple, 7–17  1.7–3 cm, appressedly cinereouslepidote on both faces, Sp antrorse or occasionally retrorse, red-brown, 2.5–4 mm, 0.6–1.1 cm apart; Inf 1 branched, glabrous; peduncle 3–5.5 mm ∅; lower peduncular Bra exceeding the internodes, ovate with linear, erose, appressed, lepidote lamina, 1.8–5.7 cm; male Inf with 15–40 slender divaricate to ascending primary Br (0.5–) 2.2–14 cm long; floral Bra as long as or slightly exceeding the pedicels, ovate, 0.8–2  0.7–1 mm, lacerate to erose, acute to apiculate; Fl divaricate; Ped slender, 0.8–3 mm; Sep white, tinged with rose, ovate to broadly ovate, 1.3–2.3  0.9–1.8 mm, glabrous, margin hyaline and distally erose, rounded to acute; Pet white to rose, ovate to oblong-ovate, 3–5  2–3.1 mm, St longer than the petals; Fil 4–4.5 mm; Anth yellow-green to green, 2 mm, mucronate; female Inf with 12–40 slender divaricate primary Br 2–8.5 cm long; floral Bra shorter than the pedicels, narrowly ovate to ovate-triangular, 1–2  0.7–0.8 mm, margin hyaline, erose, acuminate; Fl divaricate to ascending; Ped stout, 1.5–3.5 mm; Sep light brown, ovate, 1.3–1.9  1.3–2.2 mm, glabrous, margin hyaline, acuminate, acute; Pet rose, triangular, 3–4  2–2.5 mm; Ov purple, ovoid, 3.5 mm; Fr purple, drying light brown, triquetrous, 5.5–10  5–6 mm, glabrous, smooth; Se yellowish-brown, 3.5–4 mm. H. malvernii Gilmartin (Ceiba 11(2): 9, fig. 4, 1965). Type: Honduras, El Paraíso (Gilmartin 966 [US, US]). — Distr: Honduras (El Paraíso); hillsides and bank of road cuts, 700 m; known only from the type collection. Growth habit not described, flowering to 2 m tall; L numerous, L lamina 50  3.5–4.5 cm, pale appressed-lepidote, becoming glabrous

Hechtia BROMELIACEAE

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Fig. 4 Hechtia lymansmithii. (Copyright: A. Espejo-Serna)

adaxially, glabrescent abaxially, margin undulate, Sp divaricate to antrorse, 3 mm, 0.9–1.1 cm apart; Inf glabrous, 1 branched; peduncular Bra to 16 cm, reflexed, lepidote abaxially, nearly glabrous adaxially; male Inf unknown; female Inf with 8 reflexed weak primary Br to 55 cm long; floral Bra narrowly ovate, 5–10  2 mm, glabrous, acuminate; Fl divaricate to reflexed, laxly disposed, subsessile; Sep reddish, ovate, 3  1.5 mm, glabrous, mucronate; Pet whitish, 5  1 mm; Ov inferior, ellipsoid; Fr 10 mm, glabrous. H. mapimiana López-Ferrari & Espejo (Acta Bot. Mex. 102: 90–92, ills. (pp. 91, 93–95), 2013). Type: Mexico, Durango (Mercado & al. 259 [CIIDIR, IEB, UAMIZ]). — Distr: Mexico (Durango: Lerdo); thorn scrub, limestone cliffs, on rocks, 1400–1750 m. – Fig. 5. Caespitose in small groups, flowering to 2.3 m tall; Ros 40–70 cm ∅; L numerous, sheaths light brown adaxially, dark brown abaxially, broadly ovate to subquadrate, 6–15  5.5–10 cm, lustrous, glabrous on both faces, L lamina glaucous, cylindrical-conical and slightly curved upwards at the apex, 30–45  2.5–4 cm, densely whitelepidote on both faces, Sp light brown, reddish at the apex, retrorse, 3–6 mm, 2–5 cm apart; male

Inf 2–3 branched, with numerous Br per node, apparently fasciculate by the reduction of the rachis; peduncular Bra foliaceous, sheaths ovatetriangular, papery, brittle, 4.5–5.5  2.6–3.2 cm, entire, very sparsely white-lepidote to glabrous, lamina narrowly triangular, 8–10  5–7 mm, densely white-lepidote on both faces, pungent, margins spiny; primary Br oblong-cylindrical, 5.4–6.8 cm, with 2 secondary basal branches 2.8–4.5 cm long; floral Bra white, filiform, 3–5 mm, shorter than the pedicels, entire, glabrous; Fl polystichous, ascending; Ped filiform, 5.5–11 mm; Sep white tinged with pink to purple, triangular, 3.6–4  1.5–1.7 mm, acute; Pet light green, elliptic, 6.5–7.5  4–4.2 mm, rounded; St subequal; Fil 3.2–3.5 mm; Anth green, 4.2–4.5 mm; female Inf 2 branched, with numerous Br grouped apparently in fascicles by the reduction of the rachis; peduncular Bra foliaceous, sheaths ovate-triangular, 5–6  2.8–3 cm, papery, brittle, entire, very sparsely white-lepidote to glabrous, lamina 7–16.5  0.5–0.7 cm wide, pungent, densely white-lepidote on both faces, margins spiny; primary Br oblong-cylindrical, 6–9 cm, with 2 solitary flowers at the base; floral Bra white, filiform, 3–5 mm, entire, glabrous, shorter than the pedicels; Ped filiform, 7–8 mm; Fl polystichous, ascending; Sep white tinged with pink to

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Fig. 5 Hechtia mapimiana. (Copyright: A. Espejo-Serna)

purple, triangular, 3.5  2.5 mm, acute; Pet light green, ovate-triangular, 5–5.2  2.5–2.7 mm, acute; Ov green, ovoid; Fr light brown, triquetrous, 10  8–9 mm, with thin papery walls when mature. H. marnier-lapostollei L. B. Smith (Bromeliad Soc. Bull. 11: 58, ills., 1961). Type: Mexico, Oaxaca (Schwartz s.n. in Marnier-Lapostolle s.n. [US]). — Distr: Mexico (Oaxaca: San Carlos Yautepec, San Juan Lajarcia); thorn scrub and tropical deciduous forest, on the ground or on limestone rocks, 750–1500 m. Caespitose, flowering to 1 m tall; L 15–20, sheaths yellowish, suborbicular, 2 cm ∅, glabrous and lustrous, densely lepidote distally, L lamina 15–28  2 cm, pungent, densely and closely pale-lepidote on both faces, Sp antrorse, slender, 2.5–4 mm, 0.12–1.3 cm apart; Inf 1 branched, glabrous; peduncle 2–4 mm ∅; peduncular Bra exceeding the internodes, stramineous, broadly ovate with a long linear lamina, 1–4 cm, pungent; male Inf with 8–24 ascending primary Br 1.5–5.5 cm long; floral Bra stramineous, broadly ovate, 2 mm, erose apically, acuminate, longer than the pedicel; Fl divaricate; Ped 1 mm; Sep white-purpureous, triangular-ovate, 2 mm; Pet white tinged with purple, elliptic,

3–4 mm, rounded; St as long as the petals; Fil 3.5 mm; Anth green, 1.5 mm, mucronate; female Inf with 22 divaricate to ascending primary Br 2–5 cm long; floral Bra broadly ovate, erose, 2–2.5 mm; Fl divaricate; Ped 2 mm; Sep ovate-triangular, 2 mm, acute; Pet triangular to narrowly triangular, with a large callus at the base, 3.8 mm; Ov ovoid, trigonous, glabrous; Fr dark brown, 5.5–9 mm, shiny. H. matudae L. B. Smith (Phytologia 5: 395–396, t. 1, figs. 1–2, 1956). Type: Mexico, Morelos (Matuda 26351 [US]). — Lit: LópezFerrari & al. (2008: with ills.). Distr: Mexico (México: Ocuilan; Morelos: Tepoztlán); oak and tropical deciduous forests, on the ground or on cliffs and crags of volcanic rocks, 1700–1900 m. – Fig. 6. Caespitose, forming large colonies, flowering 0.8–1.6 m tall; Ros symmetrical to slightly asymmetrical in mature plants, 40–80 cm ∅; L numerous, sheaths white to yellowish, broadly oblong to square, 6–12.5  5–10 cm, lustrous basally and white-lepidote apically on both faces, L lamina extended, pendent in mature plants, glaucousgreen, 19–80  1.7–6.5 cm, densely whitelepidote on both faces, glabrescent adaxially, Sp yellowish, antrorse, 1.6–2 mm, 0.6–1.2 cm apart;

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triangular to ovate-triangular, 7–7.5  3–3.5 mm, acute, entire; Ov purple, ovoid to elongate-ovoid, 5–5.1  2.5 mm, glabrous; Fr brown, 12–15  7–8 mm.

Fig. 6 Hechtia matudae. (Copyright: A. Espejo-Serna)

Inf 2 branched; peduncle cylindrical, 2.5–3 cm ∅; peduncular Bra straw-coloured, foliaceous, densely lepidote on both faces, becoming progressively reduced distally, densely imbricate; male Inf with up to 40 ascending to divaricate primary Br 6–11.5 cm long, each with 2 basal secondary branches; floral Bra white, linear, 4–8  1 mm, acuminate, entire; Fl ascending to divaricate, densely disposed; Ped 2.2–2.7 mm; Sep rose, triangular to triangular-oblong, 4.6–5.2  1.9–2.7 mm, acute, entire; Pet rose, widely elliptic to oblongelliptic, 6.5–8.5  4–4.7 mm, rounded, entire; St subequal, shorter than the petals; Fil 2.8–4 mm; Anth purple to dark purple, 1.5–1.8 mm; female Inf with 30–35 primary Br 7.8–8 cm long, each with 2 basal secondary branches; floral Bra white, linear, 4.5–8  1.5 mm, acuminate, entire; Fl ascending to appressed, densely disposed; Ped 4.5 mm; Sep rose, linear triangular, 4–4.2  1 mm, acuminate, entire; Pet rose to dark rose,

H. melanocarpa L. B. Smith (Contr. Gray Herb. 161: 32, t. 4, figs. 8–9, 1946). Type: Mexico, Guerrero (Foster 1258 [GH]). — Distr: Mexico (Guerrero: widespread); tropical deciduous forest, on steep limestone slopes and riversides, 550–1350 m. Solitary or with few rosettes, flowering 2–3 m tall; L numerous, sheaths widely ovate, 7  5–8 cm, serrate, white-lepidote distally, L lamina green, 80–120  3.4–5 cm, glabrous adaxially, appressedly white-lepidote abaxially, Sp reddish, antrorse, 5–7 mm, 2.2–3 cm apart; Inf erect to ascending, 2 branched, sometimes the lower branches appearing verticillate, white-lepidote; peduncle cylindrical, 1.8–2 cm ∅; peduncular Bra foliaceous, pungent, margins spiny, becoming smaler distally; male Inf with >35 divaricate to ascending primary Br 15–37 cm long, with 2–4 ( 8) basal secondary branches; floral Bra light brown, ovate, acuminate, erose, 3 mm; Fl divaricate, laxly disposed, sessile; Sep green tinged with purple dots, triangular-ovate, 2.7–3.2  1.7–2.2 mm, glabrous, acute; Pet white, elliptic to oblong, 4.5–5  2.5–3 mm, rounded; St longer than the petals; Fil 4.3–5.3 mm; Anth green to yellowish, 2 mm; female Inf with >25 divaricate to ascending primary Br 8–35 cm long, with 2 basal secondary branches; floral Bra light brown, ovate, 4.7–5.2 mm, lepidote, erose, acute, long mucronate; Fl ascending; Ped stout, 1 mm; Sep green with purple dots, triangular to ovate, 3.7–4  2.5–3 mm, glabrous, acute; Pet white, ovate-triangular, 5.5–6  3.7–4 mm; Ov rose, ovoid, 4.5–5  2.6–3 mm, glabrous; Fr black, 9–14  4.5 mm, glabrous; Se reddishbrown, 5 mm. H. mexicana L. B. Smith (Contr. Gray Herb. 117: 19–20, t. 1, fig. 60, 1937). Type: Mexico, San Luis Potosí (Maury 6593 [F, GH]). — Distr: Mexico (Tamaulipas, San Luis Potosí); tropical deciduous and semi-deciduous forests, on rocks, 200–300 m.

1018

Usually forming clumps of 3 or more rosettes, flowering to 2 m tall; L numerous in spreading rosettes, sheaths yellowish to light brown, with a dark brown central spot, suborbicular, 6–9  5–9 cm, glabrous, lustrous, L lamina 80–100  2.5–4 cm, glabrous and shiny adaxially, whitelepidote abaxially, Sp light brown, divaricate to antrorse, 3–5.5 mm, 1.3–2.3 cm apart; Inf seemingly lateral, erect to ascending, densely lepidote, 2 branched; peduncle 12–15 mm ∅; peduncular Bra triangular to narrowly triangular, acuminate, densely white-lepidote; male Inf unknown; female Inf with 16–30 divaricate to ascending primary Br 2–18 cm long, with 2 basal secondary branches; floral Bra brown, broadly ovate, 3.5–4  2.5 mm, margins hyaline, densely white-lepidote, acute to acuminate; Fl divaricate, subsessile; Sep brown, broadly ovate, 3.5 mm, lepidote, rounded, mucronate; Pet white, ovate, 6 mm; Fr dark brown, 10 mm, sparsely lepidote, acute. H. montana Brandegee (Erythea 7: 9, 1899). Type: Mexico, Baja California Sur (Brandegee s.n. [US, NY]). — Lit: Felger & Yetman (2000). Distr: NW Mexico (Sonora, Baja California Sur); thorn scrub, steep ledges, on the ground or on rocks. Incl. Hechtia montana ‘Burgundy’ Kimnach & Trager (1993); incl. Hechtia gayorum L. W. Lenz (1995); incl. Hechtia gayii L. W. Lenz (1995) (nom. inval., Art. 61.1). Caespitose, forming dense colonies, flowering to 1 m tall; Ros compact, small, 30 cm ∅, rhizomatous; L numerous, sheaths light brown to stramineous, ovate to suborbicular, 3.5  3–3.5 cm, margins finely ciliolate and spinydenticulate, glabrous or sparingly lepidote, L lamina grey-green, 15–45  2–3 cm, lepidote but glabrescent adaxially, white-lepidote abaxially, margins sinuate, Sp reddish-brown, retrorse, 2–5 mm, 1.7–2 cm apart; Inf 1 branched, white-lepidote, glabrescent with age; peduncle cylindrical, 7 mm ∅; peduncular Bra narrowly triangular, 10–10.5  0.5 cm, margins spiny, becoming smaller distally; male Inf with 17–25 divaricate to ascending primary Br 3–8 cm long; floral Bra hyaline, linear to linear-triangular, 2–2.5 mm; Fl laxly disposed, divaricate to reflexed;

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Ped 4–7 mm; Sep whitish, triangular, 3 mm, lepidote, acute; Pet yellowish-white, obovate, 5–6 mm, acute; female Inf with 8–9 ascending primary Br 4.5–12 cm long; floral Bra narrowly triangular, shorter than the pedicels; Fl laxly disposed, ascending; Ped 6–8 mm; Sep whitish, purplish basally, narrowly triangular, 3–3.5  1.2–1.3 mm, sparsely lepidote, acuminate to acute; Pet yellowish-white, oblong, 4–6  1.6 mm, rounded to acute; Ov green, glabrous, smooth, 3.3  1.5 mm ∅; Fr light to reddish-brown, 10–12  7–8 mm, rostrate; Se brownish, 5–5.5 mm. As in other species (H. iltisii, H. tillandsioides, H. glomerata), abnormal flowers are sometimes produced. This is the case of the cultivated H. gayorum (here treated as synonym) that comes from the same region. Felger & Yetman (2000) mention that the Guajirío, an ethnic group native to the lower Sierra Madre of SE Sonora and adjacent SW Chihuahua, have for many years used plants (which they call “hichiconi”) of H. montana as a supplemental food. H. montis-frigidi González-Rocha & al. (Acta Bot. Mex. 109: 44–47, ills., 2014). Type: Mexico, Morelos (López-Ferrari & al. 2206 [UAMIZ]). — Distr: Mexico (Morelos: Puente de Ixtla); oak forests, on the ground or on rocks, 1750–2120 m. Caespitose in small groups, flowering to 2 m tall; Ros 60–70 cm ∅; L numerous, sheaths straw-coloured adaxially, light brown and lustrous abaxially, depressed-ovate, 5.5–6  5–7.5 cm, glabrous at the base, densely white-lepidote at the apex on both faces, L lamina dark green, narrowly triangular, 32–48.5  2.8–4.8 cm, lepidote on both faces, Sp yellowish to brownish, antrorse, 1.5–5 mm, 1–2 cm apart; male Inf 3 branched with up to 97 primary branches; peduncular Bra foliaceous, sheaths ovate triangular, 3–3.4  2.4–2.5 cm, lamina narrowly triangular, 10–13.2  0.3–0.5 cm, glabrescent adaxially, sparsely white-lepidote abaxially, acuminate, margins entire to slightly erose; primary Br ascending, 9–18 cm, sessile to shortly stalked; secondary Br longer than primary branches, 3–10 cm; tertiary Br when present 2 cm long; floral Bra light brown, triangular, 1.7–3.8 mm,

Hechtia BROMELIACEAE

equal to slightly longer than the pedicels, entire to slightly erose, lepidote; Fl polystichous, ascending; Ped 1.5–3 mm; Sep light brown, triangular, 1.5–3  1.1–2 mm, acute and shortly apiculate; Pet white-greenish, ovate-elliptic, 3.1–5  2–3 mm, rounded to slightly emarginate; St subequal; Fil 3–5.5 mm; Anth green, 1–2 mm; female Inf 2 branched with 30–60 primary branches; peduncular Bra ovate-triangular, 3–14  0.6–2.5 cm, glabrescent adaxially, densely white-lepidote abaxially, acuminate, margins entire to slightly erose; primary Br ascending, 6–23 cm, sessile; secondary Br 4–10.5 cm; floral Bra brownish, narrowly triangular to ovate, 2.8–5.6  0.6–1.6 mm, entire to slightly erose, lepidote, longer than the pedicels; Ped 1–2 mm; Fl polystichous, ascending; Sep green-brownish, triangular, 2–3  1.7–2.5 mm, densely lepidote, acute and shortly apiculate; Pet green-brownish, triangular, 3.5–4.6  2–3 mm, acute; Ov green, ellipsoid; Fr brownish, ovoid, 10–13  3–7 mm; Se brownish, 3.5–6 mm. H. mooreana L. B. Smith (Contr. US Nation. Herb. 29: 522–523, fig. 78, 1954). Type: Mexico, Guerrero (Moore 5196 [US, BM, G, UC]). — Distr: Mexico (Guerrero: Chilapa, Copalillo, Eduardo Neri, Mártir de Cuilapan, Tepecoacuilco); tropical deciduous forest, on the ground or more rarely on rocks on steep limestone slopes, 500–1300 m. Solitary or in clumps of few caulescent elongated rosettes, flowering to 1.5 m tall; stems conspicuous, 25–50  1.3–2 cm ∅; L numerous, sheaths stramineous, suborbicular to ovate, 3.8–4  3.6–4 cm, glabrous, densely white-lepidote and spiny at the margins distally, L lamina light green, squarrose, 15–35  0.85–1.5 cm, glabrous adaxially, densely white-lepidote abaxially, Sp light brown, antrorse, 2.4–3.3 mm, 1.1–2 cm apart; Inf 1 branched, glabrous; peduncle 1 cm ∅; male Inf with 15 divaricate primary Br 10 cm long; floral Bra hyaline with purple dots, narrowly triangular, 1 mm, erose, acuminate; Fl divaricate, subverticillate; Ped slender, ribbed, 2–2.5 mm; Sep white with purple dots, ovate-triangular, 1.5–1.7  0.7 mm, glabrous, rounded; Pet white, elliptic, 2.9–3.1  1.7 mm, rounded; St  equalling the petals; Fil 1.5 mm;

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Anth yellow, 1.3 mm; female Inf with 25–32 divaricate to ascending primary Br 3–10 cm long; primary Bra 1.2 mm; Fl divaricate to ascending, laxly disposed; Ped 1.9–2.1 mm; Sep white, ovate, 1.9–2.2  1–1.2 mm, margin hyaline, rounded; Pet white, triangular, 3.2–3.4  1–1.1 mm, acuminate; Ov green, ovoid, 3 mm; Fr brown to dark brown, 7–8  2–4 mm, glabrous, smooth; Se light brown, 2.2 mm. H. myriantha Mez (Bot. Jahrb. Syst. 30 (Beiblatt 67): 6, 1901). Type: Mexico, Veracruz (Liebmann s.n. [C, B]). — Distr: Mexico (Veracruz: Actopan, Coatepec, Emiliano Zapata, Jalcomulco); tropical deciduous forest and arid scrub, on the ground or on rocks on rocky slopes, 0–800 m. I: Espejo-Serna & al. (2005: 91). Incl. Hechtia macrophylla Greenman (1907). Solitary, caulescent, flowering to 3–4 m tall; Ros globose to elongated, to 1.5 m high, 1 m ∅, stems conspicuous, to 20 cm ∅; L very numerous, sheaths white, with a dark brown spot to the apex, lustrous, broadly ovate to depressed-elliptic, 5.5–6 ( 9)  9.5–11.5 cm, spiny towards the apex, L lamina green, sometimes reddish at the margins, (0.4–) 1–1.7 m  2.5–6.5 cm, glabrous adaxially, white-tomentose and conspicuously veined abaxially, Sp antrorse to retrorse or divaricate, 3–7 mm, with an ample triangular base, 1–2 cm apart; Inf seemingly lateral, whitelepidote, 3 branched; male Inf with very numerous primary Br 8–40 cm long; peduncle cylindrical, 0.6–2 cm ∅; peduncular Bra brown, triangular, 6.5–12  1.5–2.5 cm, lepidote, acute to acuminate, longer than the internodes; primary Bra brown, triangular to lanceolate, 1.5–9 cm; floral Bra broadly triangular to ovate, 3–4  3 mm, lepidote, erose, acute and apiculate; Fl ascending to divaricate, loosely arranged, very numerous on each branch, subsessile; Sep connate at the base, obovate, 3  2 mm, rounded, apiculate; Pet white, oblong or obovate, 4–4.5  2–3 mm; St shorter than the petals; Fil 3 mm; Anth yellow, 1.5 mm; female Inf with very numerous primary Br 10–36 cm long; primary Bra brown, triangular, 1–4 cm, ovate to triangular; floral Bra brown to white, ovate, 3.8–5  3 mm, margins slightly fimbriate at the base,

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attenuate, pungent; Fl ascending to divaricate, loosely arranged, very numerous on each branch; Ped 1–5 mm; Sep brown to white, ovate to triangular, 3–3.5  2.5 mm, entire, acute; Pet white, oblong to oblong triangular, 3.6–4.1  2.2–2.5 mm, acute; Ov greyish to blackish, ovoid to longovoid, 4–5  2.5 mm, densely white-lepidote; Fr dark brown, 10–12  4–5 mm, white-lepidote to glabrescent; Se reddish-brown, 4–7 mm. H. nivea I. Ramírez & C. F. Jiménez (Phytotaxa 178(2): 122–124, ills., 2014). Type: Mexico, Oaxaca (Ramírez & al. 1826b [CICY, MEXU]). — Distr: Mexico (Oaxaca: San Juan Bautista Cuicatlán); xerophytic scrub, 700–800 m. Solitary, flowering to 1.4 m tall; Ros 35–45 cm ∅; L 60–70, sheaths white-yellowish adaxially, brownish abaxially, oblong, 2.5–3  3–5 cm, glabrous on both faces, L lamina green, narrowly triangular, 21–39  1.3–2.9 cm, glabrous to sparsely white-lepidote adaxially, white-lepidote abaxially, Sp reddish, retrorse, 4–7 mm, 1.6–4.5 cm apart; Inf 1 branched; male Inf with 65–75 branches; peduncular Bra foliaceous, sheaths triangular, lamina narrowly triangular, 4–11  0.8–1 cm, margins entire, glabrous; Br ascending, 1.2–3 cm; floral Bra light brown, elliptic, 6.3–8  3.5–5.5 mm, erose, acute, glabrous; Fl polystichous; Ped 1–2 mm; Sep straw-coloured, oblong, 3.5–4  2.2–3 mm, erose, acute, glabrous; Pet white, elliptic, 4–4.4  2.2–2.5 mm, rounded; St subequal; Fil 4–4.6 mm; Anth green, 1.7–2 mm; female Inf with 40–45 branches; peduncular Bra 2.5–9  0.8–0.9 cm, triangular, acuminate, entire, glabrous; Br ascending, 0.8–2.4 cm; floral Bra brown, triangular-ovate, 5–7  3–4 mm, erose, glabrous; Fl polystichous, sessile; Sep brown, triangular, 3–3.5  2.5–3.5 mm, apically erose, glabrous; Pet green, triangular to ovate, 3–4  2–2.8 mm, acute; Ov oblong, 2.5–3 mm; Fr brown, ellipsoid, 6.3–9 mm; Se brown, 3.4–4 mm. H. nuusaviorum Espejo & López-Ferrari (Acta Bot. Mex. 78: 98–102, ills., 2007). Type: Mexico, Oaxaca (Ceja & al. 1751 [UAMIZ, IEB]). — Distr: Mexico (Oaxaca: Santiago Yosondúa); oak-pine forest, on vertical limestone cliffs, 1700–1900 m.

A. Espejo-Serna et al.

Solitary or in clumps of few rosettes, flowering to 2 m tall; Ros caulescent, 50 cm ∅, stems conspicuous, succulent, woody, to 1.2 m  15–30 cm ∅; L numerous, very succulent, soft, sheaths light brown, widely ovate to square, 3.5–8.5  3.5–7.5 cm, lustrous, white-lepidote on both faces, L lamina lime-green, 30–75  1.5–4.5 cm, glabrous adaxially, lepidote abaxially, Sp divaricate to antrorse, green to brown, 4–9 mm, 1–2 cm apart; Inf seemingly lateral, flocculose, 1 branched; peduncle 1–2.5 cm ∅; peduncular Bra brown, triangular, 2.2–20  1.5–3.5 cm, acute, acuminate, lamina erose to spiny, densely lepidote, becoming progressively reduced distally; male Inf with 8–38 ascending, short to capitate primary Br 2.5–3.2 cm long; floral Bra light brown, ovate to triangular-ovate, 5–6  4–5 mm, acuminate; Fl divaricate, very densely disposed, sessile; Sep light brown, ovate, 4–4.7  2.3–2.4 mm, acute; Pet white, elliptic to oblong, 6–6.3  2.2–2.4 mm, acute; St subequal, as long as petals; Fil 5.5–6.1 mm; Anth yellow, 1.4 mm; female Inf with 20–25 divaricate, capitate and globose primary Br of 1.2–3 cm ∅; floral Bra brown, widely ovate, 5–7  5 mm, glabrous, entire; Fl densely disposed, sessile; Sep brown with margins white with rose dots, ovate, 4.7–5  4.3 mm, entire, acute-apiculate; Pet white with rose dots on the median nerve, elliptic, 6–7  3.5–3.8 mm, entire, acute; Ov ovoid, rose, 7–7.7  2.6 mm; Fr brown, 10  5 mm, lustrous; Se dark brown, falcate, 6 mm. H. oaxacana Burt-Utley & al. (Phytoneuron 2011(59): 3, 2011). Type: Mexico, Oaxaca (Utley & Burt-Utley 7960 [MEXU, CAS, GH, M, NY, US, USF]). — Distr: Mexico (N Oaxaca); oak and pine forest, steep hillsides and karstic rock outcrops, 1650–2400 m. Usually in large colonies, flowering to (0.72–) 1.16–1.77 m tall; Ros 30–50 cm ∅, occasionally with stout stolons; L very numerous, sheaths reniform to transversely elliptic, 2.3–5.6  (3.5–) 5–8.3 cm, lepidote distally on both faces, marginally spiny and floccose, L lamina (11–) 22–40.5  (1–) 1.5–2 ( 2.5) cm, lepidote to glabrescent adaxially, lepidote abaxially, Sp

Hechtia BROMELIACEAE

reddish-brown, antrorse, occasionally retrorse, (1.5–) 2.5–3.5 ( 4.5) mm, (0.4–) 1.4–2 ( 2.5) cm apart; Inf glabrous to lepidote, rachis often weakly geniculate; peduncle (6–) 8–14 ( 18) mm ∅; peduncular Bra ascending, (2.2–) 3.4–22 cm, with narrowly linear-triangular spiny lamina; male Inf 1 branched with numerous ascending to subascending primary Br (1–) 2–11 ( 17.5) cm long, or rarely 2 branched with very short basal secondary branches; floral Bra shorter to longer than the sepals, narrowly to broadly ovate or ovate-triangular to suborbicular, (2.2–) 3–7 ( 9)  (1.7–) 2.5–5 ( 6) mm, glabrous to finely lepidote, margin finely spinulose to erose, spinulose to apiculate, acute, rounded or praemorse at the apex; Fl divaricate to ascending; Ped articulated, stout, (0.5–) 1.5–2 ( 3) mm; Sep ovate to ovate-triangular, 1.2–2.5 ( 3.5)  1.3–2.5 mm, glabrous, entire to erose, rounded to acute or praemorse; Pet greenish-white, oblong-elliptic to ovate-elliptic, 2.5–4.8  1.5–3 mm, glabrous, entire, rounded; Fil basally briefly adnate to the petals; female Inf 1 branched, with numerous ascending to subascending primary Br (1–) 2–11 ( 17.5) cm long; floral Bra shorter or longer than the sepals, narrowly ovate to broadly ovate or oblong-ovate, 3–6  3–5.5 mm, marginally hyaline, entire to finely spinulose or erose, apiculate to spinulose; Fl divaricate to ascending; Ped articulated, stout, triquetrous, 1–2 mm; Sep triangular to ovatetriangular, 2–3 ( 4)  1.2–2.5 ( 3.5) mm, glabrous or rarely weakly lepidote, entire, rounded to acute, apiculate or praemorse; Pet narrowly ovate to triangular, 3–4.5 ( 5.5)  1.5–2.8 mm, rounded to spinulose; Ov glabrous; Fr 7–9 ( 11.5)  (3–) 4–5 ( 6) mm, glabrous, smooth or somewhat reticulate. H. pedicellata S. Watson (Proc. Amer. Acad. Arts 26: 155, 1891). Type: Mexico, Jalisco (Pringle 2970 [GH, VT]). — Distr: Mexico (Jalisco: Guadalajara); tropical deciduous forest, on rocks on dry rocky bluffs and ledges, 1000 m; known from the type only. Habit unknown, flowering to 1.2 m tall; L numerous, sheaths stramineous, suborbicular, 4.5–5  3.5–4 cm, margin spiny, L lamina 25–65

1021

 2–3 cm, glabrous adaxially, white-lepidote abaxially, Sp yellow to light brown, divaricate to antrorse, 1.5–2 mm, 0.9–1.2 cm apart; Inf glabrous; peduncle cylindrical, 6–7 mm ∅; peduncular Bra brown, narrowly triangular, 3–9.5  0.7–1 cm, attenuate, pungent, margins serrulate; male Inf unknown; female Inf 1 branched, with 15–18 ascending to divaricate primary Br 2–6 cm long; primary Bra brown, triangular, 10–14  4–5 mm, acuminate; Fl laxly disposed, ascending; Ped stout, 5–6 mm; Sep ovate-triangular, 1.5–2  1 mm, acute; Pet triangular, 3–3.3  2 mm, acute; Fr brown, 9–12  5 mm; Se brown, 5 mm. H. podantha Mez (in A. & C. de Candolle, Monogr. Phan. 9: 549–550, 1896). Type [neo]: Mexico, Hidalgo (Pringle 6932 [ENCB, B, BR, CHAPA, GH, LY, MEXU, MICH, P, UC, US, WU, Z]). — Lit: Martínez-Correa (2008); Espejo-Serna & al. (2010c). Distr: C Mexico (Aguascalientes, Guanajuato, Querétaro, Hidalgo); xerophilous scrub, on limestone hills or volcanic rocks, 1100–2650 m. Solitary or in small clumps, flowering 1.5 m tall; L numerous, sheaths light yellow to brown, ovate to suborbicular, 2.1–8.8  2.3–6.1 cm, glabrous adaxially, glabrous at the base and sparsely lepidote at the apex abaxially, serrate, L lamina green, 19–53.7  0.5–1.5 cm, sparsely lepidote on both faces, Sp antrorse, 3.1–8 mm, 0.65–3.2 cm apart; Inf 2 branched, glabrous or glabrescent; peduncle cylindrical, 0.46–1.8 cm ∅; peduncular Bra foliaceous, sparsely lepidote, serrate, 6.3–17.8 cm, the upper ones sheathing, sparsely lepidote, entire; primary Bra stramineous, widely ovate, glabrous, serrulate, 2.2–4.8 cm; male Inf with up to 46 ascending primary Br 1.6–6.6 cm long; secondary Br ascending, 1.35–5.25 cm; floral Bra brown, linear to triangular, 1.2–3.1  0.2–1 mm, glabrous, entire, acute; Fl divaricate; Ped 1.4–2.7 mm; Sep brownish, triangular to ovate, 1.7–4.3  1–2.1 mm, glabrous, entire, acute; Pet green, brownish apically, elliptic to ovate, 2.5–5.4  1.4–2.8 mm, glabrous, entire, rounded; St equal, longer than the petals; Fil 1.3–4.2 mm; Anth yellow, 0.7–2 mm; female Inf with up to 47 ascending primary Br 2.5–9 cm

1022

long; secondary Br ascending, 1.4–5.3 cm; floral Bra brown, triangular, 1.1–5.3  0.1–0.6 mm, entire, acute; Fl numerous, divaricate; Ped 1.6–5.1 mm; Sep brownish, triangular to ovate, 1.5–4  0.7–2.2 mm, glabrous, entire, acute; Pet green, triangular, 1.9–4.7  1.4–2.7 mm, glabrous, entire, acute; Ov green, ovoid, 3.8–7  1.2–2.4 mm; Fr brownish, 7–15  4–6.5 mm; Se reddish-brown, 6–7 mm. H. pretiosa Espejo & López-Ferrari (Acta Bot. Mex. 83: 50–53, ills., 2008). Type: Mexico, Guanajuato (López-Ferrari & al. 3268 [UAMIZ, IEB]). — Distr: Mexico (Guajanuato: Xichú); cactus scrub, slopes and cliffs, on igneous rocks, 800–1400 m. Caespitose, flowering to 1.7 m tall; Ros with 2 or more conspicuous narrowly cylindrical stolons; L numerous, sheaths yellowish to light brown, widely ovate to square, 4–5.5  5 cm, glabrous, lustrous, L lamina 30–50  2.2–3.5 cm, glabrous adaxially, densely brownlepidote abaxially, Sp green to brown, antrorse, 2.5–5 mm, 0.5–1.5 cm apart; Inf seemingly lateral, simple to 1 branched, procumbent; male Inf with 4–9 divaricate primary Br 5–22 cm long; floral Bra rose to rose-brownish, ovate to triangular-ovate, 6–8  3–4.5 mm, with capitate gland-hairs, acuminate, margin slightly erose; Fl divaricate to reclinate, sessile to subsessile; Sep rose to rose-brownish, oblong to lanceolate-oblong, connate at the base forming a tube, 5.5–7 mm, limb 6–8.5  2.5–3.5 mm, with capitate glandhairs, acute and shortly apiculate; Pet rose, elliptic, 6–8  3.5–4 mm, with capitate gland hairs, rounded; St unequal, united at the base with the perigone tube; Fil 2–5 mm; Anth dark purple, 1–1.5 mm; female Inf simple or with 2–7 divaricate primary Br 8–19 cm long; floral Bra rose to rose-brownish, triangular to triangular-ovate, 8–13  4–4.5 mm, with capitate gland-hairs, acuminate, margin slightly and irregularly erose; Fl sessile to subsessile; Sep rose to rose-brownish, narrowly triangular, 7–11  4–5 mm, with capitate gland-hairs, acute and shortly apiculate, margin lightly erose; Pet rose, elliptic, 8–9  3–5 mm, with capitate gland-hairs, entire, rounded and shortly apiculate; Ov ovoid to

A. Espejo-Serna et al.

narrowly ovoid, 6–10  3–4 mm, with capitate gland hairs and scarcely pedate scales; Fr light brown, 9–13  8–9 mm, lustrous, sparsely lepidote; Se light brown, 5–6  0.5 mm. H. pringlei B. L. Robinson & Greenman (Amer. J. Sci. (New York) ser. 3, 50: 167, 1895). Type: Mexico, Oaxaca (Pringle 4637 [GH, AC, B, BHUPM, BKL, BM, BR, ENCB, G, GOET, HAL, ISC, K, MANCH, MEXU, MIN, MO, MSC, NY, P, PH, S, UC, US, VT, WU, Z]). — Distr: S Mexico (C Oaxaca); shrublands, oak forest and/or tropical deciduous forest, on calcareous soils or rarely on limestone cliffs. Solitary or in small clumps, flowering to 1–2 m tall; L 15–30, fleshy, sheaths stramineous to light brown, ovate to depressed-ovate, 3.1–4.5  4–7 cm, L lamina green, 13–55  1–4 cm, white-lepidote on both faces, Sp reddish to dark brown, retrorse to antrorse or divaricate, 3–5 mm, 0.23–3.9 cm apart; Inf 1 branched, farinoselepidote to glabrescent with age; peduncle 1.5 cm ∅; peduncular Bra linear to narrowly triangular, attenuate, entire, pungent, becoming gradually shorter further up; male Inf with 24–35 ascending to divaricate primary Br 3–6 cm long; primary Bra linear to lanceolate from a triangular base, 1.5–2 cm; floral Bra white, ovate, 2.5  1.6 mm, erose, longer than the sepals; Fl polystichous, divaricate, subsessile; Ped inconspicuous, lepidote; Sep whitish to light brown, ovate-triangular, 1.7–2  1.2–1.3 mm, margin scarious, erose, somewhat farinose-lepidote, rounded; Pet white, oblong to elliptic-oblong, 2.9–3  1–2 mm, obtuse; St equal, shorter than the petals; Fil 2.5 mm; Anth yellow, 0.8–1 mm, apiculate; female Inf with 20–30 ascending to divaricate primary Br 3–17 cm long; primary Bra light brown, triangular, 1–3 cm, erose, attenuate; floral Bra brown, ovate to elliptic, 2–3 mm, farinoselepidote, erose-spiny, acute to acuminate; Fl 3–3.5  1–1.7 mm; Ped stout, to 2 mm, sulcate; Sep brown, ovate, 2.4  1.3 mm, acuminate; Pet green, triangular, 3.2  1.2 mm; Ov light brown to rose, ellipsoid, 2.7–3.5  1.2 mm; Fr brown to light brown, 6–7  3–4.5 mm; Se brown, 4–4.5 mm.

Hechtia BROMELIACEAE

H. pumila Burt-Utley & Utley (Syst. Bot. 13 (2): 279–281, ills., 1988). Type: Mexico, Guerrero (Burt-Utley & Utley 7908 [MEXU, B, BH, BM, CAS, DUKE, F, GH, K, MEXU, MICH, MO, NY, TEX, UC, US, WIS]). — Distr: Mexico (Guerrero: Chilpancingo, Tepecoacuilco); thorn scrub with palms and tropical deciduous forest, rocky places on steep limestone hills, 600–1650 m. Usually forming small clumps of 2 or more rosettes, flowering 58–151 cm tall; Ros 11–34 cm ∅; L few, spreading, sheaths stramineous, broadly ovate to reniform, 1.3–2.2  2.3–4.6 cm, margins finely spiny, glabrous and lustrous on both faces, becoming densely lepidote distally, L lamina 9–25  1.3–2 cm, appressedly cinereous-lepidote on both faces, Sp red-brown or rarely stramineous, antrorse or occasionally retrorse, 1–2 mm, 0.45–1 cm apart; Inf 1 branched, glabrous; peduncle cylindrical, 3.5–8 mm ∅; peduncular Bra 0.7–7.5 cm, with ovate or broadly ovate glabrous sheaths and linear lepidote lamina, entire; male Inf with 12 subascending primary Br (1.5–) 2–4 cm long, appearing somewhat winged when dry; floral Bra ovate or ovate-triangular, 1–2.5  0.8–2 mm, glabrescent, attenuate-acuminate, erose, esp. at the apex, exceeding the pedicels; Fl sessile to subsessile; Ped articulated, stout, to 1 mm; Sep ovate or ovate-triangular, 1.1–1.8  0.7–1.3 mm, praemorse or occasionally acuminate, entire and occasionally hyaline, glabrous; Pet elliptic, 2–3.5  (1–) 1.6–2 mm, rounded, glabrous; St subequal, shorter than the petals; Fil 2.9–3 mm; Anth yellow, 1 mm; female Inf with 12–17 subcapitate primary Br 0.3–2.5 cm long; floral Bra exceeding the pedicels, triangular to ovate-triangular, 1.1–2.5  0.6–2.6 mm, glabrous, attenuateacuminate, erose distally, occasionally hyaline; Fl sessile to subsessile; Sep dark red-brown, ovate to ovate-triangular, 1.3–2.5  1.3–2 mm, glabrous, acute or praemorse, margins often hyaline; Pet triangular to narrowly ovate-triangular, 2.6–4.3  1.3–2.3 mm, rounded to acute; Fr dark red-brown, 5.5–11  3.1–5.5 mm, glabrous. H. purhepecha I. García & al. (Acta Bot. Mex. 107: 10–14, ills., 2014). Type: Mexico, Michoacán (Espejo & al. 7547 [UAMIZ, CIMI, IEB]). — Distr: Mexico (Michoacán: Los Reyes, Peribán);

1023

(sub-) deciduous tropical forest, slopes and cliffs, on rocks, 900–1000 m. Caespitose, stoloniferous, flowering to 1.7 m tall; Ros 30–50 cm ∅, at the tip of narrowly cylindrical stolons; L 15–25, sheaths white on both faces, depressed-ovate, 3–4  5–6.5 cm, glabrous, lustrous, L lamina 11–50  2.6–4.5 cm, light green with a purple stripe along the margins, glabrous adaxially, densely white-lepidote abaxially, Sp yellow to light brown, 2–4.5 mm, 0.8–1.3 cm apart; Inf seemingly lateral, 1–2 branched, erect, glabrous; male Inf 2 branched with 25–30 divaricate to ascending primary Br 4–28 cm long; secondary Br, when present, 2–4 cm; floral Bra brown-purplish to dark brown with purplish dots, triangular-ovate, 1.5–1.9  0.9–1.2 mm, erose, glabrous, apiculate; Fl divaricate, sessile; Sep brown-purplish to dark brown with purplish dots, triangular-ovate, 3.1–3.4  3–3.5 mm, entire, glabrous, rounded; Pet white, elliptic, 4.8–5  3.3–3.5 mm, rounded; St subequal; Fil 3.9–4. 2 mm; Anth yellow, 2–2.5 mm, apiculate; female Inf 1 branched with 13–20 ascending primary Br 10–21 cm long; floral Bra green with rose apex and purplish dots, ovatetriangular, 1.4–2  0.5–0.7 mm, glabrous, minutely crenate, acute; Fl sessile; Sep green with rose apex and purplish dots, ovate-triangular, 3.5–4.5  2.6–3.6 mm, entire, glabrous, rounded; Pet whitegreenish, narrowly triangular-elliptic, 5.8–6  3.2–3.8 mm, rounded; Ov narrowly ovoid, 5–6  2.6–3 mm; Fr brown-reddish to dark brown, 10–12  4–6 mm, lustrous; Se reddish, 8–9 mm. H. purpusii Brandegee (Univ. Calif. Publ. Bot. 7: 325, 1920). Type: Mexico, Vera Cruz (Purpus 8420 [UC, NY, US]). — Distr: Mexico (Veracruz: Coatepec, Jalcomulco, Totutla, Puente Nacional, Tenampa); tropical deciduous forest, on limestone cliffs, 30–1200 m. – Fig. 7. Incl. Hechtia lindmanioides B. L. Robinson (1937). Caespitose, pendulous, flowering to 1 m tall; Ros short to long caulescent, stems rhizomatous, cylindrical, 20–80  2–3 cm ∅, covered by the persistent leaf sheaths; Ros somewhat asymmetrical, 20–50 cm ∅; L 10–15, recurved and pendent, brittle, sheaths yellowish, oblong to ovate,

1024

5–7.5  3.5–5 cm, lustrous, white-lepidote distally on both faces, minutely serrate, L lamina yellowish-green, 20–50  1–1.5 cm, densely white-lepidote abaxially, long-attenuate, minutely serrate; Inf erect to pendent; male Inf 3 branched, with numerous primary Br 10–28 cm long; peduncle cylindrical, 20–32 cm, 4–5 mm ∅; peduncular Bra yellowish, narrowly triangular to linear, 2–10 cm  2–5 mm, densely whitelepidote, much longer than the internodes, decreasing in size further up, deciduous; floral Bra filiform, 1–1.5 mm, glabrous, deciduous, shorter than the pedicel; Fl polystichous, divaricate to descending; Ped filiform, 3–5 mm; Sep brownish, triangular, 1.5–2  1 mm, acute; Pet white, broadly elliptic, 4–5  1.5–2 mm, rounded; St shorter than the petals; Fil 2.5–3 mm; Anth yellow, 1 mm; female Inf 3 branched, with numerous primary Br 10–16 cm long; peduncle cylindrical, 25–50 cm, 3 mm ∅; peduncular Bra yellowish, narrowly triangular to linear, 5–13  0.2–0.5 cm, densely white-lepidote, much longer than the internodes, decreasing in size upwards; floral Bra filiform, inconspicuous, to 2.5 mm; Fl polystichous, divaricate to descending; Ped filiform, 1.5–2.5 mm; Sep brown-greenish, triangular to narrowly triangular, 0.9  0.8 mm, acute; Pet white to rose-whitish, elliptic to oblong, Fig. 7 Hechtia purpusii. (Copyright: A. EspejoSerna)

A. Espejo-Serna et al.

4–6  3 mm, rounded; Ov dark brown, ovoid, 4–5  2–3 mm; Fr light brown, 5–10  2.6–3 mm, divaricate to reflexed; Se brown, 2.5–3 mm. H. reflexa L. B. Smith (Contr. Gray Herb. 117: 18–19, t. 1, figs. 54–55, 1937). Type: Mexico, Michoacán (Langlassé 238 [US, GH, K, P]). — Distr: S Mexico (Michoacán: Aquila, Arteaga; Guerrero: José Azueta); tropical deciduous and semideciduous forest, on karstic limestone rocks, 50–650 m. Incl. Hechtia michoacana Burt-Utley & al. (2011). Usually caespitose, flowering (0.55–) 1–1.4 m tall; Ros 30–90 cm ∅; L numerous, sheaths reniform to transversally elliptic, 2.5–4.5  (3.5–) 4.4–5.7 ( 8.5) cm, marginally spiny and floccose, both faces glabrous and lustrous becoming lepidote distally, L lamina green to maroon, straight to falcate, 24–57  1.2–2.4 ( 2.9) cm, densely lepidote adaxially and lepidote abaxially, Sp red-brown, antrorse or occasionally divaricate or retrorse, (2–) 2.5–4.5 ( 5) mm, 0.7–2.5 cm apart; Inf glabrous; peduncle 3.5–7 mm ∅; peduncular Bra foliaceous, ascending, 1.1–26 cm, lepidote, becoming progressively reduced upwards, ovate-triangular with linear lamina; male Inf 2 branched, with numerous ascending to

Hechtia BROMELIACEAE

subascending reflexed primary Br 6–22.5 cm long; floral Bra adnate to and often exceeding the pedicels, ovate, 1.2–2.5  0.7–1.2 mm, finely serrulate distally, apiculate to attenuate-acuminate, glabrous; Fl divaricate to subascending; Ped slender, 0.5–2.2 mm; Sep deep to light pink or green with brownish maculation, ovate, 0.9–1.5  0.5–1.1 mm, entire to serrulate distally, incised to praemorse or apiculate distally; Pet white, cream or light to dark pink, elliptic, (1.6–) 3–3.5  1–1.7 mm, rounded; St longer than the sepals; female Inf 1 branched, with 20–22 arcuate to ascending or subascending primary Br 6.5–22.5 cm long, rarely with 1–2 short basal secondary branches; floral Bra adnate to the pedicels, shorter than the sepals, narrowly ovate to ovate, 1.5–2.5  0.8–1.8 mm, glabrous, entire to finely erose, apiculate to attenuate-acuminate or praemorse; Fl ascending to subascending; Ped slender, 1.5–3.5 ( 4.5) mm; Sep deep to light pink or green with maroon maculation, triangular to ovate-triangular, 1.2–2  0.7–1.1 mm, glabrous, entire to serrulate distally, acutepraemorse to apiculate; Pet white, cream or light to dark pink, narrowly triangular, (2.3–) 3–4.5  1–1.2 mm, glabrous, rounded; Ov ovoid, 2.8–3  1 mm; Fr green suffused with red when immature but becoming yellow-brown and reticulate at maturity, (5–) 6–9  2.5–4.5 mm, glabrous, smooth. H. reticulata L. B. Smith (Contr. Gray Herb. 117: 17–18, t. 1, fig. 46, 1937). Type: Mexico, Colima (Palmer 1352 [GH, F, UC, US]). — Distr: Mexico (Jalisco: La Huerta; Colima: Colima, Manzanillo; Michoacán: Arteaga, La Huacana); tropical deciduous forest, 40–750 m. Caespitose, flowering to 1.2 m tall; Ros 60–140 cm ∅; L numerous, densely rosulate, spreading, sheaths stramineous to light brown, orbicular to depressed-ovate, 5.2–6.8  6–7.6 cm, glabrous, lustrous, margins irregularly spiny, L lamina green to reddish adaxially, 30–70  3–4 cm, Sp brown to yellowish, antrorse, 3–4.5 mm, 0.7–1.5 cm apart; Inf barely 2 branched, glabrous; peduncle 1 cm ∅; peduncular Bra erect, the lower exceeding the internodes, progressively smaller distally, pungent;

1025

male Inf unknown; female Inf with 15–20 ascending primary Br 10–15.5 cm long; primary Bra brown, triangular, 5–15 mm, filiformacuminate; floral Bra ovate, 3.5  1 mm, acuminate, equal or barely longer or shorter than the pedicels; Fl divaricate, laxly disposed; Ped stout, conical, 2–4 mm, glabrous, sulcate; Sep green, ovate, 2.5–4  1.5 mm, margins hyaline, erose, rounded; Pet white, narrowly triangular, 3.5–4  1.5–2 mm, acute; Ov superior, green, ovoid, 3  1 mm; Fr brown, 7.5–10  3–5 mm, irregularly reticulate; Se brown, 4.8–5.4 mm. H. rosea E. Morren ex Baker (Handb. Bromel., 140, 1889). Type: Mexico (Anonymus s.n. in Hort. Liège s.n. [LG [Morren drawing], GH [photo]]). — Distr: Mexico (Oaxaca, Chiapas); tropical deciduous and semideciduous forests and thorn scrub, on the ground or on rocks, 10–600 m. I: Roguenant & al. (2016: 150–151). Incl. Hechtia roezlii hort. ex Baker (1889) (nom. inval., Art. 34.1c); incl. Dyckia desmetiana Baker (1894)  Hechtia desmetiana (Baker) Mez (1896); incl. Bromelia desmetiana hort. ex Baker (1894) (nom. inval., Art. 34.1c); incl. Hechtia meziana L. B. Smith (1935); incl. Hechtia macdougallii L. B. Smith (1969). Caespitose, flowering to 1.5 m tall; L rather few, rosulate, sheaths stramineous to light brown, suborbicular to transversally oblong, 4.5–6  5.5–7 cm, densely brown-lepidote on both faces, glabrous to the base, lustrous, margins spiny to the apex, L lamina dark green to purple, 15–90  3.6–4.2 cm, fleshy, densely lepidote on both faces, Sp divaricate to antrorse, coarse, 5–7 mm, 0.7–2.7 cm apart; Inf reddish, 1 branched, glabrescent; peduncle 1 cm ∅; peduncular Bra broadly ovate, the lower ones with a long linear lamina exceeding the internodes, the upper ones merely acuminate; male Inf with (5–) 13–25 ascending primary Br 5.5–22 cm long; Fl divaricate to ascending, laxly disposed, subsessile; floral Bra rose, ovate to oblong-ovate, 3.8–4.7  1.8–2 mm, acuminate, shorter than the sepals; Sep rose, broadly ovate, 5–5.6  3.3–3.8 mm, entire, acuminate to apiculate, glabrous; Pet rose, basally very shortly connate, elliptic, 9.5–10  3.7–5 mm, glabrous, obtuse; St subequal,

1026

shorter than the petals; Fil 7.6–8.4 mm; Anth yellow, 3 mm, shortly apiculate; female Inf with 10–25 ascending primary Br 5.8–15 cm long; floral Bra triangular, 3.5–4.5  3.3 mm, acute to acuminate, longer than the sepals; Fl divaricate, pedicellate; Ped 1 mm; Sep rose, triangular, 2.4–4.6  2.65–3.4 mm, glabrous, acuminate; Pet rose, narrowly triangular, 8–10  3.7–4.2 mm, glabrous, acute; Ov reddish, ellipsoid, 5–5.7  1.7–2.7 mm; Sty 0.9 mm; Fr green to light brown, 12–23  5.6–9 mm, smooth; Se reddish to brown, 5.5–10 mm. H. roseana L. B. Smith (Contr. Gray Herb. 117: 17, t. 1, figs. 41–42, 1937). Type: Mexico, Puebla (Rose & al. 9970 [US]). — Distr: Mexico (Puebla, Oaxaca); arid thorn scrub and cactus scrub, on limestone soils, 1500–1700 m. – Fig. 8. Caespitose, flowering to 2.9 m tall; Ros to 60 cm ∅ with 1 to several conspicuous, stout stolons to 1 m long; L 40–50, sheaths yellow adaxially, light brown abaxially, transversally elliptic, 3.5–4.5  6 cm, glabrous and lustrous on both faces, lepidote abaxially at the distal end, L lamina yellow-green to light green with red spots at the spine bases, 29–43  2.5–4 cm, glabrous adaxially, finely appressedly white-lepidote abaxially, Sp reddish-brown, divaricate, some Fig. 8 Hechtia roseana. (Copyright: A. EspejoSerna)

A. Espejo-Serna et al.

antrorse or retrorse, 4–6 mm, 1.3–3.8 cm apart; Inf 2 (rarely 1) branched, glabrous; peduncle 1.5 cm ∅; peduncular Bra light brown, linear to narrowly triangular, the lower spiny, the upper entire, gradually becoming shorter upwards; male Inf with >60 primary branches 7.5–23 cm long, the basal primary Br with 2 basal lateral secondary branches; floral Bra stramineous to light brown, ovate, 5.5–7.5  3.5–4.2 mm, erose and hyaline at the margins, much longer than the sepals; Fl divaricate to ascending, laxly disposed; Ped stout, 1–1.6 mm; Sep stramineous to light brown, widely ovate, 3.2–3.4  3.1–3.4 mm, entire, glabrous, obtuse; Pet white, elliptic, 5.3–5.8  2.5–2.6 mm, entire, glabrous, obtuse; St subequal, barely longer than the petals; Fil 2.8–4.2 mm; Anth greenish, 1.5–2 mm; female Inf with >50 primary Br 9–25 cm long, basal primary branches with 2 basal lateral secondary branches; primary Bra triangular-ovate, long acuminate, 1.5–3 cm; floral Bra triangular-ovate, 4.5–6.2 mm, acute, longer than the sepals; Fl ascending, very laxly disposed, sometimes appearing subverticillate, sessile to subsessile; Sep greenish to light brown, triangular-ovate, 3.2–3.7  2–2.2 mm, entire, glabrous, margin hyaline, acuminate; Pet white, triangular to ovate-triangular, 5–5.2  2.5–2.9 mm, glabrous,

Hechtia BROMELIACEAE

entire, acute to acuminate; Ov wholly superior, greenish to rose, ellipsoid, 4.9–5.1  2.4 mm; Fr brown, 10–12  5–5.5 mm, slightly lustrous, very faintly rugose; Se reddish-brown, 5.5–6  1 mm. H. rubicunda López-Ferrari & Espejo (Acta Bot. Mex. 107: 154–158, ills., 2014). Type: Mexico, Oaxaca (Martínez-Correa & al. 116 [UAMIZ, IEB]). — Distr: Mexico (Oaxaca: Yautepec); tropical deciduous forest, on rocks on igneous cliffs, 500–850 m. Solitary, flowering to 1.5 m tall; Ros 50–75 cm ∅; L 15–30, sheaths dark brown, glabrous and lustrous on both faces, depressed ovate, 2.7–3.7  5–6.5 cm, margins minutely spinose, L lamina red-yellowish, narrowly triangular, 28–60  3.5–6.5 cm, glabrous adaxially, densely appressedlepidote abaxially, Sp yellowish to light brown, divaricate to antrorse except in the basal portion where they are retrorse, 2–4.5 mm, 0.9–1.2 cm apart; Inf erect to arcuate; male Inf 2 branched with 45–50 primary branches; peduncular Bra foliaceous, straw-coloured, sheaths triangular, 0.8–1.2 cm, lamina linear, 1.7–15.5  0.15–0.4 cm, lepidote on both faces, margins entire, pungent; primary Br 6.6–30 cm, stalked; secondary Br ascending, 2–4.5 cm; floral Bra straw-coloured, ovate-triangular, 1–1.5  0.5–0.7 mm, erose, glabrous, attenuate; Fl polystichous, divaricate; Ped 2 mm; Sep light brown, triangular, 1.3–1.7  1–1.3 mm, acute; Pet white, elliptic, 2.7–4  2–2.3 mm, acute; St subequal; Fil 3–3.2 mm; Anth yellow, 1.5 mm; female Inf 1 branched with 35–40 branches, occasionally the lower branches with a pair of secondary basal branches; peduncular Bra foliaceous, straw-coloured, sheaths triangular, 0.8–1.2 cm, lamina linear, 1.7–15.5  0.15–0.4 cm, lepidote on both faces, margins entire, pungent; Br divaricate to ascending, 4.5–40 cm; floral Bra straw-coloured, ovatetriangular, 1–1.5  0.5–0.7 mm, erose, acuminate, glabrous, longer than the pedicels; Ped 1 mm; Fl polystichous, reflexed at anthesis; Sep green with rose apex, triangular, 1.5–1.7  1.2–1.4 mm, acute; Pet white, triangular, 3.8–4  1.8–2 mm, attenuate and acuminate; Ov rose-coloured, ovoid, 3.8–4 mm; Fr light green, ovoid, 10–12  4–4.5 mm, lustrous; Se reddish.

1027

H. santanae I. Ramírez & P. Carrillo (Phytotaxa 266(4): 262–266, ills., 2016). Type: Mexico, Jalisco (Ramírez & al. 1940 [CICY, IBUG, MEXU, US, ZEA]). — Distr: Mexico (S Jalisco); tropical deciduous forest, 780–920 m. Caespitose, flowering 1.1–1.4 m tall; Ros globose, to 30  20–40 cm ∅; L 30–40 per rosette, sheaths pale yellow with light brown spots when dry, broadly ovate to broadly oblong, 3–4  3.5–4.8 cm, finely spiny, glabrous in the middle area but with white-lepidote margins, L lamina narrowly triangular, attenuate, 12–34  1.7–2.8 cm, succulent, green, densely to sparsely silverylepidote adaxially, densely white-lepidote abaxially, Sp brownish, antrorse, 1–3 mm, 0.4–1.7 cm apart; Inf 1 branched; male Inf with 40–45 Br 5.5–13.5 cm long; floral Bra purple with a brown apex, oblong-triangular, 3.4–3.9  2.2–2.4 mm, acuminate, erose and hyaline, glabrous, shorter than the sepals; Fl erect, 5–5.5  3.8–4.2 mm, subsessile; Sep purple with a brown apex, ovate, 2.3–2.7  1.8–2.3 mm, entire, glabrous, acute; Pet white, elliptic, 4.3–4.8  3–3.2 mm, entire, rounded; St unequal, shorter than the petals; Fil 3.8–4.4 mm; Anth reddish, 2–2.2 mm; female Inf with 30–70 Br 4–9.5 cm long; floral Bra purple with a brown apex, oblong, 4–4.2  3–3.2 mm, acuminate, erose and hyaline, glabrous, longer than the sepals; Fl erect, 5.7–6  3–4 mm, pedicellate; Ped 2.8–3.2 mm; Sep purple with a brown apex, triangular, 2.4–2.9  2–2.2 mm, entire, glabrous, acute; Pet whitepurplish, oblong, 4.7–5  2–2.2 mm, entire, acute; Ov purple to reddish, oblong to ellipsoid, 3–3.2  1.7–2 mm ∅; Fr brown, 6–11  3–4 mm ∅; Se 5–6.8 mm. Material of this species was previously identified as H. laevis. H. schottii Baker ex Hemsley (Biol. Centr.Amer., Bot. 3: 318, 1884). Type: Mexico, Yucatán (Schott 645 [BM, F]). — Distr: Mexico (Yucatán: Opichén; Campeche: Calakmul); tropical deciduous forest, on the ground or on limestone rocks, 30–260 m. Forming extensive colonies, flowering 2–2.5 m tall, rhizomatous; L numerous, sheaths stramineous to light brown, suborbicular, 4–5  4–5 cm,

1028

glabrous, lustrous, margins spiny, L lamina green, 50–100  3.5–4.5 cm, glabrous adaxially, finely cinereous-lepidote abaxially, Sp mostly divaricate, some retrorse or antrorse, 5–6 mm, 1.4–3 cm apart; Inf seemingly lateral, 2–3 branched, densely and finely white-flocculose throughout; peduncle 1 cm ∅; peduncular Bra light brown, triangular, 4.5–5 cm, acuminate; male Inf with numerous ascending to erect primary Br 4–20 cm long; secondary Br 1.5–8 cm; primary Bra light to dark brown, ovate, 1–6  0.6–1.5 cm, entire, acuminate; floral Bra brown, ovate to broadly ovate, 4–5 mm, acute, apiculate, about equalling the sepals; Fl ascending, subverticillate and densely to laxly disposed, subsessile; Sep brown, ovate, 2.8–3  2.2–2.4 mm, floccose-lepidote, margins hyaline, erose, obtuse; Pet basally shortly connate, white, elliptic, 4.4–4.8  2.7–2.8 mm, glabrous, entire, obtuse; St unequal, barely shorter than the petals; Fil 2.3–3.9 mm; Anth greenish, 1.7 mm, apiculate; female Inf with numerous ascending to erect primary Br 10–20 cm long; secondary Br 2–8 cm; primary Bra light to dark brown, ovate, 1–6  0.6–1.5 cm, entire, acuminate; floral Bra brown, ovate to broadly ovate, 4–5 mm, acute, apiculate, about equalling the sepals; Fl ascending, subverticillate and densely to laxly disposed, subsessile; Sep brown, ovate, 4–4.4  3–3.3 mm, floccose-lepidote, margin hyaline, erose, obtuse to acute; Pet basally shortly connate, white, triangular to ovate, 5–5.2  3–3.3 mm, glabrous, entire, acute; Ov rose, ellipsoid, 4.5  2.3 mm, white-lepidote; Fr light brown, 8–9  4.5–4.7 mm, white-lepidote to glabrescent, acute; Se reddishbrown, 3.5–6 mm. H. sphaeroblasta B. L. Robinson (Proc. Amer. Acad. Arts 35: 323, 1900). Type: Mexico, Guerrero (Nelson 2044 [GH, US]). — Distr: Mexico (Puebla, Guerrero, Oaxaca); tropical deciduous forest, on the ground or rarely on rocks, 800–1900 m. Caespitose, flowering 2.5–3.5 m tall; L numerous, sheaths stramineous, orbicular to ovate, 5.6–8  5.8–11 cm, lepidote adaxially, glabrous and lustrous abaxially, margins spiny, L lamina green, 25–110  3–5.5 cm, inconspicuously

A. Espejo-Serna et al.

finely white-lepidote on both faces, Sp brownish to reddish, antrorse, 2.5–5 mm, 1.5–3.2 cm apart; Inf seemingly lateral, 2 branched, glabrous; peduncle 2.5–2.8 cm ∅; peduncular Bra green to light brown, foliaceous, linear, 24–50  0.5–0.7 cm, margin spiny, pungent; male Inf with numerous ascending flattened primary Br 28–40 cm long and with 2–15 divaricate secondary Br 3–12 cm long; primary Bra stramineous, ovate-triangular, 1.5–2 cm, attenuate; floral Bra scarious, triangular, 1–1.5 mm; Fl divaricate, single to subverticillate, subsessile; Sep light brown, ovate, 2–2.3  1.6 mm, glabrous, entire, obtuse; Pet white, suborbicular to elliptic, 3–3.4  2–2.2 mm, glabrous, entire, obtuse; St subequal, barely shorter than the petals; Fil 2.5–2.7 mm; Anth yellowish, 1 mm; female Inf with numerous ascending flattened primary Br 22–42 cm long, with 8–15 ascending secondary Br 2.5–10 cm long; primary Bra stramineous, ovate-triangular, 1.2–2.5 cm, attenuate; floral Bra white, ovatetriangular, 1.5–1.8 mm, erose, apiculate, shorter than the sepals; Fl ascending to divaricate, subverticillate, subsessile to shortly pedicellate; Ped stout, 0.8–1.2 mm; Sep white, ovate, 1.9–2.3  1.6 mm, glabrous, entire, obtuse; Pet white, triangular, 2.5–3  1.5 mm, glabrous, entire, acute; Ov green, ellipsoid, 2–2.5  1.2 mm, glabrous; Fr light brown, 5.8–6.8  2.5–4 mm; Se dark brown, 4.7–5 mm. H. stenopetala Klotzsch (Allg. Gartenzeitung 3: 402, 1835). Type: Mexico, Veracruz (Schiede & Deppe s.n. [B, HAL]). — Distr: Mexico (C Veracruz); tropical deciduous forest, on the ground and on rocks of lava flows, 100–700 m. Incl. Hechtia besseriana J. Verschaffelt (1874); incl. Hechtia glabra Brandegee (1920). Caespitose, flowering 0.5–2 m tall; Ros  elongated, short- to long-caulesent, stems rhizomatous, 1.6 cm ∅; L numerous, sheaths yellowish-brown, ovate to widely ovate to oblong, 3.5–6  2–4.6 cm, lustrous towards the base, brown-lepidote toward the apex, margins finely serrate and densely lepidote, L lamina green, sometimes tinged with red, 28–56 ( 70)  1.3–3.5 cm, densely lepidote on both faces, Sp divaricate to antrorse, light brown, 1.5–2.3 mm,

Hechtia BROMELIACEAE

1029

1.4–2.2 cm apart; Inf 2 branched; male Inf with numerous primary Br 10–15 cm long; peduncle 9–11 mm ∅, glabrous; peduncular Bra yellowish, narrowly triangular to linear, 6–12  0.2–1 cm, white-lepidote, entire, decreasing in size upwards; floral Bra whitish-brown, oblong to elliptic, 0.7–1.2 mm, erose, acute, equal or shorter than the pedicels; Fl polystichous, divaricate; Ped 0.8–1 mm; Sep elliptic to broadly triangular, 1.6–2.1  1–1.7 mm, rounded; Pet white, broadly elliptic to broadly oblong, 2.5–3  1.6–2 mm, rounded; St longer than the petals; Fil 2.5–3 mm; Anth green, 1.1–1.4 mm; female Inf with numerous primary Br 10–15 cm long; peduncle cylindrical, 8–13 mm ∅; peduncular Bra yellowish, narrowly triangular to linear, (2–) 11–12 ( 18)  (0.5–) 0.8–1 cm, deciduous; floral Bra oblong to triangular, 0.9–1.1  0.5 mm, rounded, equal or longer than the pedicels; Fl polystichous, divaricate; Ped obconical, 0.7–1 mm; Sep green, triangular to ovate, 1.4–2.3  1–2 mm, rounded or acute; Pet white, triangular, 2.5–3  1.3–1.6 mm, acute; Ov green, ellipsoid, 3–4  1.3–2 mm; Fr dark brown, 6–8  3–4 mm; Se brown, 2.5–2.9 mm.

acuminate, glabrous adaxially, white-lepidote abaxially, margins irregularly dentate; floral Bra light brown, linear to very narrowly triangular, 1.6–2.5 mm, shorter than the pedicels; Fl polystichous, ascending to divaricate; Ped conical, 2.5–3.5 mm, sulcate; Sep brownish, triangular, 4.8–5.3  2–2.3 mm, entire, glabrous, acute; Pet white, elliptic, 5.7–6  3–3.3 mm, entire, glabrous, rounded; St subequal, shorter than the petals; Fil 5.3–5.6 mm; Anth yellow, 2 mm; female Inf with numerous ascending primary Br 2–10 cm long; primary Bra stramineous to light brown, ovate-triangular, 2.5–4.6  1.4–1.6 cm, glabrous adaxially, white-lepidote abaxially, margins irregularly spiny-dentate, acuminate; floral Bra light brown, narrowly triangular to linear, 2–2.5 mm, shorter than the pedicels; Fl densely disposed to agglomerated, ascending; Ped stout, conical, 2.5–3.5 mm, sulcate; Sep brownish, elliptic to ovate-triangular, 3.5 mm, erose, hyaline, rounded; Pet white, triangular, 4.5–5.2  2.3–2.5 mm, entire, glabrous, acute; Ov green, ellipsoid, 4 mm; Fr green to light brown, 10–14  7–9 mm; Se brown-reddish, 4–4.8 mm.

H. subalata L. B. Smith (Contr. Gray Herb. 117: 15–16, t. 1, fig. 29, 1937). Type: Mexico, Durango (Rose 3467 [US]). — Distr: Mexico (Durango, Zacatecas, Nayarit, Jalisco); tropical shrubland, on the ground or on rocks, 900–1750 m. Caespitose, flowering to 1.5 m tall; L numerous, sheaths stramineous, square to transversely oblong, 4–4.5  4.5–4.7 cm, glabrous, lustrous, erose to spiny at the margins, L lamina green to glaucous, 20–36  1.5–3 cm, densely appressedly white-lepidote on both faces, long attenuate and coiled in the apical portion, margins repand, Sp light brown, divaricate to retrorse or some antrorse, 2.2–4.5 mm, 0.65–1.3 cm apart; Inf 1 branched, white-lepidote, glabrescent with age; peduncle 1.3–1.5 cm ∅; peduncular Bra light brown, amplexicaul, ovate to oblong-ovate, with a short narrowly triangular lamina, 5–5.6  1.5–2.3 cm, irregularly dentate, white-lepidote abaxially, margins irregularly dentate; male Inf with numerous ascending to erect primary Br 4–8 cm long; primary Bra stramineous to light brown, ovate to lanceolate, 4–6.6  1.6–2 cm,

H. tehuacana B. L. Robinson (Proc. Boston Soc. Nat. Hist. 31: 265–266, 1904). Type: Mexico, Puebla (Pringle 8578 [GH, B, BM, ENCB, G, GOET, K, MEXU, NY, P, UC, US, VT, Z]). — Distr: Mexico (Puebla, Oaxaca); xerophytic shrubland, calcareous hills, 1650–1750 m. Incl. Hechtia pueblensis Burt-Utley & al. (2011). Solitary or in small groups, flowering to 2.2 m tall; Ros 50 cm ∅; L numerous, sheaths yellow adaxially, brownish abaxially, square to ovate, 4.4–7  4.3–7.2 cm, glabrous, distally densely lepidote on both faces, serrate, L lamina green, 25–46  0.8–1.8 cm, sparsely lepidote to lepidote on both faces, Sp light brown to reddish, divaricate to antrorse or sometimes retrorse, 3.8–8.9 mm, 0.53–2.5 cm apart; Inf 2 branched, glabrescent; peduncle cylindrical, 1–2 cm ∅; basal peduncular Bra foliaceous, brown, 7.3–16.5 cm, sparsely lepidote, serrate, upper peduncular Bra caudate, entire; male Inf with up to 86 ascending primary Br 3.6–8 cm long; secondary Br basal, ascending, 1.9–4.9 cm; primary Bra ovate-triangular, sparsely

1030

A. Espejo-Serna et al.

lepidote, serrate, 2–7.3 cm; floral Bra brownish, triangular, 2–3.5  0.2–1.2 mm, entire, glabrous, acuminate; Fl divaricate, densely disposed; Ped stout, 1–2.7 mm, glabrous; Sep brownish, ovate, 1.6–4.3  0.6–2.1 mm, glabrous, entire, acute; Pet green, oblong, 3.2–4.7  1.7–3.2 mm glabrous, entire, rounded; St equal, longer than the petals; Fil 2.3–4.2 mm; Anth yellow, 0.5–1.9 mm; female Inf with up to 44 ascending primary Br 5.1–6 cm long; secondary Br ascending, 1.3–2.4 cm; primary Bra brown, ovate-triangular, 3.1–4.6 cm, sparsely lepidote, serrate; floral Bra brownish, triangular, 1.5–2.8  0.4–1.2 mm, entire, glabrous, acute; Fl divaricate, densely disposed; Ped stout, 2.8–4.4 mm; Sep brownish, ovate, 1.3–2.6  0.6–1.2 mm, glabrous, entire, acute; Pet green, triangular, 2.1–3.6  0.9–1.4 mm, glabrous, entire, acute; Ov green, ovoid, 2 mm; Fr brownish, 7–10  4.4–7.5 mm ∅; Se brownish, 3–3.8 mm.

or nearly equalling the sepals; Fl ascending, subverticillate, sessile; Sep green, ovate to elliptic, 4.5–6  2.5–4 mm, margin hyaline, densely lepidote, carinate, obtuse; Pet white, elliptic, connate basally, 5–8.5  2.2–3.5 mm, obtuse, slightly lepidote abaxially; St unequal, shorter than the petals; Fil 3–6 mm; Anth green, sagittiform, 1.5–2 mm; female Inf with 11–17 ascending to appressed primary Br 3–23 cm long; primary Bra 1.2–1.5 cm, attenuate, acute; Fl ascending, subverticillate, sessile to subsessile; floral Bra brownish, triangular-ovate, 4–6 mm, shorter than the sepals; Sep brownish, ovate to broadly ovate, 4.5–7  4 mm; Pet white, elliptic, 7.5–10  4.5–5 mm, glabrous, entire, rounded; Ov green to brownish, ovoid, densely lepidote; Fr brown, 8–12  5–6 mm, lepidote, glabrescent. Vernacular names: “Texas False Agave”, “Guapilla de Texas”.

H. texensis S. Watson (Proc. Amer. Acad. Arts 20: 374–375, 1885). Type: USA, Texas (Havard 85 [GH, GH]). — Distr: USA (Texas), Mexico (Coahuila, Nuevo León, Durango, Zacatecaas); xerophytic scrub and rosette-rich shrubland, limestone hills and ledges, on the ground or on rocks, 900–1850 m. Incl. Hechtia scariosa L. B. Smith (1937); incl. Hechtia zacatecae L. B. Smith (1937). Caespitose, flowering 0.55–2 m tall; L numerous, sheaths light brown, broadly ovate to suborbicular, 2.5–5.5  3–5.5 cm, glabrous and lustrous on both faces, densely serrulate to serrulate, L lamina green or green with red to bright rose stripes, 20–45  1–5 cm, glabrous and lustrous adaxially, densely appressedly white-lepidote abaxially, Sp yellow to dark castaneous, antrorse to divaricate or some retrorse, 4–6 mm, 1–2.7 cm apart; Inf seemingly lateral, lepidote, 1 to rarely 2 branched; peduncle strongly flattened at the base, 3.5–7.5 mm ∅; peduncular Bra stramineous to dark brown, ovate to triangular, 2.7–5  1 cm, acuminate, shorter than the internodes; male Inf with 7–18 ascending slender primary Br 4–26 cm long; primary Bra stramineous, ovate-triangular, 3.7–12 mm, attenuate; secondary Br 1.7–4 cm; floral Bra rose with a broad hyaline margin, suborbicular, 4.5–7 mm, erose, apiculate, shorter than

H. tillandsioides (André) L. B. Smith (Contr. US Nation. Herb. 29: 431, 1951). Type: Mexico (Anonymus s.n. [K]). — Distr: Mexico (San Luis Potosí, Querétaro, México, Guerrero); tropical deciduous forest and shrublands, on perpendicular limestone cliffs and ledges, 600–1800 m. I: Roguenant & al. (2016: 145, as H. caerulea).  Bakeria tillandsioides André (1889) (incorrect name, Art. 11.4)  Bakerantha tillandsioides (André) L. B. Smith (1934); incl. Vriesea glaucophylla hort. ex Mez (1934) (nom. illeg., Art. 53.1); incl. Niveophyllum caeruleum Matuda (1965)  Hechtia caerulea (Matuda) L. B. Smith (1972); incl. Hechtia integerrima M. B. Foster (1968). Solitary to caespitose, pendulous, flowering to 1.6 m tall, shortly caulescent; Ros asymmetrical, 30–80 cm ∅; L numerous, recurved and pendent, sheaths white to light brown, square to transversally oblong, 5–7.5  3.5–5 cm, whitelepidote on both faces, glabrescent, entire and sinuate, L lamina light green adaxially, whitelepidote abaxially, 30–80  1.5–4 cm, longattenuate, minutely serrate to entire; Inf erect to arching or pendent, 1–2 branched, glabrous; peduncle 6–13 mm ∅; peduncular Bra light brown to purplish-brown, foliaceous, linear to very narrowly triangular, 5–70  0.8–1 cm, entire

Hechtia BROMELIACEAE

to minutely serrate, glabrous adaxially, densely white-lepidote abaxially, involute, long-attenuate, longer than the internodes, becoming gradually shorter upwards; male Inf with 20–25 ascending primary Br 8–30 cm long; primary Bra light brown, linear to triangular, 10–20  3–8 mm, acuminate; secondary Br when present 2–6, 4–14 cm, divaricate to diffuse; floral Bra brown to purplish, linear, 2–3  0.5 mm, shorter than the pedicels; Fl polystichous, divaricate; Ped filiform, 5–7 mm; Sep purplish to rose, triangular to narrowly triangular, 1.5–2.8  1–1.7 mm, glabrous, entire, rounded; Pet lilac, ovate to elliptic, 5–6  3.3–3.8 mm, rounded; St subequal, longer than the petals; Fil 3–4.5 mm; Anth green, 1 mm; female Inf with 14–16 ascending primary Br 25–35 cm long; primary Bra brown-purplish, triangular, 2–7  0.8–1.2 cm, long-attenuate to acuminate; secondary Br 7–12, 5–16 cm, ascending; floral Bra light brown, linear to triangular, 1.6–1.8  1 mm, acuminate; Fl polystichous, divaricate; Ped filiform, 2.3–4 mm; Sep green to purplish, triangular, 2–2.2  1–1.3 mm, acute; Pet lilac, elliptic to oblong, 4–5  2.3–2.6 mm, rounded; Ov green, 3.5  3 mm; Fr green, narrowly ovoid, 7.7–8  3 mm, reflexed and pointing downwards at maturity. H. zamudioi Espejo & al. (Acta Bot. Mex. 83: 55–59, ills., 2008). Type: Mexico, Querétaro (Zamudio & Hernández 11285 [IEB, MEXU, NY, UAMIZ]). — Distr: Mexico (Querétaro: Peñamiller, Pinal de Amoles, San Joaquín); arid and montane shrubland, on vertical calcareous cliffs or rocky bluffs and ledges, 850–1650 m. Caspitose, forming extensive colonies, flowering to 2 m tall; Ros spreading, 60–70 cm ∅; L numerous, sheaths straw-yellow, widely ovate to subquadrate, 8.5–9  8.8–10.2 cm, lustrous, glabrous on both faces, with a yellow exudate, L lamina blue-green, slightly discolorous, 30–48  3.5–5 cm, densely white-lepidote on both faces, Sp light brown to reddish, antrorse, slender, 1.5–2 mm, 0.6–1.5 cm apart; Inf glabrous; peduncle cylindrical, 1.2–2 cm ∅; peduncular Bra foliaceous, straw-yellow, narrowly triangular to triangular, 10–30  1.8–2.5 cm, attenuate, pungent, the basal ones imbricate and

1031

longer than the internodes, becoming progressively reduced distally, sheaths subquadrate, 2–2.5 cm long, entire, lamina densely whitetomentose on both faces, spiny along the margins; male Inf 1–2 branched, with numerous primary Br 15–25 cm long, with 2 basal secondary Br 5–9.5 cm long; primary Bra light brown, narrowly triangular to lanceolate, 5.5–7  1.5–1.7 cm, glabrous, acuminate, hyaline and minutely spiny along the margins; floral Bra filiform, white, inconspicuous, to 3 mm, entire, glabrous; Fl divaricate to diffuse, laxly disposed; Ped filiform, 3–6 mm; Sep green, triangular to ovate or oblong, 3.5–4.5  2–3.5 mm, hyaline, acute; Pet green, elliptic, 6–8  3.5–4 mm, rounded; St subequal, shorter than the petals; Fil 6–7 mm; Anth green, 3 mm; female Inf 1 branched with numerous diffuse to ascending primary Br 6–10 cm long; primary Bra light brown, lanceolate to narrowly triangular or narrowly ovate, glabrous, 3.5–7  0.9–1.5 cm, acuminate, hyaline and minutely spiny along the margins; floral Bra white, filiform, inconspicuous, to 3 mm, entire, glabrous; Fl divaricate, laxly disposed; Ped filiform, 5–9 mm; Sep green, triangular to narrowly triangular, 3–4  1.2–1.4 mm, apiculate; Pet green, narrowly triangular to narrowly ovate, 5–8  2.5–3 mm, acute to rounded; Ov green, ovoid to ellipsoid, 5–7  3–4 mm; Fr green, light brown when mature, 8–12  5–6 mm; Se light brown, 5–6 mm.

References Burt-Utley, K. & Utley, J. F. (2011) Contributions toward a revision of Hechtia (Bromeliaceae, Pitcairnioideae) I. New and noteworthy species of Hechtia from Mexico. Phytoneuron 2011(59): 1–17, ills. Espejo-Serna, A. (2012) El endemismo en las Liliopsida mexicanas. Acta Bot. Mex. 100: 195–257. http://www. redalyc.org/articulo.oa?id=57424406008 Espejo-Serna, A. [& al. 2005], Lopez-Ferrari, A. R. & Ramirez-Morillo, I. (2005) Bromeliaceae. In: Sosa, V. (ed.): Flora de Veracruz, 136. Veracruz (MX): Instituto de Ecologia A. C. Espejo-Serna, A. [& al. 2007], Lopez-Ferrari, A. R., Martinez-Correa, N. & Pulido-Esparza, V. A. (2007) Bromeliad flora of Oaxaca, Mexico: Richness and distribution. Acta Bot. Mex. 81: 71–147, maps. http:// www.redalyc.org/articulo.oa?id=57408105

1032 Espejo-Serna, A. [& al. 2010a], Lopez-Ferrari, A. R. & Ramirez-Morillo, I. (2010a) Bromeliaceae. In: Rzedowski-Rotter, J. & Calderón de Rzedowski, G. (eds.): Flora del Bajio y de Regiones Adyacentes, 165. Pátzcuaro (MX): Instituto de Ecología. Espejo-Serna, A. [& al. 2010b], López-Ferrari, A. R. & Zamudio, S. (2010b) Reencuentro con Hechtia epigyna Harms (Bromeliaceae). Acta Bot. Mex. 90: 11–18, ills., map. http://www.redalyc.org/pdf/574/57412073003.pdf Espejo-Serna, A. [& al. 2010c], Martinez-Correa, N. & Lopez-Ferrari, A. R. (2010c) Neotypification of Hechtia podantha (Bromeliaceae). J. Bot. Res. Inst. Texas 4(1): 221–223. Espejo-Serna, A. [& al. 2012], López-Ferrari, A. R., CejaRomero, J. & Mendoza-Ruiz, A. (2012) Neotipificación de Hechtia lanata (Bromeliaceae; Hechtioideae), especie endémica de Oaxaca, México. Bot. Sci. 90(4): 385–388, ills. http://www.scielo.org.mx/pdf/bs/v90n4/v90n4a3.pdf Espejo-Serna, A. [& al. 2013], López-Ferrari, A. R., Martínez-Correa, N. & Till, W. (2013) A note on the typification of Hechtia galeottii (Hechtioideae, Bromeliaceae). Phytotaxa 94(2): 41–48, ills., map. https://doi.org/ 10.11646/phytotaxa.94.2.2. Felger, R. S. & Yetman, D. (2000) Roasting the Hechtia out of it: The use of Hechtia montana (Bromeliaceae) as a food plant in Sonora, Mexico. Econ. Bot. 54(2): 229–233. https://doi.org/10.1007/BF02907827. Kemble, B. (2016) Hechtia lanata. Cact. Succ. J. (US) 88(3): 104–109, ills. https://doi.org/10.2985/015.088.0302. Kopfstein, R. (2015) An ethnobotanical note: Hechtia edulis. J. Bromeliad Soc. 64(4): 258–263, ills. López-Ferrari, A. R. [& al. 2008], Espejo-Serna, A., CejaRomero, J. & Mendoza-Ruiz, A. (2008) Hechtia matudae, a spectacular, though neglected Bromeliad from Mexico. J. Bromeliad Soc. 58(2): 56–60, ills. López-Ferrari, A. R. [& al. 2011], Espejo-Serna, A., CejaRomero, J. & Mendoza-Ruiz, A. (2011) Hechtia lundelliorum, another spectacular and little known bromeliad from Mexico. J. Bromeliad Soc. 60(5): 211–215, ills. Martínez-Correa, N. (2008) Sistematica del complejo de especies de Hechtia podantha Mez (Pitcairnioideae, Bromeliaceae). Iztapalapa (MX): Universidad Autónoma Metropolitana; unpubl. M.Sc. thesis. McVaugh, R. (1989) Flora Novo-Galiciana. A descriptive account of the vascular plants of Western Mexico. Vol. 15: Bromeliaceae to Dioscoreaceae. Ann Arbor (US): University of Michigan Herbarium. Mez, C. (1896) Bromeliaceae. In: Candolle, C. De (ed.): Monographiae phanerogamarum. Vol. 9. Paris (FR): Masson & Cie. https://doi.org/10.5962/bhl.title.45961.

A. Espejo-Serna et al. Mez, C. (1935) Bromeliaceae. In: Engler, A. & Diels, L. (eds.): Das Pflanzenreich; IV(32), Heft 100 [parts 2–4]. Berlin (DE): Wilhelm Engelmann. http://bibdigital.rjb. csic.es/ing/Libro.php?Libro=592 Pulido-Esparza, V. A. [& al. 2004], Lopez-Ferrari, A. R. & Espejo-Serna, A. (2004) Flora bromeliológica del estado de Guerrero, Mexico: Riqueza y distribución. Bol. Soc. Bot. México 75: 55–104. http://www.redalyc. org/articulo.oa?id=57707504 Ramírez Morillo, I. M. & Jimenez-Nah, C. F. (2012) A new species of Hechtia (Hechtioideae, Bromeliaceae) from Puebla, Mexico. Phytotaxa 42: 1–8. https://doi.org/ 10.11646/phytotaxa.42.1.1. Ramírez Morillo, I. [& al. 2014a], Jiménez, C. F., FernándezConcha, G. C. & Pinzón, J. P. (2014a) Three new species and growth patterns in Hechtia (Bromeliaceae: Hechtioideae). Phytotaxa 178(2): 113–127, map, ills. https://doi.org/10.11646/phytotaxa.178.2.3. Ramírez Morillo, I. [& al. 2014b], Carnevali, G., Jiménez Nah, C. & Pinzón, J. P. (2014b) Portraits of Hechtia (Bromeliaceae: Hechdioideae [sic!]): Hechtia isthmusiana Burt-Utley. J. Bromeliad Soc. 64(1): 26–33, ills. Ramírez-Morillo, I. [& al. 2016], Carrillo-Reyes, P., TapiaMuñoz, J. L. & Cetzal-Ix, W. (2016) An addition to genus Hechtia (Hechtioideae; Bromeliaceae) from Jalisco, Mexico. Phytotaxa 266(4): 261–270, map, ills. https://doi.org/10.11646/phytotaxa.266.4.3. Roguenant, A. [& al. 2016], Lecoufle, M. & RaynalRoques, A. (2016) Les Broméliacées. Approche panoramique d’une grand famille “américaine”. Paris (FR): Belin. Rzedowski, J. (1991) El endemismo en la flora fanerogamica Mexicana: Una apreciación analitica preliminar. Acta Bot. Mex. 15: 47–64, ills. http://www. redalyc.org/articulo.oa?id=57401505 Siekkinen, A. (2015) The mystery of Hechtia argentea — Part 1 / Part 2. J. Bromeliad Soc. 65(1): 48–57, (2): 100–110, ills. Smith, L. B. (1938) Bromeliaceae. In: North American Flora 19(2): 61–228, keys. New York (US): New York Botanical Garden. http://biodiversitylibrary.org/page/28708822 Smith, L. B. & Downs, R. J. (1974) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 1: Pitcairnioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/4393694 Utley, J. & Burt-Utley, K. (1994) Bromeliaceae. In: Davidse, G., Sousa S., M. & Chater, A. O.(eds.): Flora Mesoamericana 6: 89–156. Mexico (MX): Universidad Nacional Autonoma. http://www.tropicos.org/Name/ 42000361?projectid=3&langid=66

Neobergia BROMELIACEAE U. Eggli

Neobergia E. L. Smith (J. Bromeliad Soc. 33 (2): 73, 1983). — Distr: Cultivated only. = Neoregelia  Billbergia. Smith (1983) lists the combination B. nutans  N. spectabilis (=  Neobergia ‘Perneri’), and later, B. nutans  N. carolinae was mentioned as  Neobergia ‘Noddy’. The plants are like a robust B. nutans but become red during flowering, and have a

shortened,  erect (rather than arching or hanging) inflorescence.

References Smith, E. L. (1983) Bigeneric hybrids — A listing. J. Bromeliad Soc. 33(2): 69–75. http://journal.bsi.org/V33/2/

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_97

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Neoglaziovia BROMELIACEAE U. Eggli

Neoglaziovia Mez (in Martius, Fl. Bras. 3(3): 426, 1894). Type: Bromelia variegata Arruda. — Bromelioideae — Lit: Smith & Downs (1979: 2036–2038, Fl. Neotropica). Distr: NE Brazil. Etym: Gr. ‘neos’, new (to avoid a homonym); and for Auguste F. M. Glaziou (1828–1906), French botanical traveller collecting 1861–1895 in Brazil. Perennial terrestrial Ros plants, stemless, with underground rhizomes, forming dense to open colonies; L 3–10 per rosette, with indistinct entire sheath, lamina very long and narrow, stiff and succulent, margins with laxly arranged short Sp 2–4 ( 6) mm; Inf terminal, shorter than the leaves, unbranched, peduncular Bra longer than the internodes, leaf-like, fertile part few- to manyflowered, lax or dense; Fl shortly pedicellate or sessile, glabrous; Sep free,  symmetrical, red; Pet free, symmetrical, bright purple, with 2 scales at the base; St included; Fil free; Ov completely inferior; Fr fleshy berries. Neoglaziovia is easily recognized by the rosettes with few ascending to erect, narrowly linear and succulent leaves. Its systematic position within Bromelioideae is not well-resolved. Schulte & al. (2009) found it as basal sister of the “core bromelioids”, but with limited support. Evans

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

& al. (2015) did not even find the 2 studied species (N. variegata, N. burle-marxii) to form a monophyletic clade, and both appear as separate entities as part of an extensive polytomy in their EuBromelioid clade. Leal & al. (2006) and Lemos Pereira & Maciel Quirino (2008) report hummingbird pollination (mainly by Chlorostilbon aureoventris) for N. variegata. — An intergeneric hybrid with Orthophytum has been formally named Orthoglaziovia. N. burle-marxii Leme (J. Bromeliad Soc. 40(3): 101–103, ills., 1990). Type: Brazil, Bahia (BurleMarx s.n. [HB, RB]). — Distr: Brazil (Bahia); Caatinga vegetation, sandy soil. L to 3 m, lamina linear, narrowing gradually to the tip, 2.5 cm wide at the base, very rigid and thick, strongly canaliculate, upper face inconspicuously white-lepidote, lower face uniformly densely white-lepidote in longitudinal rows; Inf to 70 cm, erect, peduncular Bra not completely covering the peduncle, lower ones leaf-like, upper ones narrowly lanceolate, reddish, densely whitelepidote on both faces; fertile part of the Inf dense, 30- to 40-flowered, with an apical tuft of small bracts; Fl sessile, spreading-ascending; Sep suborbicular; Pet long obovate, 13–18 mm. N. concolor C. H. Wright (Curtis’s Bot. Mag. 136: t. 8348 + text, 1910). Type: K, GH [photo]. — Distr: NE Brazil (Bahia); Caatinga vegetation, sandy soils.

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_91

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U. Eggli

green to brown-green, smooth, lower face with broad white bands, margins somewhat revolute; Inf erect to  inclined, densely white-floccose; peduncular Bra narrow, entire or slightly serrulate; fertile part of the Inf lax, 10- to 60-flowered; lower floral Bra  as long as the flowers, linear, upper to 3 mm, triangular; Ped to 4 mm; Fl somewhat spreading; Sep obtuse to minutely mucronate, 6–7 mm; Pet obtuse, to 13 mm. — Cytology: 2n = 100 (tetraploid) (Gitaí & al. 2014). The species is has been extensively used as a source for fibre for making textiles especially in rural areas (Xavier 1942), and fibres are classified as having low to medium strength in comparison with other lignocellulosic fibres (Almeida & al. 2008). The leaves are also harvested to feed to animals, and overharvesting has caused the disappearance of the species from some areas in Bahia (Silveira & al. 2011).

References Fig. 1 Neoglaziovia variegata. (Copyright: U. Eggli)

L to 0.6 m but probably becoming longer, lamina to 2.5 cm wide, long-acuminate, both faces uniformly appressedly white-lepidote; Inf erect, peduncular Bra not described; lower floral Bra linear, longer than the flowers, upper ones to 4 mm, triangular; Ped 5–7 mm; Fl spreading; Sep broadly ovate, rounded; Pet broadly round, to 20 mm. N. variegata (Arruda) Mez (in Martius, Fl. Bras. 3(3): 427, t. 80, fig. 1, 1894). Type: not typified. — Distr: NE Brazil (Piauí, Ceará, Rio Grande do Norte, Paraiba, Bahia, Minas Gerais); Caatinga vegetation on stony to sandy ground. I: Leme & Marigo (1993: 156–157). – Fig. 1.  Bromelia variegata Arruda (1810)  Billbergia variegata (Arruda) Schultes fil. (1830)  Agallostachys variegata (Arruda) Beer (1856); incl. Bromelia linifera hort. ex Beer (1856) (nom. inval., Art. 34.1c); incl. Dyckia glaziovii Baker (1889). L to 1.5 m, lamina 1.5–2 cm wide, acuminatepungent, both faces laxly lepidote, upper face

Almeida, J. R. M. d’ [& al. 2008], Almeida, A. L. F. S. d’ & Carvalho, L. H. (2008) Mechanical, morphological, and structural characteristics of Caroa Neoglaziovia variegata fibres. Polymers & Polymer Composites 16(9): 589–595. https://doi.org/10.1177/096739110801600902 Evans, T. M. [& al. 2015], Jabaily, R. S., Gelli de Faria, A. P., Oliveira F. de Sousa, L. de, Wendt, T. & Brown, G. K. (2015) Phylogenetic relationships in Bromeliaceae subfamily Bromelioideae based on chloroplast DNA sequence data. Syst. Bot. 40(1): 116–128. https://doi. org/10.1600/036364415X686413. Gitaí, J. [& al. 2014], Paule, J., Zizka, G., Schulte, K. & Benko-Iseppon, A. M. (2014) Chromosome numbers and DNA content in Bromeliaceae: Additional data and a critical review. Bot. J. Linn. Soc. 176(3): 349–368, ills. https://doi.org/10.1111/boj.12211. Leal, F. C. [& al. 2006], Lopes, A. V. & Machado, I. C. (2006) Polinização por beija-flores em uma area de caatinga no Municipio de Floresta, Pernambuco, Nordeste do Brasil. Revista Brasil. Bot. 29(3): 379–389. https://doi.org/10.1590/S0100-84042006000300005. Leme, E. M. C. & Marigo, L. C. (1993) Bromeliads in the Brazilian wilderness. Rio de Janeiro (BR): Marigo Comunicação Visual. Lemos Pereira, F. R. de & Maciel Quirino, Z. G. (2008) Fenologia e biologia floral de Neoglaziovia variegata (Bromeliaceae) na caatinga paraibana. Rodriguésia 59(4): 835–844, ills. Schulte, K. [& al. 2009], Barfuss, M. H. & Zizka, G. (2009) Phylogeny of Bromelioideae (Bromeliaceae) inferred

Neoglaziovia BROMELIACEAE from nuclear and plastid DNA loci reveals the evolution of the tank habit within the subfamily. Molec. Phylogen. Evol. 51(2): 327–339. https://doi.org/10.101 6/j.ympev.2009.02.003. Silveira, D. G. [& al. 2011], Pelacani, C. R., Antunes, C. G. C., Rosa, S. S., Duarte Souza, F. V. & Santana, J. R. F. de (2011) Resposta germinativa de sementes de caroá [Neoglaziovia variegata (Arruda) Mez]. Ci. Agrotecnol. 35

1037 (5): 948–955. https://doi.org/10.1590/S1413-705420 11000500012. Smith, L. B. & Downs, R. J. (1979) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 3: Bromelioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/i399940. Xavier, L. P. (1942) O caroá, história, cultura e distribuição geográfica. Rio de Janeiro (BR): Ministerio da Agricultura.

Neophytum BROMELIACEAE U. Eggli

Neophytum M. B. Foster (Bromeliad Soc. Bull. 8: 73, 1958). — Distr: Cultivated only. = Neoregelia  Orthophytum. Smith & Downs (1979: 2064) mention the hybrid N. bahiana  O. navioides (now classified as Sincoraea), for which the formal name  Neophytum lymanii M. B. Foster was published in 1964 (illustrated by Roguenant & al. (2016: 587)). Many more cultivars are offered in the horticultural trade, many of them producing rosettes with numerous gracefully ascending to arching leaves that turn red in strong light.

References Roguenant, A. [& al. 2016], Lecoufle, M. & RaynalRoques, A. (2016) Les Broméliacées. Approche panoramique d’une grand famille “américaine”. Paris (FR): Belin. Smith, L. B. & Downs, R. J. (1979) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 3: Bromelioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/sta ble/i399940

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_98

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Ochagavia BROMELIACEAE U. Eggli

Ochagavia Philippi (Anales Univ. Chile 13: 168, 1856). Type: Ochagavia elegans Philippi. — Bromelioideae — Lit: Smith & Downs (1979: 1527–1533, Fl. Neotropica); Zizka & al. (2002: monograph). Distr: C to S-C Chile (incl. Juan Fernández archipelago). Etym: For Silvestre Ochagavía Errázuriz (1820–1883), Chilean lawyer, politician and businessmen, minister of education 1853–1854, and founder of the Ochagavia Winery in 1851. Incl. Rhodostachys Philippi (1858). Type: Rhodostachys andina Philippi. Incl. Ruckia Regel (1868). Type: Ruckia ellemetii Regel. Incl. Placseptalia Espinosa (1947). Type: Placseptalia rebecae Espinosa. Perennial terrestrial rosette plants, shortly to distinctly caulescent, forming extensive stands through offsetting; L numerous, quite densely arranged, L sheath ovate to orbicular, merging into the lamina, conspicuously veined, coriaceous, quite succulent, marginally spinose, L lamina narrowly triangular to linear, gradually attenuate towards the tip, to 80 (
120) cm, rigid, coriaceous and quite succulent, canaliculate or flat, upper

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

face glabrous or sparsely lepidote near the base, lower face densely to sparsely white-lepidote to glabrescent, marginal spines 1–4.5 mm, antrorse to spreading, innermost L at flowering time rosered, with reduced lamina; Inf terminal, single, condensed and capitate to subcorymbose, with 7–>50 densely arranged flowers, sessile or to 17.5 cm pedunculate, peduncular Bra similar to innermost rosette leaves; floral Bra shorter than the flowers, acute, pungent, membranous, rosered to white, with spinose margins; Fl 3.5–7.9 cm, hermaphrodite, actinomorphic, sessile, with a conspicuous epigynous tube to 19 mm; Sep free,  symmetrical, chartaceous, whitish to pale rosered; Pet free, erect, obovate to spatulate, without basal appendages, rose-red; St usually exserted; Fil free or basally united with the petals (only O. elegans); Ov inferior; Sty exserted and longer than the stamens; Sti lobes conduplicate, hardly contorted; Fr berries, 2–4.5 cm (without the persistent perianth remains), brown to black when dry; Se numerous, flattened, glabrous, unappendaged, dark brown to black. A small genus of 4 species, endemic to Chile between 31 330 and 38 140 S, essentially confined to the mediterranean climate zone. The leaves are distinctly succulent, with a water storage volume of 66% reported for O. litoralis (Horres & Zizka 1995: as O. carnea). Ochagavia is closely related to the monotypic genus Fascicularia, which differs by retuse or

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_92

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praemorse sepals, blue-violet petals with appendages, and included stamina and style, but sterile plants are virtually impossible to distinguish. The molecular phylogenetic analysis of Horres & al. (2007) confirms the long-suspected close relationship between the two genera. They are, together with Deinacanthon, Greigia and Bromelia, part of a weakly supported clade that is sister to the remaining genera of Bromelioideae (Givnish & al. 2011), confirming similar placements in the phylogenies of Schulte & al. (2005), Schulte & Zizka (2008), and Schulte & al. (2009). The phylogeny of Silvestro & al. (2014) places Fascicularia within Ochagavia, and the combined clade is sister to Greigia. Evans & al. (2015) confirm the inclusion of Fascicularia and Deinacanthon in Ochagavia (Greigia not studied). Plants are only occasionally found in cultivation, esp. in Botanical Gardens. Hummingbird pollination has been reported for O. elegans (Skottsberg 1928, Bernardello & al. 2001). The following names are of unresolved application but are referred to this genus: Rhodostachys leiboldiana Mez (1896)  Ochagavia leiboldiana (Mez) L. B. Smith & Looser (1934). O. andina (Philippi) Zizka & al. (Willdenowia 32(2): 343, 2002). Type: Chile (Germain s.n. [SGO 46378, FR [photo]]). — Distr: C-S Chile (Región VI O’Higgins: Cachapoal to Región VIII: Bíobío); 700–2500 m. Rhodostachys andina Philippi (1858). L lamina (14.3–) 30–40 cm, (0.5–) 0.9–1.7 cm wide, linear, faintly canaliculate, upper face glabrous to lepidote near the base, lower face densely white-lepidote; Inf globose to ovoid,  30-flowered, to 15 cm pedunculate; Fl 5.3–6.8 cm, epigynous tube 5–7 mm; St exserted. Insufficiently known. Most collections date from the 19. or early 20. century, and the taxon is likely very rare today. O. carnea (Beer) L. B. Smith & Looser (Revista Univ. (Santiago) 18(8): 1078, 1934).

U. Eggli

Type: [lecto — icono]: Paxton’s Fl. Gard. 2, t. 65, 1851. — Distr: C to S-C Chile (Regíon V Valparaiso to Región IX Araucanía); understorey of woodlands, in ravines and on riverbanks, (60-) 200–1080 m. I: Rauh (1990: 212: fig. 97); Roguenant & al. (2016: 607). Bromelia carnea Beer (1856)  Rhodostachys carnea (Beer) Mez (1896); incl. Bromelia longifolia Lindley (1851) (nom. illeg., Art. 53.1); incl. Bromelia lindleyana Lemaire (1853) (nom. inval., Art. 34.1b); incl. Rhodostachys grandiflora Philippi (1859)  Ochagavia grandiflora (Philippi) Mez (1934); incl. Hechtia ellemeetii Hort. Utrecht (1866) (nom. inval., Art. 32.1c); incl. Ruckia ellemetii Regel (1867); incl. Pourretia argentea hort. ex Wittmack (1895) (nom. inval., Art. 32.1c); incl. Rhodostachys chamissonis Mez (1896)  Ochagavia chamissonis (Mez) L. B. Smith & Looser (1934). Stems to >25 cm; L lamina (38–) 50–80 (
120) cm, 1.2–3.4 (
4.8) cm wide, linear, conspicuously canaliculate, upper face glabrescent, lower face sparsely lepidote to glabrescent, only near the base more densely lepidote; Inf globose to elongate, to >50-flowered, to 17.5 cm pedunculate; Fl (3.5–) 4.5–7.9 cm, epigynous tube 3–7 mm; St exserted. O. elegans Philippi (Anales Univ. Chile 13: 168, 1856). Type: Chile, Juan Fernández (Philippi 941 [B, F, GH [photo], W †]). — Distr: Chile: Juan Fernández archipelago: Isla Robinson Crusoe, endemic; rocky exposed cliffs, 200–600 m. I: Wilkin (1996). Stems to 72 cm; L lamina (7.5–) 10–24 cm, 1–2.9 cm wide, narrowly triangular, upper face flat or slightly canaliculate, glabrous to  lepidote, lower face densely white-lepidote; Inf capitate, to 20-flowered, 2.5 cm pedunculate; Fl 5–7 cm, epigynous tube 10–19 mm; St included or slightly exserted. O. litoralis (Philippi) Zizka & al. (Willdenowia 32(2): 340, 2002). Type: Chile, Colchagua (Volckmann s.n. [SGO 46374]). — Distr: Coastal C Chile (Región IV Coquimbo to Región VI

Ochagavia BROMELIACEAE

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canaliculate, upper face glabrous to sparsely lepidote, lower face densely white-lepidote; Inf globose to ovoid, 15- to 35-flowered, 8–12 cm pedunculate; Fl 4–6.3 cm, epigynous tube 3–6 mm; St exserted.

References

Fig. 1 Ochagavia litoralis (Engraving from Revue Horticole, sér. 4, 40: facing p. 211, as Hechtia pitcairniifolia, 1868). (Copyright: Public Domain)

O’Higgins: Colchagua); coastal rocks and cliffs, often in extensive stands, 10–250 m. I: Roguenant & al. (2016: 608). – Fig. 1. Rhodostachys litoralis Philippi (1859)  Fascicularia litoralis (Philippi) Mez (1896); incl. Hechtia pitcairniifolia Hort. Berol. ex B. Verlot (1868)  Bromelia pitcairniifolia (Hort. Berol. ex B. Verlot) K. Koch (1868)  Rhodostachys pitcairniifolia (Hort. Berol. ex B. Verlot) Bentham (1883)  Fascicularia pitcairniifolia (Hort. Berol. ex B. Verlot) Mez (1896); incl. Ochagavia lindleyana Mez (1934); incl. Placseptalia rebecae Espinosa (1947). Stems to >20 cm; L lamina 17–40 cm, 1–3 cm wide, narrowly triangular, not or only slightly

Bernardello, G. [& al. 2001], Anderson, G. J., Stuessy, T. F. & Crawford, D. J. (2001) A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernandez Islands (Chile). Bot. Rev. (Lancaster) 67(3): 255–308. https:// doi.org/10.1007/BF02858097. Evans, T. M. [& al. 2015], Jabaily, R. S., Gelli de Faria, A. P., Oliveira F. de Sousa, L. de, Wendt, T. & Brown, G. K. (2015) Phylogenetic relationships in Bromeliaceae subfamily Bromelioideae based on chloroplast DNA sequence data. Syst. Bot. 40(1): 116–128. https://doi. org/10.1600/036364415X686413. Givnish, T. J. [& al. 2011], Barfuss, M. H. J., Ee, B. van, Riina, R., Schulte, K., Horres, R., Gonsiska, P. A., Jabaily, R. S., Crayn, D. M., Smith, J. A. C., Winter, K., Brown, G. K., Evans, T. M., Holst, B. K., Luther, H., Till, W., Zizka, G., Berry, P. E. & Sytsma, K. J. (2011) Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: Insights from an eightlocus plastid phylogeny. Amer. J. Bot. 98(5): 872–895, ills., maps. https://doi.org/10.3732/ajb.1000059. Horres, R. & Zizka, G. (1995) Untersuchungen zur Blattsukkulenz bei Bromeliaceae. Beitr. Biol. Pfl. 69(1): 43–76, ills. Horres, R. [& al. 2007], Schulte, K., Weising, K. & Zizka, G. (2007) Systematics of Bromelioideae (Bromeliaceae) — evidence from molecular and anatomical studies. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution: Poales. Aliso 23(1): 27–43, ills. https://doi.org/10.5642/aliso.20072301.05 Rauh, W. (1990) Bromelien. Ed. 3. Stuttgart (DE): Verlag Eugen Ulmer. Roguenant, A. [& al. 2016], Lecoufle, M. & Raynal-Roques, A. (2016) Les Broméliacées. Approche panoramique d’une grand famille “américaine”. Paris (FR): Belin. Schulte, K. & Zizka, G. (2008) Multi locus plastid phylogeny of Bromelioideae (Bromeliaceae) and the taxonomic utility of petal appendages and pollen characters. Candollea 63(2): 209–225. http://www. ville-ge.ch/cjb/publications_candollea_632.php Schulte, K. [& al. 2005], Horres, R. & Zizka, G. (2005) Molecular phylogeny of Bromelioideae and its implications on biogeography and the evolution of CAM in the family. Senckenberg. Biol. 85(1): 113–125, maps. Schulte, K. [& al. 2009], Barfuss, M. H. & Zizka, G. (2009) Phylogeny of Bromelioideae (Bromeliaceae) inferred

1044 from nuclear and plastid DNA loci reveals the evolution of the tank habit within the subfamily. Molec. Phylogen. Evol. 51(2): 327–339. https://doi.org/10.10 16/j.ympev.2009.02.003. Silvestro, D. [& al. 2014], Zizka, G. & Schulte, K. (2014) Disentangling the effects of key innovations on the diversification of Bromelioideae (Bromeliaceae). Evolution 68(1): 163–175. https://doi.org/10.1111/ evo.12236. Skottsberg, C. (1928) Pollinationsbiologie und Samenverbreitung auf den Juan Fernández-Inseln. In: Skottsberg, C. (ed.): The natural history of Juan Fernández and Easter Island, Botany, 2(4): 503–547, ills. Uppsala

U. Eggli (SE): Almqvist & Wiksells Boktryckeri. http:// biodiversitylibrary.org/page/28693416 Smith, L. B. & Downs, R. J. (1979) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 3: Bromelioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/i399940 Wilkin, P. (1996) Ochagavia elegans. Curtis’s Bot. Mag., ser. nov., 13(1): 22–26, t. 287. https://doi.org/10.1111/ j.1467-8748.1996.tb00532.x. Zizka, G. [& al. 2002], Trumpler, K. & Zöllner, O. (2002) Revision of the genus Ochagavia (Bromeliaceae, Bromelioideae). Willdenowia 32(2): 331–350, ills., map, key. http://www.jstor.org/stable/3997641.

Orthocohnia BROMELIACEAE N. Schütz

Orthocohnia G. Lawn (J. Bromeliad Soc. 63 (2): 138, 2013). — Distr: Cultivated only. = Orthophytum  Deuterocohnia. The only known hybrid is the cultivar ‘Surprise’, raised in a Florida nursery in 1996 from the cross

Deuterocohnia lorentziana (treated as synonym of D. brevifolia in this handbook)  Orthophytum estevesii (pollen parent). It is similar to D. brevifolia, but forms larger plants.

N. Schütz (*) Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_99

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Orthoglaziovia BROMELIACEAE U. Eggli

Orthoglaziovia Lawn (J. Bromeliad Soc. 61(5): 234, 2012). — Distr: Cultivated only. = Orthophytum  Neoglaziovia. A single cultivar ‘Rosita’ has been named for the cross N. variegata  O. albopictum (now classified as

Sincoraea), obtained in cultivation. The plants are roughly intermediate between the parents, with an almost sessile, cone-shaped inflorescence with pale pinkish flowers.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_100

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Orthomea BROMELIACEAE U. Eggli

Orthomea E. L. Smith (J. Bromeliad Soc. 33 (2): 75, 1983). = Orthophytum  Aechmea. Reported combinations include A. fasciata var. purpurea  O.

navioides (the latter now classified as Sincoraea) (introduced into cultivation as cultivar ‘Powderpuff’) and A. recurvata var. benrathii  O. vagans (unnamed).

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_101

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Orthophytum BROMELIACEAE U. Eggli and E. J. Gouda

Orthophytum Beer (Flora 37: 347, 1854). Type: Prantleia glabra Mez [typification according to L. B. Smith & Downs, Fl. Neotrop. 14(3): 1696, 1979]. — Bromelioideae — Lit: Smith & Downs (1979: 1696-1710, Fl. Neotropica); Leme (2008: key to the mello-barretoi-complex); Louzada & al. (2010: cytology); Louzada & Wanderley (2010: partial monograph of the vagans-clade); Louzada (2012: unpublished monograph, with key); Louzada & al. (2014: molecular phylogeny); Leme & al. (2017: classification). Distr: E Brazil; Atlantic Forest, Cerrado and Caatinga vegetations, usually saxicolous. Etym: Gr. ‘orthos’, erect, straight; and Gr. ‘phyton’, plant; for the erect stems with well-spaced leaves of the type species. Incl. Prantleia Mez (1891). Type: Prantleia glabra Mez. Incl. Cryptanthopsis Ule (1908). Type: Cryptanthopsis saxicola Ule. Perennial terrestrial or saxicolous rosette plants; Ros sessile or short- to long-caulescent, sometimes elongating-desintegrating when flowering,

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] E. J. Gouda Curator University Botanic Gardens, Utrecht, The Netherlands e-mail: [email protected]

monocarpic but offsetting, sometimes with short stolons; L few to many, laxly to densely rosulate or spirally arranged on elongate stems, sheath inconspicuous or  indistinctly separated from the lamina, L lamina linear to narrowly triangular, or ovate-triangular, coriaceous to succulent, flat or canaliculate, glabrous to densely lepidote, margins serrate and with antrorse to retrorse spines, apex acuminate,  pungent; Inf terminal, elongate and rosette desintegrating with inflorescence growth, and then with spikes, spikes of spikes, or glomerules in the axils of the primary bracts, or congested and sessile in the centre of the caulescent rosette; peduncular Bra resembling the leaves but progressively diminishing in size; floral Bra subcoriaceous to coriaceous, narrowly triangular to ovate-triangular, sometimes strongly carinate, shorter to longer than the sepals, green to green-yellowish or red to dark wine-red, margins serrate, apex acuminate and/or pungent; Fl sessile to shortly pedicellate, hermaphrodite; Sep narrowly triangular to ovate-triangular, free or rarely forming a short tube, acuminate and/or pungent; Pet often spatulate, free or forming an epigynous tube to 2 mm long, white to green, greenish-yellow or lilac-rose, upper spreading or erect, obtuse to obtuse-cucullate or acute, basal appendages echinatiform, cupuliform, scutelliform or sacciform (absent in Subgen. Orthocryptanthus); St included; Fil of the inner whorl adnate to the petals; Ov inferior,  trigonous, glabrous to densely lanate-lepidote; Sty long and slender; Sti lobes simple, erect, narrow; Fr subglobose to

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globose capsules, with persistent sepals; Se ovoid, striate, few to numerous per fruit. The majority of the 55 species has been described in the past 20 years or so (Leme & al. 2017). There are two centres of diversity, one in the Serra do Espinhaço Range in Bahia and Minas Gerais, the other in the inselberg regions of Minas Gerais and Espírito Santo. Many of the species are narrow endemics. Until recently, the genus also included species with sessile rosettes with congested, sessile inflorescences and white flowers now segregated as Sincoraea (see separate treatment in this handbook). Orthophytum is placed in subfamily Bromelioideae, where it belongs to a group of early-diverging genera (Schulte & Zizka 2008, Schulte & al. 2009). Schulte & al. (2009) found a position as sister group of Ananas for the 3 taxa of Orthophytum s.s. analyzed, though with low support. Silvestro & al. (2014) and Evans & al. (2015) analyzed more species and found that Orthophytum is not monophyletic. In the analysis of Silvestro & al. (2014) Orthophytum s.l. falls in 2 clades (Orthophytum s.s. versus the taxa now segregated as Sincoraea), being sister to Cryptanthus p.p. and the combined clade sister to Ananas. The phylogeny obtained by Evans & al. (2015) differs insofar as the species now segregated as Sincoraea appear in sister-group position to 2 Aechmea species and Ronnbergia petersii (as sister to Lapanthus, within a well-supported “Hohenbergia-Orthophytum clade”), while the species of Orthophytum s.s. form a well-supported clade with Aechmea floribunda and Canistrum triangulare in the extensive polytomy that makes up their Eu-Bromelioid clade. The most detailed analysis is that of Louzada & al. (2014), which is largely congruent with the clades found by Silvestro & al. (2014). Orthophytum s.s. accordingly is closely related to the morphologically often similar genera Cryptanthus (see separate treatment in this handbook) and Lapanthus (2 species, not considered to include succulents). The traditional distinction of Orthophytum s.s. (petals with basal appendages) from Cryptanthus (petals without basal appendages) is thus probably not warranted, esp. because the presence/absence

U. Eggli and E. J. Gouda

of petal appendages is not phylogenetically informative according to Schulte & Zizka (2008). On the other hand, Cryptanthus appears to differ consistently from Orthophytum in its generally low chromosome numbers (2n = 34, 36, or 54, vs. a polyploid series of 2n = 50, 100, 150 in Orthophytum and the species now segregated as Sincoraea) (Louzada & al. 2010, Louzada & al. 2014). Lapanthus is similar to Orthophytum s.s. but differs by completely erect petals, free filaments of the inner whorl, and lanceolate (vs. saccate or cupuliform) petal appendages (Louzada & Versieux 2010). More detailed analyses and a more complete study of Cryptanthus are necessary to further elucidate the relationship between that genus and Orthophytum s.s. Traditionally, Orthophytum was divided into 2 main groups, based on whether the inflorescence is sessile or long-pedunculate, and Leme (2004) provided a further break-down into 6 “subcomplexes”. These groupings were only partly recovered in the molecular phylogenies of Louzada & al. (2014) and Leme & al. (2017). Based on their results, Leme & al. (2017) divide Orthophytum into 5 subgenera, but only Subgen. Orthophytum (in terminal sister-position to Subgen. Clavanthus) is strongly supported, while the topology of the remaining clades is less clearly resolved, and Subgen. Capixabanthus is shown as polyphyletic in the molecular data. All subgenera are morphologically well circumscribed and (with the exception of the wide-spread Subgen. Orthophytum) show neatly defined geographical ranges: [1] Subgen. Orthophytum: Plants (nearly) stemless; L thin to thick-coriaceous; Sep free; Pet erect except the suberect to recurved apical part, white or green, appendage echinatiform; Sti simple-dilated (rarely simple-patent). — 43 spp., widespread throughout the range of the genus, to 500 (rarely 600–1200) m. [2] Subgen. Clavanthus Leme & al. 2017: Plants stemless; L strongly coriaceous; Sep free; Pet erect and corolla tubular-clavate, green, green with white apex, or greenish-yellow, appendages scutelliform (rarely sacciform); Sti

Orthophytum BROMELIACEAE

simple-dilated. — 6 species from Minas Gerais 650–1300 m. [3] Subgen. Orthocryptanthus Leme & al. 2017: Plants long-caulescent; L coriaceous; Sep free; Pet spreading-recurved, lilac-rose or white, appendages absent; Sti simple-erect to simple-patent. — 2 species from Minas Gerais, 1000–1400 m. [4] Subgen. Capixabanthus Leme & al. 2017 (= “Vagans-clade of earlier classifications): Plants stemless to long-caulescent; L thin to thick-coriaceous; Sep free; Pet suberect or rarely spreading, green with withe apex or white, appendages cupuliform or sacciform; Sti simple-dilated. — 7 species from Espirito Santo and neighbouring Bahia and Minas Gerais, to 600 m. [5] Subgen. Krenakanthus Leme & al. 2017: Plants caulescent; L thin; Sep connate for – of their length; Pet spreading, roselilac to purple towards the apex, appendages cupuliform; Sti conduplicate-spiral. — Only O. roseolilacinum from Minas Gerais, 250–810 m, not succulent. Succulence: The degree of succulence varies throughout the genus, but can be very strongly pronounced in taxa from more xeric habitats. Horres & Zizka (1995) specifically mention O. disjunctum, but their material was probably misidentified as cultivated plants of this species are hardly succulent. Horticulture: Horticulturally, Orthophytum s.s. is of limited importance, and its species are not commonly seen in cultivation. Due to the small size and the xeromorphic features, many species are well-adapted to cultivation together with other xerophytes and succulents. Plants can easily be propagated from the short basal shoots produced before flowering, or from shoots arising at the base of the inflorescence, depending on the species. The following intergeneric hybrids have been recorded: Orthophytum  Aechmea = Orthomea (see separate entry in this handbook); Orthophytum  Cryptanthus = Orthotanthus (see

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separate entry in this handbook); and Orthophytum  Nidularium = Ortholarium (not involving succulent taxa). O. argentum Louzada & Wanderley (Phytotaxa 28: 27–29, ills., 2011). Type: Brazil, Bahia (Louzada & al. 110 [SP, HUEFS]). — Distr: E Brazil (Bahia: Rio de Contas); quartzite rock outcrops in Campo Rupestre vegetation, shaded or rarely open places, 900–1300 m. Incl. Orthophytum toscanoi ssp. atropurpureum P. J. Braun & Esteves (2010). [1] Ros open, shortly caulescent, 24–34 cm tall, with short rhizomatose offsets, stems completely covered by the leaves; L arching, sheath distinguishable from the lamina, chartaceous, ovate, castaneous, both faces glabrous, margins serrulate, L lamina narrowly triangular, 15–20  2–2.5 cm, rigid, coriaceous, canaliculate, with long attenuate pungent tip, cinereous, densely lepidote on both faces, margins densely serrate, Sp slender, dark brown, 3.5–4 mm but gradually shorter towards the leaf tip, retrorse; Inf pedunculate, branched with sessile glomerate branches; peduncle elongate, erect, 11–18 cm, woolly-lepidote, dark winered; peduncular Bra narrowly triangular with long-attenuate pungent tip, divergent, much exceeding the internodes, leaf-like, coriaceous, margins densely serrate and lepidote; fertile Inf part 12 cm, with 7–8 condensed spicate branches, lax but dense at the tip, axis wholly exposed, slender, straight, glabrescent, dark wine-red; primary Bra leaf-like, linear-triangular or ovate-lanceolate with pungent tip, patent or reflexed, the lower longer than the subtended branch, the upper shorter, coriaceous, both faces subdensely lepidote, margin serrate; floral Bra triangular,  15  10 mm, clasping the flowers, coriaceous, with pungent tip, prominently veined, carinate, dark wine-red, sparsely lepidote, margins serrate or serrulate; Fl 17 mm, sessile; Sep triangular, 10  4 mm, asymmetrical, coriaceous, dark wine-red, glabrous, entire, adaxial ones carinate, abaxial one ecarinate; Pet elliptic, 13  5 mm, obtuse, green with white blades, apically spreading to recurved, with 2 fimbriate appendages well above the base and conspicuous callosities; St included, exceeding the stigma; outer Fil free, inner Fil adnate

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to the petals, 8 mm; Anth 3–4 mm, elliptic; Ov trigonous, 3 mm, green; Sti simple, erect; Fr and Se not known. O. braunii Leme (Pabstia 5(1): 14–15, 1994). Type: Brazil, Bahia (Pereira & Braun E343 [HB]). — Distr: E Brazil (Bahia); on quartzitic sandstone rocks and sandy soils, 400–1250 m. [1] Ros distinctly elongate, caulescent, 20 cm at flowering time; L 10, suberect to curved, rosulate, sheath inconspicuous, outer face furrowed, L lamina narrowly triangular, 13  2.5 cm, canaliculate, leathery, abaxially distinctly veined, both faces greenish and very densely white-lepidote with strongly appressed membranous scales, upper face becoming glabrescent to the long-caudate tip, margins laxly spinose, Sp 3–5 mm; Inf to 16 cm, erect; peduncle 6 cm, greenish, white-lanate; peduncular Bra leaf like; fertile Inf part 8 cm, dense near the tip, with 7 laxly arranged spicate branches at the base with 2 congested flowers each; primary Bra suberectly curved, lower ones leaf-like and  5 longer than the subtended branch, upper ones somewhat leaf-like and 2 as long as the subtended branch; floral Bra ovate,  20  9 mm, acuminate, green, both faces white-lepidote, veined, carinate or ecarinate, laxly spined with 1.5–2 mm long hooked recurved spines; Fl subsessile, 27 mm; Sep lanceolate, 13  4 mm, almost free, subsymmetrical, slenderly acuminate, whitish, white-lepidote, carinate, margins near the tip very minutely retrorsely spinulose; Pet subspatulate with broadly acute and minutely apiculate tip, 21  5 mm, pale greenish but white towards the tip, with 2 fimbriate appendages above the base and 2 elongate acute calli 10 mm long; St details not described; Anth 2 mm, obtuse; Ov broadly obovoid, 5  4 mm, white, white-lepidote; Sty not described; Sti simple, erect, lobes free, greenish; Fr and Se unknown. O. diamantinense Leme (J. Bromeliad Soc. 58(6): 257–259, ills., 2008). Type: Brazil, Minas Gerais (Leme & al. 1825 [HB]). — Distr: E Brazil (Minas Gerais); among rocks and in rock crevices in full sun or more often partial shade, 1070–1100 m. I: Leme & al. (2017: fig. 14D).

U. Eggli and E. J. Gouda

[2] Ros acaulescent, with short basal rhizomatous offsets, plants 30–37 cm tall at flowering time; L 6, laxly rosulate but forming a distinct rosette before anthesis, sheath inconspicuous, subreniform, 2.7  5.5 cm, strongly corrugate, densely white-lepidote towards the tip and abaxially, thick, L lamina sublinear-attenuate, 35–41  2.2–3.8 cm, long caudate, thick and strongly coriaceous, subspreading-arcuate, distinctly canaliculate mainly towards the base, dark red, adaxially glabrous, abaxially conspicuously veined and densely appressedly and coarsely whitelepidote, margins densely spinose at the base but more laxly so towards the tip, Sp narrowly-triangular, spreading (basal ones) to antrorse-uncinate (apical ones), 2–3 mm, 3–10 mm apart; Inf pedunculate, erect, without forming adventive rosettes; peduncle 22–28 cm, greenish to reddish bronzecoloured, densely white-lanate; peduncular Bra not distinct from the rosette leaves, well-spaced; fertile Inf part subellipsoid to capitate, 4.5–6  3.5–4.5 cm, branched with 5–6 congested glomerate branches; primary Bra spreading or nearly so, the lowest many times longer than the fascicles, canaliculate, the basal ones resembling the peduncular bracts, the upper ones with a broadly ovate sheath and a long, narrowly subtriangular to sublinear-caudate or nearly subulate pungent lamina 5–26  1.8–3 cm, densely and coarsely whitelepidote abaxially mainly towards the base, glabrescent to glabrous adaxially, greenish at the base and dark red towards the apex, conspicuously veined abaxially, densely to laxly spinose; Inf branches densely 3- to 5-flowered, sessile; floral Bra ovate-triangular, 17–25  8–17 mm, acuminate with a short mucro, densely spinulose towards the apex, distinctly carinate mainly towards the apex, glabrescent except the white-lepidote to -lanate tip; Fl 38–41 mm, sessile, tubular, with inconspicuous epigynous tube, unscented; Sep suboblong, 18–23  4–8 mm, acute and shortly acicular-mucronulate, subsymmetrical, green except for the hyaline membranous entire to crenulate margins, glabrous except for the white-lanate tip, adaxial ones alate-carinate, abaxial one ecarinate; Pet sublinear-subspatulate, 30–33  5–7 mm, obtuse-cucullate, erect, green except for the white apex, with 2 irregularly shaped broadly

Orthophytum BROMELIACEAE

laminate laciniate-crenulate appendages 4–5 mm above the base, and with 2 oblong callosities; outer Fil 25–26 mm, inner Fil 24 mm and adnate to the petals for 16–17 mm; Anth subellipsoid to sublinear, 3.5–4 mm; Ov 6–8  5–6 mm, trigonous, glabrous, green; Sty not described; Sti weakly conduplicate, 1.5 mm ∅; Fr trigonous; Se reniform, dark brown. Compared with O. mello-barretoi and the non-succulent O. graomogolense in the protologue, and differing mostly in leaf characters. O. eddie-estevesii Leme (J. Bromeliad Soc. 50 (2): 55–57, ills., cover-ill., 2000). Type: Brazil, Minas Gerais (Esteves Pereira 499 in Lemé 4693 [HB, UFG]). — Lit: Braun & Esteves Pereira (2009: 60-62, with ills.). Distr: E Brazil (Minas Gerais); rocky soils in Campo Rupestre grasslands, 1300 m. I: Leme & al. (2017: fig. 14E). [2] Ros shortly caulescent, with slender stolons to 10 cm, 17–19 cm tall at flowering time; L subdensely arranged, recurved, sheaths inconspicuous, L lamina narrowly-triangular, 10–15   2.5 cm, attenuate-caudate, strongly coriaceous, distinctly canaliculate, completely covered by a thick layer of white trichomes on both faces, margins densely spinose, Sp narrowly triangular, nearly straight to slightly retrorse, densely whitelepidote except for the yellowish apical portion, 3  1 mm; Inf erect, to 15 cm; peduncle 6–7 cm, densely white-lanate, green; peduncular Bra leaflike, progressively shorter upwards, almost completely covering the axis; fertile Inf part subcorymbose, branched with 5 densely arranged nearly sessile glomerate branches, ellipsoid in overall outline, 5–6  2 cm ∅; primary Bra leaf-like with broadly ovate-triangular base, long-caudate, the basal ones distinctly exceeding the subtended branches, the others gradually shorter and only slightly longer than the subtended branch, margins densely spinose; Br 4  1.5 cm in outline, densely 2- to 3-flowered; floral Bra of the fascicles triangular, 22–23  12 mm, acuminate, densely spinulose towards the tip, carinate, reaching the middle of the sepals, greenish-hyaline towards the base and brightly orange-red at the apex, veined, membranous, sparsely white-lepidote, floral Bra of the

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simple apical inflorescence part broadly oblong, 30  20 mm, equalling of the sepal length, irregularly curved, ecarinate, with pungent tip, densely spinulose; Fl 45 mm, sessile, with a 1.5 mm long epigynous tube, unscented; Sep narrowly lanceolate, 30  6 mm, acuminate, entire, reddish-orange towards the apex, submembranous, glabrous, carinate; Pet sublinearspatulate, 36  5 mm, strongly obtusecucullate, erect at anthesis, greenish-yellow, with 2 densely fimbriate appendages 8 mm above the base and 2 conspicuous longitudinal callosities which nearly equal the anthers; Fil 26 mm, the inner adnate to the petals for 20 mm; Anth 3 mm, base obtuse, apex acute; Ov trigonous, 9 mm; Sty long-filiform; Sti simpleerect, 2.5 mm ∅, lobes densely papillose, recurved; Fr and Se unknown. This is probably the most attractive species of Subgen. Clavanthus due to the conspicuously white-lepidote leaves and the contrasting reddish-orange sepals and the greenish-yellow petals. It is similar to O. mello-barretoi. O. harleyi Leme & M. Machado (J. Bromeliad Soc. 56(3): 108–110, ills., 2006). Type: Brazil, Bahia (Machado 180 [HB, CEPEC]). — Distr: E Brazil (Bahia); granite rock outcrops, on rocks or between shrubs, 700–830 m. [1] Ros acaulescent, with slender stolons to 6 cm, 13–15 cm tall at flowering time; L 25–35, densely rosulate but becoming well-spaced at flowering time, sheath inconspicuous; L lamina narrowly triangular-attenuate, 10–11  1.3–1.9 cm, attenuate-caudate, fleshy-coriaceous, suberectrecurved to spreading-recurved, nearly flat near the base to slightly canaliculate towards the tip, densely and coarsely white-lepidote on both faces mainly towards the base, trichomes (almost) totally obscuring the greenish to reddish-brown colour, veined mainly abaxially, margins subdensely spinose, Sp triangular, slightly retrorseuncinate, 3–4 mm, 3–7 mm apart; Inf 12 cm, erect; peduncle 6.5 cm, densely white-lanate, pale green but the colour almost completely obscured by the trichomes; peduncular Bra leaflike, suberect-arcuate to nearly spreading, not completely covering the axis; fertile Inf part

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simple, densely congested, 4.5–5.5 cm, densely 25- to 35-flowered, without vegetative shoots; basal floral Bra like the upper peduncular bracts, almost leaf-like, much exceeding the flowers, upper ones ovate-triangular, 24–35  8–13 mm, acuminate-caudate, ecarinate, exceeding the sepals, greenish towards the base, reddish-brown towards the apex, colour in part obscured by a dense layer of coarse white trichomes on both faces, veined, thinly coriaceous, strongly recurved, margins subdensely to densely spinulose; Fl 28–34 mm, sessile or nearly so, suberect, tubular with spreading-ascending limb, with an epigynous tube 0.5 mm; Sep sublinearlanceolate, 14–16  3–5 mm, acuminate, shortly caudate, entire, pale green, veined, membranous mainly along the margins, white-lanate at the base, subdensely to densely white-floccose towards the apex, shortly fimbriate, obtusely carinate to carinate; Pet sublinear-spatulate, 24–28  4–6 mm, subobtuse and inconspicuously apiculate, white towards the apex, with 2 densely and irregularly scalloped and long-lacerate predominantly downwardly oriented appendages 2.5–3 mm above the base and 2 conspicuous longitudinal callosities which nearly equal the filaments; outer Fil 20–21 mm, inner Fil 18 mm, adnate to the petals for 12–13 mm; Anth 2–2.5 mm; Ov trigonous, 3–5  4–5 mm, whitelanate; Sti simple-erect, 1 mm ∅, lobes suborbicular, remotely crenulate; Fr globose, 7 mm ∅, white-lanate, greenish; Se reniform, black-purple, striate. Compared with O. braunii and the non-succulent O. saxicola in the protologue. According to the original description, flowers in cultivated plants opened in late afternoon and stayed open throughout the night — unique for the genus as far as known. O. jacaraciense Leme (Selbyana 30(2): 144–145, ills., 2010). Type: Brazil, Bahia (Reis s.n. in Leme 6987 [HB]). — Distr: E Brazil (Bahia); on interior sand dunes formed from decomposed quartzite rocks, sun-exposed at the fringe of shrubs, 900–1000 m. – Fig. 1. [1] Ros acaulescent, elongating at flowering time and plants then 20–22 cm tall; L 5 at flowering time, in a distinct rosette before

U. Eggli and E. J. Gouda

Fig. 1 Orthophytum jacaraciense (type collection). (Copyright: E. M. C. Leme)

flowering, sheath inconspicuous, L lamina narrowly lanceolate, 1–12.5  2 cm, acuminatecaudate, strongly coriaceous, suberect-arcuate to nearly spreading, nearly flat near the the base to canaliculate towards the apex, yellowish-green or sometimes reddish towards the apex, dull, finely veined abaxially, subdensely white-lepidote at the base mainly adaxially and glabrous towards the apex on both faces, margins straight, subdensely to densely spinose, Sp narrowly triangular, mostly retrorse-uncinate, pale castaneous towards the apex, 2–3 mm, 3–7 mm apart; Inf erect, 18 cm; peduncle 12 cm, erect, light green, finely and sparsely white-lanate to glabrescent; peduncular Bra like the rosette leaves, spreading-arcuate, slightly decreasing in size towards the inflorescence tip; fertile Inf part branched, 6.5 cm, branches 3, laxly arranged, strongly condensed and shortly spicate, 0.5–1.5 cm apart, densely 2to 3-flowered, tip of the main axis similarly

Orthophytum BROMELIACEAE

fasciculate-rosulate, to 3 cm, densely 8-flowered; primary Bra spreading-arcuate to reflexed, nearly flat, basal ones resembling the upper peduncular bracts, 3 as long as the subtended branches but gradually shorter towards the inflorescence tip, light to yellowish-green, sometimes reddish towards the apex, finely veined abaxially, glabrous, densely to subdensely spinulose; floral Bra ovate to ovate-subtriangular or broadly subtriangular, those of the branches 21–22  12–13 mm, those of the main axis 30–32  18 mm, acuminate, slightly pungent, equalling to exceeding the sepals, strongly suberect-recurved, finely veined, glabrous, margins densely spinulose; Fl 34–36 mm, sessile, narrowly tubular, unscented, with inconspicuous epigynous tube; Sep narrowly lanceolate, 17  5 mm, acuminate-caudate, entire, green, glabrous, the adaxial ones carinate, the abaxial one ecarinate; Pet narrowly sublinear-spatulate, 27–28  4–5 mm, obtuse to obtuse-emarginate, slightly cucullate, pale green except for the white apex, with 2 thick scalloped appendages 4 mm above the base and 2 conspicuous longitudinal callosities slightly shorter than the filaments; Fil greenish, 20–22 mm, the inner adnate to the petals for 13 mm; Anth 2–2.5 mm; Ov 5  7 mm, trigonous, glabrous; Sti conduplicate, 2 mm ∅, white, lobes obovate, obtuse, suberect, finely and shortly scalloped-lacerate; Fr and Se unknown. Compared with the non-succulent O. maracasense in the protologue. O. lemei E. Pereira & I. A. Penna (Bol. Mus. Bot. Munic. [Curitiba] 62: 3–4, t. 4 (p. 8), 1985). Type: Brazil, Bahia (Leme & al. 440 [HB]). — Distr: E Brazil (Bahia, Minas Gerais); arid stony ground, 600–1100 m. [1] Ros largely absent, stoloniferous, plants 50 cm tall at flowering time; lowermost L to 26 cm, sheath inconspicuous, 1  4 cm, membranous, glabrous, transversely pleated, margins very minutely spinose, L lamina very long and narrowly triangular, 25  3.5 cm, acuminate, with pungent tip, thick and coriaceous, dark green, canaliculate, both faces densely white-lepidote with scales forming an appressed membrane, abaxially densely veined with prominent veins,

1057

margins spinose, Sp to 3 mm, retrorse; Inf erect, branched; peduncle erect, 15 cm, red, at first white-lanate but becoming glabrous; peduncular Bra similar to the basal leaves but without sheath; fertile Inf part lax, 30 cm, rachis red, first whitelanate but later glabrous; Br condensed-spicate; lower primary Bra similar to the peduncular bracts, much longer than the subtended branches, upper ones ovate-acuminate, much shorter than the spikes; Br condensed strobilate spikes, subglobose, 2  1.5 cm ∅, subsessile; floral Bra ovate, 10 mm, acuminate-spinose, reaching the tips of the sepals, keeled, veined, glabrous, winered, margins almost entire or pectinately spinose; Fl 10–15 mm, without epigynous tube; Sep ovateto narrowly triangular, 7–10  3 mm, glabrous, red, adaxial pair at first strongly keeled and later winged, abaxial one ecarinate; Pet spatulate, 12 mm, blunt, green with white lobes, with 2 strongly fimbriate appendages 4 mm above the base, without callosities; St 8 mm; Anth 2 mm; Ov subglobose, 3 mm, not further described; Fr globose; Se reniform. Flower description completed from Louzada (2012). O. mello-barretoi L. B. Smith (Bol. Mus. Nac. Rio de Janeiro, Bot. ser. 2, 15: 2, t. 1, figs. c-e, 1952). Type: Brazil, Minas Gerais (Mello Barreto 2121 [R, US [photo]]). — Lit: Leme (2008). Distr: E Brazil (Minas Gerais); among rocks in campo, 1000–1400 m. I: Leme & al. (2017: fig. 14C). [2] Ros somewhat elongating at flowering time, 10–20 cm tall, with stolons; L to 25 cm, sheath elliptic, 4 cm, L lamina arching-recurved, very narrowly triangular, with pungent tip, strongly canaliculate, adaxially densely and coarsely whitelepidote near the base and subdensely whitelepidote and soon glabrescent towards the tip, margins laxly spinulose, Sp straight or curved, 2 mm, 5–10 mm apart; Inf branched; peduncle erect, 3.5–16 cm, densely white-lanate; peduncular Bra not distinct from the rosette leaves but more laxly arranged; fertile Inf part very densely subglobose or slightly interrupted at the base, 3–5 cm ∅, with congested glomerate branches, main axis head-like, densely flowered; primary

1058

Bra leaf-like, all but the uppermost with a long lamina; floral Bra broadly ovate, 12–15  8–10 mm, acuminate, straight, erect or nearly erect, slightly shorter than the sepals, serrulate, with conspicuously white-lanate tip; Fl with almost no epigynous tube; Sep (sub-) oblong, 17–20 mm, acute and mucronate, densely white-lanate at the tip, serrulate; Pet oblong-elliptic, 25 mm, obtuse, white, with 2 fimbriate appendages 7 mm above the base; inner Fil high-connate with the petals; Ov subglobose, 6 mm, strongly compressed, glabrous; Fr subglobose; Se black-purple, striate. The type locality is Jaboticatubas in the Serra do Cipó according to Leme & Coelho de Paula (2008). — The recently described O. piranianum is similar according to its protologue. O. piranianum Leme & C. C. Paula (J. Bromeliad Soc. 58(2): 112–116, ills., 2008). Type: Brazil, Minas Gerais (Leme & Paula 7189 [HB, RB]). — Distr: E Brazil (Minas Gerais: Grão Mogol); sandy to rocky soils on quartzitic sandstone outcrops in Campo Rupestre vegetation, full sun or partial shade, 1300 m. I: Leme & al. (2017: fig. 14G). [2] Ros stemless, desintegrating-elongating at flowering time and plants to 19 cm tall, offsetting with basal rhizomes, without new shoots from the inflorescence; L 12, laxly arranged, sheath inconspicuous, strongly corrugate, L lamina narrowly triangular-attenuate, 15–16  3 cm, longcaudate, coriaceous, subspreading-recurved, distinctly canaliculate esp. when water-stressed, both faces coarsely cinereous-lepidote obscuring the bronze leaf colour, abaxially veined, margins (sub-) densely spinose, Sp mostly spreading-retrorse, 2–3 mm, 3–7 mm apart; Inf erect; peduncle 9 cm, pale bronze-coloured, densely and finely white-lanate; peduncular Bra not distinct from the rosette leaves; Inf branched but remaining congested-condensed and capitate in outline,  4  3.5 cm, rachis not visible; primary Bra spreading-recurved, longer than the subtended fascicles, with a suborbicular base and a narrowly triangular-attenuate canaliculate lamina similar to the rosette leaves; Br 4 cm, suberect, sessile, 2.7  2.5 cm in outline, 4- to 5-flowered; floral Bra ovate-triangular, 20–24  12–16 mm,  as

U. Eggli and E. J. Gouda

long as the sepals but recurved, acuminate with short acicular mucro, distinctly carinate, green, finely veined, abaxially densely and coarsely white-lepidote throughout, adaxially densely and coarsely white-lepidote towards the apex, margins densely and coarsely spinose towards the tip; Fl 31–33 mm, narrowly tubular, with inconspicuous epigynous tube, unscented; Sep narrowly triangular-lanceolate, 18  4.5–5 mm, acuminate and shortly acicular-mucronulate, entire, pale green except for the hyaline membranous margins, subdensely to densely and coarsely white-lepidote abaxially, the adaxial ones alate-carinate, the abaxial one ecarinate; Pet sublinear-subspatulate, 24–25  4.5–5 mm, obtuse-cucullate, claw green, limb white, with 2 irregularly lacerate-crenulate obovate to suborbicular appendages 4 mm above the base and 2 conspicuous longitudinal callosities much shorter than the inner filaments; Fil greenish, outer 17 mm, inner 16 mm, adnate to the petals for 9 mm; Anth suboblong, 2.5 mm; Ov  5–6  4.5 mm, trigonous, white, densely sublanate; Sti 1.5 mm ∅, white, lobes weakly conduplicate, densely papillose; Fr and Se unknown. Similar to O. mello-barretoi, but differing by the densely white-lepidote leaves and floral bracts without glabrous tip, different sepals and a whitesublanate ovary. O. piranianum grows sympatrically with the hardly succulent O. graomogolense, from which it is not easily distinguished. O. riocontense Leme (J. Bromeliad Soc. 55(4): 156–158, ills. (p. 162), 2006). Type: Brazil, Bahia (Reis jr. s.n. in Leme 5787 [HB]). — Distr: E Brazil (Bahia); on quartzitic sandstone rock outcrops in cavities of steep rocks, 700–1200 m. [1] Ros acaulescent, elongating-disintegrating at flowering time and plants then 40–45 cm tall, basally offsetting, without new shoots from the inflorescence; L 10, subdensely arranged, suberect-arcuate, sheath inconspicuous, L lamina narrowly triangular, 28–32  4 cm, long-caudate, distinctly succulent, distinctly canaliculate esp. towards the tip, adaxially densely white-lepidote near the base but glabrous otherwise, abaxially densely appressedly white-lepidote completely obscuring the leaf colour and forming a membrane, distinctly veined, margins (sub-) densely

Orthophytum BROMELIACEAE

spinose, Sp distinctly uncinate-retrorse, yellowish to pale brown, 2–3 mm, 4–11 mm apart; Inf erect; peduncle dark red, white-lanate; peduncular Bra not distinct from the rosette leaves, slightly reduced in size further up; fertile Inf part branched, 22 cm; Br 10–11, as condensed sessile subglobose-strobilate spikes 1.7–2  2–2.5 cm ∅, 8- to 15-flowered, laxly (near the base) to densely (more distally) arranged, 1–4 cm apart, at the tip of the main axis forming an inconspicuous head of 2 or 3 fascicles, rachis dark red, white-sublanate to glabrous; primary Bra straight or nearly so, spreading or the upper ones reflexed, slightly canaliculate to nearly flat, the basal ones resembling the peduncular bracts, distinctly longer than the subtended fascicles but the distal ones abruptly shorter, the upper ones with a triangular or broadly ovate-triangular base and a long-acuminate lamina, 2–6  1.3–1.6 cm, distinctly longer to slightly shorter than the subtended fascicles, glabrous adaxially, densely white-lepidote abaxially, green (basal ones) to reddish (upper ones), distinctly veined abaxially, densely to laxly spinulose; floral Bra ovate-triangular, 12–15  8–11 mm, acuteapiculate, densely spinulose, basal ones carinate, the other ecarinate, with strongly recurved tip, dark red, finely veined, glabrescent to glabrous; Fl 19–23 mm, densely arranged, tubular with hardly spreading-ascending limb, with inconspicuous epigynous tube, unscented; Sep slightly asymmetrical, broadly ovate, 9–11  5–5.5 mm, entire, red except for the hyaline margins, glabrous, tip with a distinct slightly pungent mucro 1.5 mm long, adaxial ones alate-carinate; Pet subspatulate, 15–17  4–4.5 mm, rounded to inconspicuously cucullate-emarginate, base whitish, central portion green, distally white, with 2 densely scalloped-fimbriate appendages 4 mm above the base and 2 conspicuous longitudinal callosities almost extending to the anthers; Fil white near the base, greenish near the tip, outer 11 mm, inner 9 mm, adnate to the petals for 5 mm; Anth green, 2.5 mm; Ov 4 mm, subtrigonous, greenish-white, glabrous; Sti 1 mm, white, with reniform erect lobes; Fr and Se unknown. Compared with O. maracasense (not considered to be succulent) in the protologue, differing, amongst other characters, by the strongly

1059

succulent leaves. O. riocontense is amongst the most succulent members of the genus. O. rubrum L. B. Smith (Smithsonian Misc. Collect. 126: 34, 180, Fig. 83, 1955). Type: Brazil, Bahia (Foster 2444 [US]). — Distr: E Brazil (Bahia); in Caatinga vegetation on a table mountain,  600 m. [1?] Ros distinct but elonganting at flowering time and plants 50–60 cm tall; L numerous, covered at first with white appressed scales, sheaths suborbicular, 2–3 cm, pale brown, becoming glabrous and glossy, L lamina linear-triangular, 55–60  2–3 cm, long-attenuate and subfiliform-acute, adaxially glabrescent and dark glossy green, abaxially cinereous-lepidote, margins laxly serrate; Sp antrorse, greenish or brownish, 2 mm; Inf densely conglomerately compound; peduncle elongate, densely white-lepidote, often slightly curved; peduncular Bra leaf-like, divergent to spreading; fertile Inf part with 3–5 spikes, ellipsoid to globose in outline, densely many-flowered, 2–4  2.5 cm ∅, red, sometimes the lowest spike somewhat remote from the others; primary Bra spreading, lower ones leaf-like, upper soon shorter and triangular, upper ones  as long as the spikes, with pungent tip; floral Bra broadly ovate, 20 mm, acuminate, bright pinkish-red, the lower ones divergent, the upper suberect, all with pungent tip, veined abaxially with lines of white trichomes, soon glabrous; Fl narrowly tubular and hardly opening; Sep triangular, 12 mm, mucronulate, the adaxial ones very broadly alatecarinate, green to red at the tip; Pet 15 mm, white, with 2 appendages well above the base; St included; Ov subglobose, yellowish-green. Treated as dubious and not recollected since the first description, which was based on cultivated material (Louzada 2012). O. schulzianum Leme & M. Machado (J. Bromeliad Soc. 55(4): 175–177, ills., 2006). Type: Brazil, Minas Gerais (Machado & Schulz s.n. in Leme 5881 [HB]). — Distr: E Brazil (C Minas Gerais; Serra do Espinhaço: Diamantina plateau); rock outcrops, 1260 m. – Fig. 2. [2] Ros subdensely few-leaved, elongatingdisintegrating at flowering time and plants to

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Fig. 2 Orthophytum schulzianum. (Copyright: E. M. C. Leme)

19 cm tall, with slender stolons; L spreading to spreading-ascending, sheath inconspicuous, L lamina narrowly triangular, 12  3 cm, attenuate-caudate, coriaceous, distinctly canaliculate with upright-incurved margins, completely covered by a thick layer of coarse white trichomes on both faces, distinctly veined abaxially, margins densely spinose, Sp narrowly triangular, nearly straight to slightly retrorse, 2 mm, 2–3 mm apart; Inf erect; peduncle 10 cm, densely white-lanate, green; peduncular Bra leaf-like but gradully reduced in size further up, suberect; fertile Inf part densely branched and subcorymbose, ellipsoid-capitate, 5–6  3 cm ∅, with 5 strobilate sessile glomerules, each 2- to 3-flowered; primary Bra suberect, completely covered by a dense thick layer of coarse white trichomes on both faces, greenish-white near the base, slightly darker towards the apex, the basal ones  leaf-like, much longer than the fascicles but gradually

U. Eggli and E. J. Gouda

shorter higher up, uppermost slightly longer than the fascicles from a broadly ovate-triangular base, 2.8  2.5 cm, margins densely spinose; floral Bra suberect, greenish-white, veined, densely and coarsely white-lepidote mainly abaxially, those of the fascicles narrowly triangular, acuminate, strongly carinate, slightly shorter than the sepals, membranous, 29  13 mm, densely spinulose towards the tip, those of the simple main axis narrowly triangular to broadly ovate, longapiculate, ecarinate, slightly shorter to equalling the sepals, 31–35  10–16 mm, densely spinulose; Fl 46 mm, erect, narrowly tubular, without epigynous tube, unscented; Sep subsymmetrical, narrowly lanceolate, 26–27  7 mm, acuminate, entire to remotely denticulate near the apex, greenish-white, membranous, densely to subdensely and coarsely white-lepidote except for the white-lanate apex, carinate; Pet sublinearspatulate, 37  6 mm, obtuse-cucullate, green except for the white apex and apical margins, with 2 laminate obovate and apically denticulate appendages at the base and 2 conspicuous longitudinal callosities 25 mm long; outer Fil 31 mm, inner Fil 28 mm, adnate to the petals for 20 mm; Anth 3.5 mm; Ov 7 mm, trigonous: Sti simple-erect, 1 mm ∅, lamina green near the base, white towards the apex, suberect; Fr and Se unknown. Compared with O. eddie-estevesii and O. mellobarretoi in the protologue. Versieux & al. (2010: 13) argue that the species is probably closest to O. diamantinense, and that the differences need further evaluation. O. striatifolium Leme & L. Kollmann (J. Bromeliad Soc. 57(4): 152–154, ills. (p. 151, 153), 2007). Type: Brazil, Espírito Santo (Kollmann & al. 7085 [MBML, HB]). — Distr: E Brazil (Espírito Santo); rock outcrops in the Atlantic Forest, in shallow organic soils in full sun. – Fig. 3. [1] Ros subdensely 10- to 14-leaved, elongating-disintegrating at flowering time and plants then to 13 cm tall, with slender rhizomes 5 cm long, without new shoots from the inflorescence; L sheath inconspicuous, L lamina narrowly triangular-lanceolate, 8–11  1.4–1.5 cm, long attenuate-caudate, thickly coriaceous, arcuate to

Orthophytum BROMELIACEAE

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arranged, narrowly tubular with distinctly spreading-recurved limb, with inconspicuous epigynous tube, unscented; Sep symmetrical or nearly so, narrowly lanceolate, 15–16  4–5 mm, attenuate towards the acuminate-caudate apex, entire, greenish-yellow, veined, glabrous, the adaxial ones carinate, the abaxial one ecarinate; Pet sublinear-spatulate, 27–28  5–6 mm, obtuseemarginate, very minutely apiculate-caudate, white except for the pale greenish central portion, with 2 densely and irregularly fimbriate-scalloped appendages 3.5 mm above the base and 2 conspicuous longitudinal callosities reaching the filaments; outer Fil 18 mm, inner Fil 17 mm, adnate to the petals for 11–12 mm; Anth 2.5–3 mm, green; Ov  5  4.5–5 mm, trigonous, greenish-white; Sti simple-erect, 2 mm ∅, lobes spreading-recurved, white; Fr and Se unknown. Compared with the non-succulent O. estevesii in the protologue. O. toscanoi Leme (J. Bromeliad Soc. 53(1): 23–24, 28, ills., cover ill., 2003). Type: Brazil, Bahia (Reis s.n. [HB]). — Distr: E Brazil (Bahia: Poço da Moça); amongst rocks in rather humid Fig. 3 Orthophytum striatifolium. (Copyright: E. M. forest, 500–900 m. C. Leme) [1] Ros acaulescent, subdensely 12-leaved, with short basal offshoots, 30 cm tall at spreading, canaliculate mainly towards the tip, flowering time; L sheath inconspicuous, lamina pale reddish-brown, adaxially subdensely white- narrowly-triangular, 10  2.5 cm, strongly corilepidote with trichomes in rows along the veins, aceous, attenuate-caudate, slightly canaliculate to abaxially densely and coarsely white-lepidote nearly flat, with nearly subulate tip, completely completely obscuring the leaf colour, distinctly covered by a thick layer of white trichomes on veined, margins subdensely spinose, Sp narrowly both faces, abaxially distinctly veined, margins triangular, 1.5–2 mm, 4–7 mm apart, basal ones densely spinose-serrate for most of the length but nearly straight, upper distinctly antrorse-uncinate; entire near the tip, Sp triangular, nearly straight to Inf erect; peduncle 4–8 cm, green but densely slightly retrorse, 3–5 mm, 1–8 mm apart; Inf white-lanate; peduncular Bra similar to the rosette erect; peduncle 13 cm, densely white-lanate, leaves, subspreading-arcuate to reflexed, upper greenish bronze-coloured; peduncular Bra similar ones slightly shorter and congested below the to the rosette leaves but upper ones gradually fertile inflorescence part, fertile Inf part simple, shorter, all white-lanate at the base, exposing the densely capitate-rosulate, 6- to 9-flowered, axis, suberect-arcuate; fertile Inf part corymbose, 2.5 cm, 3–3.5 cm ∅ in outline; floral Bra laxly branched at least towards the base, 15 cm, triangular-ovate, 17–23  10–13 mm, acuminate, Br 8, sessile and shortly rosulate-strobilate distinctly canaliculate, navicular, ecarinate to spikes, 1–1.2  1.5–1.7 cm ∅ in outline, 4- to obtusely carinate, strongly recurved, yellowish- 7-flowered; primary Bra like the upper peduncular green to orange (in full sun), (nearly) glabrous, bracts but gradually smaller, 20–55  12–17 mm, veined, subdensely spinose; Fl 32–34 mm, densely distinctly exceeding the subtended branches but

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gradually shorter further up, basal ones spreading, the upper slightly reflexed and slightly exceeding the subtended branch, margins on the basal (sub-) densely spinose; floral Bra ovate, 10–11  7 mm, acuminate, strongly carinate and recurved, equalling to exceeding the sepals, green, veined, coriaceous, sparsely white-sublanate, subdensely and minutely spinulose; Fl 16 mm, densely arranged, narrowly tubular with hardly flaring limb, with inconspicuous epigynous tube, unscented; Sep subsymmetrical, narrowly ovate, 7  3 mm, tip spinescent-acuminate, entire, green, rigid, sparsely white-sublanate, carinate; Pet sublinear, 11  2–3 mm, obtuse to slightly emarginate and with a hook-like minute apiculus, green except for the white apical part, with 2 densely fimbriate appendages 3 mm above the base and 2 conspicuous longitudinal callosities which nearly reach the filaments; Fil 6 mm, inner adnate to the petals for 3 mm; Anth 1 mm; Ov 3 mm, trigonous; Sti simple-erect, 1 mm ∅, lobes suberect; Fr and Se unknown. Compared with O. lemei in the protologue, but with shorter leaves with longer and more densely arranged marginal spines, shorter inflorescences, and green floral bracts and sepals. O. triunfense J. A. Siqueira & Leme (Fragm. Atlantic Forest N.E. Brazil, 311–312, ill., 2007). Type: Brazil, Pernambuco (Siqueira-Filho 1255 [UFP, HB]). — Distr: E Brazil (Paraíba, Pernambuco); granite outcrops in full sun, 720–1200 m. I: Herndon (2015). [1] Ros acaulescent, subdensely 7- to 8-leaved, elongating-disintegrating at flowering time and plants then 10–16 cm tall, propagating with slender stolons, without new shoots from the inflorescence; L sheath inconspicuous, L lamina narrowly-triangular, 11–20  2–3.5 cm, attenuatecaudate and with pungent tip, strongly coriaceous, suberect-arcuate, distinctly canaliculate, densely and coarsely white-lepidote on both faces with the trichomes almost totally concealing the greenish colour, distinctly veined abaxially, margins (sub-) densely spinose, Sp acicular, nearly straight to retrorse, 2–3 mm, 3–8 mm apart; Inf erect; peduncle 4–6 cm, densely white-lanate, pale green but

U. Eggli and E. J. Gouda

colour almost completely concealed by the trichomes; peduncular Bra similar to the rosette leaves, exposing the peduncle at the base only, suberect, slightly arcuate; fertile Inf part densely branched and subcorymbose, 3.5–4.5  2 cm ∅ in outline, subdensely 12-flowered in total; Br 5, almost sessile spikes, 1.6 cm, each 2flowered; primary Bra leaf-like but slightly shorter, suberect to nearly spreading, slightly arcuate, greatly exceeding the subtended fascicles, subdensely spinose; floral Bra of the fascicles triangular, 13  7 mm, acuminate, densely spinulose, strongly carinate, yellowish, veined, densely and coarsely white-lepidote on both faces, those of the apical part of the main axis more leaf-like, densely spinulose, distinctly exceeding the flowers to about equalling the sepals, suberect, otherwise like the other floral bracts; Fl 26 mm, narrowly tubular, with an epigynous tube 0.5 mm long, unscented; Sep narrowly triangular to ovate-lanceolate, 11  4 mm, acuminate-spinescent, entire, yellowish, submembranous along the margins, densely and coarsely white-lepidote, the adaxial ones carinate, the abaxial one obtusely carinate; Pet sublinearspatulate, 17–20  4 mm, acute and minutely apiculate, basal pale greenish, the apical white, with 2 shortly scalloped appendages 3.5 mm above the base and 2 conspicuous longitudinal callosities which nearly reach the anthers; Fil 9 mm, the inner adnate to the petals for 6 mm; Anth 2 mm, yellowish-green; Ov 5 mm, trigonous; Sti simple-erect, 1 mm ∅, yellowish; Fr and Se unknown. Herndon (2015) compares the taxon with the non-succulent O. magalhaesii, O. disjunctum, O. gurkenii and material traded (but most probably incorrectly identified) as O. leprosum. O. vasconcelosianum Leme (Phytotaxa 205 (4): 287–289, ills., 2015). Type: Brazil, Minas Gerais (Leme & al. 8673 [RB, HB]). — Distr: E Brazil (Minas Gerais: Alvarenga); on rocks on the summits of inselbergs with Campo Rupestre vegetation, 1060–1440 m. I: Leme & al. (2017: fig. 16B-E). [3] Ros long caulescent, stem 20–57 cm long, with 1–2 short shoots produced at the base of the

Orthophytum BROMELIACEAE

inflorescence; L densely and equally spaced along the stem, spreading, sheath inconspicuous, subtrapeziform, 1–1.3  1.8–2 cm, greenish towards the tip, whitish to hyaline towards the base, thintextured, densely white lepidote and densely spinulose near the apex, glabrous and entire towards the base, densely and conspicuously veined, L lamina very narrowly triangular, 5–11  0.8–1.3 cm, with attenuate but not pungent apex, coriaceous, canaliculate with upturned margins mainly when water-stressed, densely and coarsely white-lepidote on both faces, green but the colour completely obscured by the trichomes, abaxially veined, margins densely spinulose, Sp subtriangular, straight to prevailingly antrorse-uncinate, reddish, 0.5–1 mm, 1.5–4 mm apart; Inf sessile at the stem tip, 3–4 cm, corymbose and fasciculately compound at the base but unbranched distally; Br 9–12, densely arranged, nearly sessile, slightly pulvinate, 1.7–1.8 cm, 3- to 7-flowered; primary Bra leaflike, gradually shorter towards the tip of the inflorescence, distinctly exceeding the subtended fascicles; floral Bra of the fascicles narrowly triangular, 10–12  4–8 mm, acute to acuminate, distinctly carinate,  equalling the middle of the sepals, greenish to wine-coloured, veined, densely and coarsely white-lepidote, not pungent, margins entire, floral bracts of the main axis more leaf-like, equalling to exceeding the flowers; Fl 19–24 mm, densely arranged, with inconspicuous epigynous tube, distinctly fragrant; Sep narrowly lanceolate, 9–10  2.5–3 mm, acuminate, entire, greenish except for the wine-red margins, not pungent, coarsely white-lepidote, (obtusely) carinate; Pet lanceolate to narrowly obovate, 15–18  4.5–5 mm, acute to inconspicuously emarginate, subspreading-recurved, lilac-rose, without appendages but with 2 conspicuous longitudinal callosities  as long as the inner filaments, callosities inconspicuously glandulose at the base; outer Fil 8–9 mm, inner Fil 5–6 mm; Anth 2.5–3 mm; Ov obovoid, trigonous, 4–6  3–4 mm, whitish, coarsely white-lepidote; Sty rose-lilac for most of its length; Sti simple-erect, somewhat conduplicate-patent, lobes spreading, short to elongate, 0.7–1 mm, rose-lilac with white tip; Fr globose, 4–7  3.5–6 mm, greenish; Se only 3–5 per fruit, 2.2–3  1–2 mm.

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The species is unique in the genus on the base of several characters and somewhat intermediate between O. zanonii of Subgen. Capixabanthus and Cryptanthus (Subgen. Hoplocryptanthus) glaziovii, with which it shares fragrant flowers and unappendaged petals. The lilac-rose petals and the few-seeded fruits are also notable. O. zanonii Leme (J. Bromeliad Soc. 54(2): 72–73, ills. (incl. pp. 70–71), 2004). Type: Brazil, Espírito Santo (Leme & al. 5930 [HB]). — Lit: Louzada & Wanderley (2010: 23-25, with ills.). Distr: E Brazil (Espírito Santo: Pancas); slightly inclined rock outcrop in deciduous forest or nearby open sunny places, on rocks, 430–550 m; only known from the type locality. I: Leme & al. (2017: fig. 13G). – Fig. 4. [4] Ros forming large and dense clumps, with branched stems 20–50 cm long and 1.5–1.8 cm ∅, propagating by branching and rooting of mature stem sections, as well as new shoots produced at the inflorescence base; L densely and

Fig. 4 Orthophytum zanonii. (Copyright: E. M. C. Leme)

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equally spaced along the stem, spreading-recurved, sheath broadly ovate, 3.5  2.5 cm, palecoloured, densely white-lepidote and densely spinulose near the tip, glabrous towards the base, completely covering the stem, coarsely veined, disintegrating into fibres along older stem parts, L lamina narrowly-triangular, 10–25  1–1.2 cm, attenuate-caudate, with a pungent tip, strongly coriaceous, distinctly canaliculate with upturned margins, adaxially densely white-lepidote, abaxially completely covered by a thick layer of white scales, greenish to reddish but the colour  completely obscured by the white scales, sometimes cross-banded, margins densely spinulose, Sp uncinate, antrorse, reddish-brown, 0.5–1 mm, 3–5 mm apart; Inf sessile, densely compound, corymbose, ellipsoid to subcapitate in outline, 3–3.5  2.5–3 cm ∅; Br 5–8, dense, nearly sessile, complanate, 2–2.2  1.5 cm, 2- to 3-flowered; outer primary Bra transitional to the inner rosette leaves that sometimes become red-flushed during flowering time, gradually shorter nearer the inflorescene tip but distinctly exceeding the branches; floral Bra of the fascicles narrowly triangular, 18–19  6–10 mm, acuminate with a pungent tip, strongly carinate,  equalling the sepals, greenish, veined, densely white-lanate except for the apical spine, margins spinulose, those of the apical part of the main axis more leaf-like, exceeding the flowers, some rudimentary floral bracts present; Fl 25 mm, densely arranged, narrowly tubular and hardly opening, with inconspicuous epigynous tube, unscented; Sep narrowly oblong-lanceolate, 13–14  3–4 mm, acuminate, entire, greenish, rigid but not coriaceous, densely white-lanate, the adaxial ones alate-carinate, the anterior one carinate or nearly so; Pet sublinearspatulate, 18  3.5–4 mm, obtuse-cucullate, green except for the whitish apical margins, with 2 downwardly oriented cucullate crenulate appendages 5 mm above the base and 2 conspicuous longitudinal callosities  extending to the filaments; outer Fil 15 mm, inner Fil 6 mm, adnate to the petals; Anth 3.5 mm, strongly compressed laterally at anthesis; Ov 7  6 mm, trigonous but slightly complanate; Sty 1.7 mm; Sti simple-erect, lobes suberect, densely papillose, greenish; Fr and Se unknown.

U. Eggli and E. J. Gouda

Compared with the non-succulent O. vagans in the protologue but distinct in leaf and inflorescence characters. O. pseudovagans is another similar species with elongate stems.

References Braun, P. J. & Esteves Pereira, E. (2009) Succulent and xeromorphic Bromeliads of Brazil: Part 5: Orthophytum estevesii & O. eddie-estevesii. Cact. Succ. J. (US) 81(2): 59–63, ills. https://doi.org/10.2985/015.081.0202. Evans, T. M. [& al. 2015], Jabaily, R. S., Gelli de Faria, A. P., Oliveira F. de Sousa, L. de, Wendt, T. & Brown, G. K. (2015) Phylogenetic relationships in Bromeliaceae subfamily Bromelioideae based on chloroplast DNA sequence data. Syst. Bot. 40(1): 116–128. https://doi. org/10.1600/036364415X686413. Herndon, A. (2015) Introducing Orthophytum triunfense. J. Bromeliad Soc. 64(4): 264–271, ills. Horres, R. & Zizka, G. (1995) Untersuchungen zur Blattsukkulenz bei Bromeliaceae. Beitr. Biol. Pfl. 69(1): 43–76, ills. Leme, E. M. C. (2004) Studies on Orthophytum, an endemic genus of Brazil – Part I. J. Bromeliad Soc. 54(1): 36–43, ills. Leme, E. M. C. (2008) Studies on Orthophytum — Part IX. The “subcomplex mello-barretoi” and another new species. J. Bromeliad Soc. 58(6): 257–261, ills., key. Leme, E. M. C., Heller, S., Zizka, G. & Halbritter, H. (2017) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318(1): 1–88, ills., maps, key. https://doi.org/10.11646/ phytotaxa.318.1.1 Leme, E. M. C. & Coelho de Paula, C. (2008) Studies on Orthophytum — Part VIII: Two new species from Grão-Mogol State Park, Minais Gerais, Brazil. J. Bromeliad Soc. 58(3): 106–117, ills. Louzada, R. B. (2012) Revisão taxonômica e filogenia de Orthophytum (Bromeliaceae, Bromelioideae). São Paulo (BR): Universidade de São Paulo, unpublished Ph.D. thesis. Louzada, R. B. & Versieux, L. M. (2010) Lapanthus (Bromeliaceae, Bromelioideae): A new genus from the southern Espinhaço Range, Brazil. Syst. Bot. 35(3): 497–503, ills., key. https://doi.org/10.1600/036364410 792495908. Louzada, R. B. & Wanderley, M. G. L. (2010) Revision of Orthophytum (Bromeliaceae): The species with sessile inflorescences. Phytotaxa 13: 1–26, ills., key. Louzada, R. B. [& al. 2010], Palma-Silva, C., Corrêa, A. M., Kaltchuck-Santos, E. & Wanderley, M. G. L. (2010) Chromosome number of Orthophytum species (Bromeliaceae). Kew Bull. 65(1): 53–58. https:// doi.org/10.1007/s12225-010-9175-6.

Orthophytum BROMELIACEAE Louzada, R. B. [& al. 2014], Schulte, K., Wanderley, M. G. L., Silvestro, D., Zizka, G., Barfuss, M. H. & Palma-Silva, C. (2014) Molecular phylogeny of the Brazilian endemic genus Orthophytum (Bromelioideae, Bromeliaceae) and its implications on morphological character evolution. Molec. Phylogen. Evol. 77: 54–64. https://doi.org/ 10.1016/j.ympev.2014.03.007. Schulte, K. & Zizka, G. (2008) Multi locus plastid phylogeny of Bromelioideae (Bromeliaceae) and the taxonomic utility of petal appendages and pollen characters. Candollea 63(2): 209–225. http://www.ville-ge.ch/cjb/ publications_candollea_632.php Schulte, K. [& al. 2009], Barfuss, M. H. & Zizka, G. (2009) Phylogeny of Bromelioideae (Bromeliaceae) inferred from nuclear and plastid DNA loci reveals the evolution of the tank habit within the subfamily. Molec.

1065 Phylogen. Evol. 51(2): 327–339. https://doi.org/10.10 16/j.ympev.2009.02.003. Silvestro, D. [& al. 2014], Zizka, G. & Schulte, K. (2014) Disentangling the effects of key innovations on the diversification of Bromelioideae (Bromeliaceae). Evolution 68(1): 163–175. https://doi.org/10.1111/evo.12236. Smith, L. B. & Downs, R. J. (1979) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 3: Bromelioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/i399940 Versieux, L. M. [& al. 2010], Louzada, R. B., Lage Viana, P., Mota, N. & Wanderley, M. G. L. (2010) An illustrated checklist of Bromeliaceae from Parque Estadual do Rio Preto, Minas Gerais, Brazil, with notes on phytogeography and one new species of Cryptanthus. Phytotaxa 10: 1–16, ills. https://doi.org/10.11646/phytotaxa.10.1.1

Orthotanthus BROMELIACEAE U. Eggli

Orthotanthus L. E. Smith (J. Bromeliad Soc. 38(2): 75, 1983). — Distr: Cultivated only. = Orthophytum  Cryptanthus. Smith (1983) lists the cultivar ‘What’ with the parentage O. saxicola var. rubra  Cryptanthus ‘It’, and Giroux (2009) described the cultivar ‘Blazing Bonsai’ with the parentage O. navioides (now classified as Sincoraea)  C. microglazioui.

References Giroux, L. (2009) An unusual bigeneric hybrid with C. [ryptanthus] microglaziouii [sic!]. Cryptanthus Soc. J. 24(1): 18–19, ills. Smith, E. L. (1983) Bigeneric hybrids — A listing. J. Bromeliad Soc. 33(2): 69–75. http://journal.bsi.org/ V33/2/

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_102

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Puckia BROMELIACEAE N. Schütz and F. Krapp

Puckia D. Butcher (J. Bromeliad Soc. 52: 52, 2002). — Distr: Cultivated only. Incl. Dyckipu hort. (nom. inval., ICN Art. 29.1).

= Puya  Dyckia. The only known cultivar is the hybrid ‘Sparkle’ (= P. laxa (female)  (D. fosteriana  D. platyphylla)). It originated in a nursery in Venice, Florida, and was first invalidly named  Dyckipu.

N. Schütz (*) Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany e-mail: [email protected] F. Krapp Guxhagen, Germany e-mail: fl[email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_103

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Pucohnia BROMELIACEAE N. Schütz and F. Krapp

Pucohnia G. H. Anderson ex D. A. Beadle (Prelim. List. Cult. Grex Bromeliad, 200, 1991). — Distr: Cultivated only. = Puya  Deuterocohnia. The only known cultivar is ‘George Anderson’ (= P. laxa (female)  D. schreiteri) (Anderson 1986: 103). The greenish-grey leaves are said to be longer than

those of either parent. Plants produce numerous offsets at their base.

References Anderson, G. H. (1986) Hybrids by design and by chance. J. Bromeliad Soc. 36(3): 99–103, 109, ills.

N. Schütz (*) Abteilung Botanik, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany e-mail: [email protected] F. Krapp Guxhagen, Germany e-mail: fl[email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_104

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Puya BROMELIACEAE J. M. Manzanares

Puya Molina (Sag. Stor. Nat. Chili, 160, 351, 1782). Type: Puya chilensis Molina. — Puyoideae — Lit: Gilmartin (1972: synopsis Ecuador); Smith & Downs (1974: 66–190, Fl. Neotropica); Gómez & Grau (2009: synopsis Argentina); Jabaily & Sytsma (2010: phylogeny Chile); Jabaily & Sytsma (2013: biogeography and evolution); Zizka & al. (2013: taxonomy Chile). Distr: C America to C-S Chile. Etym: Span. ‘puya’, tip of the lancet used to drive oxen; probably for the pungent leaf tips or the leaf marginal teeth. Incl. Pourretia Ruiz & Pavón (1794) (nom. illeg., ICN Art. 52.1). Type: Pourretia lanuginosa Ruiz & Pavón. Monocarpic or polycarpic rosette-forming dwarf to giant herbs, terrestrial, sometimes on rocks, monocarpic taxa with solitary sessile or single-stemmed rosette, polycarpic taxa with usually offsetting rosettes with short to long branched stems, stems sometimes fleshy; Ros few- or more often many-leaved; L with a distinct wide and usually slightly to strongly fleshy sheath, L lamina narrowly triangular, coriacous to slightly or rarely distinctly succulent, persisting for several

J. M. Manzanares (*) Herbario Nacional (QCNE), Sección Botánica del Museo Ecuatoriano de Ciencias Naturales del Instituto Nacional de Biodiversidad, Quito, Ecuador e-mail: [email protected]

years, young ascending, becoming increasingly spreading to reflexed with age, adaxially usually glabrous, abaxially completely or at least between the veins covered with a cinereous indumentum, tip attenuate, pungent in the majority of the species, margins spinose-serrate, Sp retrorse or antrorse, or both; Inf terminal, usually pedunculate, erect, simple racemes or spikes, or branched, lax or dense, cylindrical or pyramidal in outline, or strobiliform, axes covered with a lanate, ferruginous, tomentose or floccose indumentum, or glabrous, peduncle usually exceeding the leaves; peduncular Bra leaf-like or reduced; lower primary Bra similar to the peduncular bracts or reduced, branches when present stipitate (i.e. with the basal portion sterile), polystichous, dense, subdense or lax; floral Bra longer or shorter than the sepals, imbricate or remote, persistent, entire or serrulate; Fl actinomorphic, hermaphrodite, subsessile or pedicellate, mostly diurnal; Sep free, convolute, entire, symmetrical or  asymmetrical; Pet free, frequently in shades of dark green or purple, without appendages at the base, becoming spirally contorted after anthesis; Ov superior; Fr loculicidal and septicidal capsules; Se alate. The genus is composed of 223 species and 7 varieties of which 55 species and 5 varieties are considered succulent. The degree of succulence is comparable to that of many bona-fide succulents from other families. Usually, succulence is confined to the leaf sheaths, and/or to the fleshy stems. CAM is present in about of

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_94

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all taxa, and has been found to have evolved repeatedly (Jabaily & Sytsma 2010). Classification: Puya was traditionally included in subfamily Pitcairnioideae. All recent molecular phylogenetic studies of the family agree, however, that the genus is sister to the Bromelioideae (Givnish & al. 2007, Schulte & Zizka 2008, Schulte & al. 2009, Jabaily & Sytsma 2010, Givnish & al. 2011). To accommodate this finding, the monotypic subfamily Puyoideae was erected by Givnish & al. (2007: 16). The genus has been found to be monophyletic by Jabaily & Sytsma (2010), while Givnish & al. (2011) found some support that the genus rather represents a paraphyletic grade at the base of the Bromelioideae, but monophyly cannot be rejected. Monophyly is also supported by the diagnostic character of the petals that form a tightly coiled spiral after anthesis. The 2 subgenera traditionally recognized have not been confirmed by the molecular phylogeny of Jabaily & Sytsma (2010), but they are here maintained on practical grounds. Puya represents one of the largest radiations in the family and spans a wide ecological amplitude in a diversity of wet to dry habitats. Its species are often characteristic elements of the Páramo floras. Puya is also a good example of a rapid specieslevel radiation in the Andes (Jabaily & Sytsma 2013): The greatest diversity is found at middle elevations and mid-latitudes in S America. The origin of the genus is in present-day C Chile, and speciation progressed from S to N tracking the final Andean uplift. Pleistocene glaciation cycles are the likely drivers for speciation. Hybrid speciation also appears to play a considerable role (Jabaily & Sytsma 2010, Jabaily & Sytsma 2013: 214). The genus is traditionally divided in 2 subgenera based on the inflorescence architecture: [1] Subgen. Puya: Inf branched, branches sterile in their apical parts. [2] Subgen. Puyopsis (Baker) L. B. Smith 1970: Inf unbranched or branched, fertile over the whole length. Ecology: Puya species native to high-elevation Páramo vegetations in the Andes often have compact

J. M. Manzanares

inflorescences with a dense woolly indumentum. Miller (1994) measured flower temperatures and found them to be above ambient temperatures in the species studied. Hypothetically, the dense indumentum serves for thermal insulation and protects the flowers from freezing damage. Hornung-Leoni & al. (2013) report that the flowers of species of subgen. Puyopsis (e.g. P. coerulea, P. venusta) are visited by specialist nectarivorous hummingbirds, while species of subgen. Puya (e.g. P. chilensis, P. alpestris) are visited by more generalist passerine birds that perch on the sterile distal branch parts to exploit the flowers with their generally highly diluted but abundant nectar. According to Salinas & al. (2007) High Andean Puya species are an important source of nutrients for hummingbirds, at least in Peru. Horticulture: Puya has limited horticultural potential due to the large size of many of its species. The smaller-growing species are sometimes encountered in specialist collections. In cultivation, hybrids with Dyckia (= Puckia, see separate entry) and Deuterocohnia (= Pucohnia, see separate entry) have been obtained. The following names are of unresolved application but are referred to this genus: Pourretia auruacensis Linden (1853); Pourretia lepidota Linden (1853); Pourretia speciosa Linden (1853); Pourretia yuccoides Linden (1871); Puya acris hort. ex Gentil (1907) (nom. inval., ICN Art. 32.1c). P. adscendens L. B. Smith (Phytologia 17: 76, t. 1: figs. 17–18, 1968). Type: Peru, Amazonas (Sagástegui 6073 [TRP, US [photo]]). — Distr: Peru (Amazonas: small area in Chachapoyas); ravines, semi-desert areas, 2800 m. [2] Polycarpic, forming dense groups, 2 m tall at flowering time; L sheath fleshy, lamina 40–60  1.5–2 cm, adaxially glabrous, abaxially covered with appressed cinereous scales, margins laxly serrate with brown antrorse Sp 5 mm long; Inf laxly branched, pyramidal, greenish, except for the petals densely cinereous-tomentose with finely stellate scales, peduncle exceeding the leaves; peduncular Bra leaf-like; primary Bra broadly ovate, 5 cm, with an attenuate apex,

Puya BROMELIACEAE

shorter than the subtended branches but slightly exceeding the naked sterile branch bases; Br divergent, ascending, 15–30 cm, laxly flowered with  secund flowers; floral Bra broadly ovate, 20 mm, with an attenuate brown apex, covering the lower of the sepals; Ped rather slender, 5 mm; Sep oblanceolate, 23 mm, broadly rounded and mucronulate; Pet green-bluish, 45 mm; St included; Anth yellow; Sty as long as the petals; Sti yellow. As in many other Andean species, the inflorescence is totally covered by dense cinereous scales, and only the petals are glabrous. Only few inflorescences are present in a dense group of plants. The flowers are pollinated by hummingbirds. P. aequatorialis André (Énum. Bromél., 5, 1888). Type: Ecuador, Imbabura (André 3564 [K]). — Distr: N Ecuador (Carchi, Imbabura, Pichincha, Azuay, Loja); dry inter-Andean valleys, 2200–2400 m. [2] Polycarpic, forming dense groups, 1.5–2 m tall at flowering time, stem 40  20–25 cm, branched; L sheath 5.5–4.5 cm, fleshy, lamina 35–50  1.5–2 cm, adaxially green, abaxially with appressed cinereous scales, margins serrate with yellow retrorse hooked Sp 5 mm long; Inf peduncle exceeding the leaves, 60–90 cm, floriferous part 50–100 cm, simple, cylindrical, lax, red-violet or green, glabrous; peduncular Bra small, exposing most of the peduncle; Br divergent, 30–50 cm, laxly flowered; floral Bra ovate, 9–10 mm, attenuate towards the tip, much shorter than the sepals, margins entire; Ped rather slender, to 15 mm; Sep linear, 5 mm, broadly rounded and apiculate; Pet white, green or violet, 45 mm; St included; Anth yellow; Sty included; Sti surrounded by the anthers. Very common in the Andes of N Ecuador. Natural hybrids between green- and violet-flowered populations result in an array of different flower colours from green to violet. The plants are used locally to plant fences. This is also one parent of the natural hybrid P. pichinchae (see there). P. aequatorialis var. aequatorialis — Distr: N Ecuador (Carchi, Imbabura, Pichincha, Azuay, Loja); dry inter-Andean valleys, 2200–2400 m. I:

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Oliva-Esteve (2002: 228); Manzanares (2005: 304–305). [2] Floral Bra merely reaching to the base of the sepals, to 10 mm; Ped 2–6 mm at anthesis; Pet green or violet. P. aequatorialis var. albiflora André (Énum. Bromél., 5, 1888). Type: Ecuador, Pichincha (André 3596 [K]). — Distr: N Ecuador (Pichincha). I: Gilmartin (1972: Fig. 8); Manzanares (2005: 306).  Puya aequatorialis fa. albiflora (André) Gouda (s.a.) (nom. inval., ICN Art. 29.1). [2] Floral Bra exceeding the base of the sepals, to 22 mm; Ped 9–15 mm at anthesis; Pet white. White-flowering plants commonly grow among populations with violet flowers. Populations from the area of the Río Pisque (Pichincha) have longer and more slender peduncles than those registered elsewhere. Gouda & al. (2012+) treat this variety as a mere form of P. aequatorialis. P. alpestris (Poeppig) Gay (Fl. Chil. 6: 12, 1853). Type: Chile, Bío Bío (Poeppig 891 p.p. [W †, BM]). — Lit: Zizka & al. (2013: with ills.). Distr: C to S-C Chile; stony arid to semi-humid habitats, 0–2200 m.  Pourretia alpestris Poeppig (1833). [1] Polycarpic, forming dense groups, 1.5–2.5 ( 5) m tall at flowering time, stems prostrate, covered with old leaves; L arched and recurving, sheath fleshy, 2.5–9.5  1.8–10 cm, margins in the upper part laxly serrate, L lamina 50–130  1.5–4.5 ( 5) cm, adaxially green with  cinereous scales, abaxially with dense coating of white scales, margins laxly serrate with uncinate Sp 4 mm long; Inf peduncle to 2.5 m, exceeding the leaves, floriferous part 0.25–1 m, pyramidal in outline, with 15–100 racemose branches, axes greenish, cinereous-tomentose; peduncular Bra oblong, reflexed,  deciduous; primary Bra like the upper peduncular Bra, shorter than the subtended branch; Br divergent, 25–30 cm, laxly to densely flowered in the lower , distally sterile with reduced bracts; floral Bra elliptic, 15 mm, shorter than the sepals; Ped slender, 10 mm; Sep oblong, acute, 20–25 mm; Pet blue-green, 50 mm;

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St included; Anth yellow-orange; Sty included; Sti yellow, surrounded by the anthers. P. alpestris ssp. alpestris — Lit: Zizka & al. (2013: with ills.). Distr: C-S Chile (Maule, Bío Bío, Araucanía), Argentina (San Juan)?; 0–2000 m. Incl. Puya whytei Hooker fil. (1868); incl. Puya pumila Ravenna (2000). [1] Plants 1.2–1.5 ( 2) m tall at flowering time; L lamina 1.5–2.5 ( 3.2) cm wide; Inf with 20 ( 40) laxly few-flowered branches. Gómez & Grau (2009) report the taxon from S Argentina (Neuquén), and Subils (2009: 346) from C Argentina (San Juan). The Neuquén record appears doubtful according to Zizka & al. (2013: 391), since the plants were described with violet petals. The plants from San Juan are described with white to lead-blue flowers. P. alpestris ssp. zoellneri Zizka & al. (Brittonia 65(4): 393, 2013). Type: Chile, Valparaíso (Zizka 8004 [FR]). — Distr: C to C-S Chile (Coquimbo, Valparaiso, Libertador O’Higgins, Maule: N of La Serena to E of Curicó); 0–2200 m. Incl. Pitcairnia alpestris L. H. Bailey (1916) (nom. inval., ICN Art. 38.1a). [1] Plants (2–) 2.5–5 m tall at flowering time; L lamina to 5 cm wide; Inf with (40–) 50–80 ( 100) densely many-flowered branches. P. asplundii L. B. Smith (Phytologia 6: 439, t. 2: figs. 14–15, 1959). Type: Ecuador, Imbabura (Asplund 20222 [S]). — Distr: N-C Ecuador (Pichincha, Cañar); shrub and Páramo vegetations, 3000–4000 m. I: Manzanares (2005: 364–365). [2] Polycarpic, forming dense groups, 2–3 m tall at flowering time; L sheath fleshy, orbicular, 6–7  5 cm, margins in the upper part serrate, L lamina 40–50  2–3 cm, adaxially glabrous, abaxially covered with cinereous scales, margins serrate with brown antrorse and retrorse Sp 5–7 mm long; Inf peduncle exceeding the leaves, 0.7–0.9 m, fertile part 0.5–0.7 m, laxly branched, cylindrical, covered with cinereous indumentum; peduncular Bra leaf-like, remote; primary Bra broadly ovate, 3–4 cm, margins in the upper part serrulate, with an attenuate apex, shorter than the subtended branch; Br divergent, strobilaceous or

J. M. Manzanares

ellipsoid, 5–9  1–4 cm, densely flowered, with a distinct stipe 1–4 cm long; floral Bra ovate, 25  15 mm, acute, reaching the middle of the sepals, margins entire; Ped stout, 5–7 mm; Sep obovate, 23  8 mm, rounded and  retuse; Pet green or bluish, 50 mm; St exposed; Anth green; Sty included. Gilmartin (1972: 30) treats this species as synonym of P. glomerifera based on an examination of the floral bracts (size and margins) and the margins of the primary bracts. Manzanares (2005: 364–365) discusses the differences between the two species in detail. Plants with a globose spike are often mis-identified as P. glomerifera, but careful examination of the flowers allows to easily distinguish between the species. — Locally used for fences as the groups grow very dense and are impossible to penetrate. P. berteroniana Mez pro sp. (in A. & C. de Candolle, Monogr. Phan. 9: 477, 1896). Type: Chile, Valparaiso (Bertero 115 [P, B, G]). — Lit: Zizka & al. (2013: with ills.). Distr: C Chile (Valparaíso: Quillota); dry and rocky areas near sea-level. I: Oliva-Esteve (2002: 230). [1 + 2] Polycarpic, forming dense groups, 5 m tall at flowering time, stems prostrate, simple or branched, covered with the bases of old leaves; L arched and recurving, sheath fleshy, 7  6 cm, margins entire, L lamina 100  2.5–5 cm, adaxially green and glabrous, abaxially with white appressed scales, margins laxly serrate with antrorse and retrorse yellow-green uncinate Sp 5–10 mm long; Inf peduncle exceeding the leaves, 0.6 m, densely white-lanate when young, floriferous part 0.6–1 m, densely branched with 80–100 branches, slenderly pyramidal in outline; peduncular Bra broadly lanceolate, reflexed,  deciduous; primary Bra like the upper peduncular Bra, much shorter than the subtended branch; Br 25–35 cm, densely flowered in the lower , sterile on the remainder with reduced bracts; floral Bra elliptic, acute, 30 mm, shorter than the sepals, white-lepidote; Ped slender, 15 mm; Sep narrowly oblong, 20–25  6.5 mm, broadly acute or obtuse, green; Pet elliptic, 50 mm, blue-green; St included; Fil blue-green; Anth orange; Sty

Puya BROMELIACEAE

included; Sti green, surrounded by the anthers; Fr ovoid, as long as the sepals. The correct identification of this name has been problematic for a long time. According to Zizka & al. (2013: 399–400), it should be considered to represent a natural hybrid, based on the observation of a plant intermediate between the locally sympatric P. alpestris ssp. zoellneri and P. venusta in the Pichidangui area (Zizka 8119, CONC, FR). Its inflorescence resembled that of P. alpestris but without the sterile terminal parts. Plants identified as P. berteroniana are also reported from the region of Coliguay (Varadarajan & al. 1490, SEL) from open cliffs in forest vegetation, sympatric with P. chilensis. More studies are needed to clarify the existence of natural hybridization, and the application of the name. According to the taxonomy of Zizka & al. (2013), material usually named P. berteroniana is attributable to P. alpestris ssp. zoellneri. P. boliviensis Baker (Handb. Bromel., 126, 1889). Type: Chile, Antofagasta (Gaudichaud s.n. [P, B, F, US]). — Lit: Jabaily & Sytsma (2010). Distr: N Chile (Antofagasta); stony coastal deserts and hills, 0–670 m. Incl. Puya copiapina Philippi (1895). [1] Polycarpic, forming dense groups, 2 m tall at flowering time; L sheath fleshy, L lamina 100  6 cm, succulent, both faces green and sparsely lepidote when young but soon completely glabrous, margins serrate with green uncinate stout Sp 10 mm long; Inf peduncle to 60 cm, exceeding the leaves, fertile part 50–70 cm, laxly branched, with 20–25 branches, broadly pyramidal in outline, green, tomentose-lepidote; lower peduncular Bra narrowly triangular and coarsely serrate, upper broadly lanceolate and entire, reflexed, primary Bra like the upper peduncular bracts, much shorter than the subtended branch; Br shortly stipitate, 20–30 cm, laxly flowered in the lower , sterile distally with reduced bracts; floral Bra lanceolate-elliptic, acute, 25–30 mm, shorter than the sepals, brown, tomentose-lepidote; Ped slender, 15 mm, tomentose-lepidote; Sep oblong-lanceolate, 30  6 mm, broadly acute, green, tomentose-lepidote; Pet elliptic, 50 mm, yellow or green-yellow, with a dark green spot at

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the base; St included; Anth dark; Sty included, longer than the stamens; Sti green; Fr subglobose. Despite the name, the taxon is endemic to Chile, but at the time when it was discovered, that part of Chile belonged to Bolivia. Flowering appears to be restricted to years with above-average precipitation associated with El Niño events. P. bravoi Aráoz & A. Grau (J. Bromeliad Soc. 58(5): 199–200, ills., 2008). Type: Argentina, Salta (Aráoz & al. 1596 [LIL, LPB, MO]). — Lit: Gómez & Grau (2009: with ill.). Distr: Argentina (Salta: near Santa Victoria); rockygrassy slopes and on rock walls, 3100 m; only known from the type locality. [2] Monocarpic, 1.5–2 m tall at flowering time; Ros solitary, stemless; L numerous, sheath fleshy, suborbicular, 9 cm, L lamina 60  6 cm, adaxially green, abaxially white-lepidote, margins entire except at the base and the top and there with few Sp 2–3 mm long; Inf peduncle as long as the leaves or shorter, 20–60 cm, stout, brown, floriferous part 100 cm, densely branched, cylindrical, brown, strobiliform, compact, all brown-lanate except for the petals; peduncular Bra densely imbricate making the peduncle appear nearly as thick as the inflorescence proper, apex reflexed; primary Bra ovate with triangular apex, longer than the subtended branch, lanate, brown, margins entire; Br shortly stipitate, spreading, densely 2to 3-flowered; floral Bra 5 mm, elliptic, acute, shorter than sepals and the pedicel, lanate, brown; Ped fleshy, 12 mm, lanate; Sep elliptic, acute, 35  8 mm, brown, lanate; Pet elliptic, blue, 60–70 mm; St included; Anth yellow; Sty included, longer than the stamens; Sti green; Fr ellipsoid, 15–35 mm; Se alate, 4–5  2.5–3.5 mm. An extraordinary monocarpic species, known from a single population with a few hundred individuals only. P. cardenasii L. B. Smith (Lilloa 14: 94, fig. 6, 1948). Type: Bolivia, Cochabamba (Cárdenas 4082 [US, GH]). — Distr: Bolivia (Cochabamba); rocky-grassy slopes in moderately moist open puna, (2950-?) 3800–3900 m. [2] Polycarpic, 1.5 m tall at flowering time; L sheath fleshy, L lamina 50  3 cm, arching,

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green, sparsely lepidote, margins laxly serrate with broad straight or curved brown Sp 8 mm long; Inf peduncle exceeding the leaves, to 75 cm, stout, lanate, floriferous part 25–50 cm, densely branched, cylindrical, pale lanate, glabrous at fruiting time; lower peduncular Bra leaf-like, the upper reflexed, base ovate and apex long attenuate, margins laxly serrate; primary Bra ovate, acuminate, 6  5 cm,  as long as the subtended branch, pale lanate, margins obscurely serrulate; Br shortly stipitate, densely 5- to 7-flowered; floral Bra ovate, acute or acuminate, 35–40  25–30 mm, exceeding the sepals; Ped slender, obconical, 10 mm; Sep broadly lanceolate, acute, 25  7 mm, thinly coriaceous; Pet blue, 45 mm; Fr subglobose. Insufficiently known and only collected twice recently. Petal colour was described as blue in the protologue, but as pale green in one of the recent collections (Solomon 9682, MO), made at a lower altitude (2950 m). P. castellanosii L. B. Smith (Phytologia 18: 140, 1969). Type: Argentina, Salta (Castellanos s.n. [BA 45819]). — Lit: Gómez & Grau (2009: with ill.). Distr: N Argentina (Salta: Valles Calchaquíes); dry mountain slopes with Monte vegetation, 2800–3000 m. I: Varadarajan (1989: 4); Grau & al. (2010: 12). – Fig. 1. Fig. 1 Puya castellanosii (Argentina, Salta, Valle de Lerma, beyond El Alisal, 2800 m). (Copyright: G. Rivera)

J. M. Manzanares

[1] Polycarpic, forming dense colonies of 3–8 rosettes aggregated into a massive clump, 1–2 m tall at flowering time; L sheath fleshy, reniform, 6  8 cm, L lamina 60–80  4.5 cm, broadly channelled, both faces densely lepidote, margins serrate with retrorse and antrorse curved Sp 5 mm long; Inf peduncle exceeding the leaves, 0.6 m, stout, green, lepidote, floriferous part 1–1.5 m, laxly branched, with 25–30 branches, pyramidal in outline, dark purplish, densely tomentoselepidote; lower peduncular Bra narrowly triangular, upper deltoid, reflexed, purplish; primary Bra ovate, with acute apex, much shorter than the subtended branch, lepidote, purplish; Br shortly stipitate, 25–30 cm, spreading, subdensely flowered in the lower , sterile distally with reduced bracts; floral Bra lanceolate, acute, 30 mm, shorter than the sepals, purplish, lepidote; Ped slender, 15 mm, tomentose-lepidote; Sep triangular-ovate, acute, 30  7 mm, purplish, lepidote; Pet elliptic, white-blue, 50–60 mm, St included; Anth yellow; Sty included; Sti green; Fr subglobose, shorter than the sepals. P. cerrateana L. B. Smith (Contr. US Nation. Herb. 29: 533, fig. 88, 1954). Type: Peru, Ancash (Cerrate 369 [US, USM]). — Distr: N & C Peru (Cajamarca, Ancash); rocky ground, 2000–3560 m.

Puya BROMELIACEAE

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[2] Polycarpic, forming dense groups united by thick stems, 1–1.5 m tall at flowering time; L sheath fleshy, elliptic, 9  6 cm, mostly glabrous, margins serrate in the upper part, L lamina 40–50  3–5 cm, adaxially glabrous, abaxially covered with fine white appressed scales, margins laxly serrate, Sp red-brown, uncinate, 7 mm; Inf peduncle exceeding the leaves, erect, 1–1.2 m,  lepidote, glabrous at anthesis, floriferous part 0.3–0.4 m, 7–10 cm ∅, simple, cylindrical, densely flowered, densely cinereous-lepidote; lower peduncular Bra leaf-like, upper quickly deciduous, lanceolate, attenuate, 5–9  1.5 cm, longer than the internodes, margins  denticulate esp. in the lower part; floral Bra 50–70  30 mm, ovate, acuminate, adaxially glabrous and nearly black, abaxially covered with a short-lanose dusty-white tomentum, longer than the sepals but with the upper coiling-reflexed at anthesis, margins entire; Fl tubular; Ped obconical, 10–15 mm, lanate; Sep elliptic, 35–40  12 mm, broadly acute, adaxially glabrous, abaxially lanate; Pet olive-green turning brown, 56 mm, apex recurved; St  exserted. P. chilensis Molina (Sag. Stor. Nat. Chili, 160, 351, 1782). Type [neo]: Chile, Bío Bío (Quezada 174 [CONC]). — Lit: Zizka & al. (2013: 396–398, ills. p. 392). Distr: C to C-S Chile (Coquimbo, Valparaiso, Libertador O’Higgins, Maule, Bío Bío); dry stony ground, 0–900 m. I: Rauh (1990: 128: fig. 131, 437); Duval (1990: 118: figs. 43–44); Oliva-Esteve (2000: 404). – Fig. 2.  Pitcairnia chilensis (Molina) Cat. Loddiges ex Loudon (1830); incl. Pourretia coarctata Ruiz & Pavón (1798)  Pitcairnia coarctata (Ruiz & Pavón) Persoon (1805)  Puya coarctata (Ruiz & Pavón) Fischer (1846); incl. Puya suberosa Molina (1810); incl. Puya gigantea Philippi (1864)  Puya chilensis var. gigantea (Philippi) Baker ex Mez (1896); incl. Puya quillotana W. Weber (1984). [1] Polycarpic, forming dense masses, 4–4.5 m tall at flowering time, stems prostrate, simple or branched, to 5 m; L sheath fleshy, broad, L lamina 80–100  5 cm, strongly channelled, adaxially soon glabrous, abaxially densely lepidote, margins serrate with antrorse and retrorse curved Sp

Fig. 2 Puya chilensis. (Copyright: M. Wisnev)

5–10 mm long with white scales; Inf peduncle exceeding the leaves, 1–1.5 m, stout, green, soon glabrous, floriferous part 1–1.5 m, densely branched, with 80–100 branches, cylindrical in outline, green, sparsely tomentose-lepidote; lower peduncular Bra leaf-like, upper broadly ovate, acute, reflexed, brown, lepidote; primary Bra triangular-ovate, acute, much shorter than the subtended branch, margins sparsely serrate, lepidote, brown-green; Br shortly stipitate, 30–40 cm, spreading, subdensely flowered in the lower , sterile distally with reduced bracts; floral Bra 50 mm, narrowly elliptic, acute or acuminate, as long as or shorter than the sepals, green, lepidote; Ped slender, 15 mm, lepidote; Sep ellipticoblong, broadly acute or obtuse, 35  13 mm, green, lepidote; Pet elliptic, yellow or yellowgreen, 50 mm; St included; Anth yellow; Sty included, longer than the stamens; Sti green. One of the best-known and most widespread species of Puya. When grazing too close to the plants, sheep sometimes get trapped by the heavily

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armed leaves with their mix of antrorse and retrorse spines. Quezada & al. (2014) studied photosynthesis over the whole range of the species. Populations from northern (more arid) sites had a predominance of CAM, those of the southern (wetter) sites had weakly expressed CAM. P. claudiae Ibisch & al. (Revista Soc. Boliv. Bot. 2(1): 131–132, ills., (incl. p. 127), 1998). Type: Bolivia, Santa Cruz (Ibisch & Ibisch 98040 [LPB]). — Distr: Bolivia (Santa Cruz); disturbed shrub areas, 2000 m. [2] Polycarpic, forming small clusters, 1.5 m tall at flowering time, stem simple or branched, to 0.5–1 m; L sheath fleshy, broad, L lamina 100–150  5 cm, green, adaxially glabrous, abaxially lepidote, margins serrate with antrorse curved Sp 15 mm long, black, white-lepidote at the base; Inf peduncle as long as or longer than the leaves, 0.4–0.5 m, stout, red-green, soon glabrous, floriferous part 1–1.5 m, densely branched, with 17–20 branches, pyramidal in outline, red, densely covered with white scales; lower peduncular Bra leaf-like, upper triangular, attenuate, lepidote, margins serrulate; primary Bra triangular, apiculate, 2.3–7  3–3.5 cm, much shorter than the subtended branch, glabrous, red, margins serrulate-denticulate; Br shortly stipitate for 5 cm, 20–40 cm, spreading, laxly flowered, red; floral Bra 10–15  8–10 mm, triangular, acute, longer than the pedicels, green-red, margins entire; Ped slender, 5–13 mm, white-tomentose; Sep elliptic, apiculate, 20–30  6–7 mm, red, green at the base, lepidote; Pet oblong, green, 50 mm; St included, 45 mm; Anth yellow; Sty included, 50 mm; Sti green; Fr 13 mm; Se broadly triangular, white-alate. After flowering, the very robust plants produce 2 new shoots at the base of the inflorescence, and thus form branched stems. The green leaves with the contrasting black marginal spines covered with white woolly scales at their base are very attractive. P. cochabambensis R. Vázquez & Ibisch (Bromelie 2000(1): 16–18, ills., 2000). Type: Bolivia, Cochabamba (Vásquez 3493 [LPB]). —

J. M. Manzanares

Distr: Bolivia (Cochabamba: Ayopaya; La Paz); on rocks, 3200–3600 m. [2] Polycarpic, forming dense colonies of several individuals, branched, 0.8 m tall at flowering time; L sheath fleshy, white, 2.5  2.5 cm, L lamina 18  1.5 cm, green, adaxially glabrous, abaxially completely covered by white scales, margins serrate with antrorse and retrorse Sp 2–3 mm long; Inf peduncle erect, to 50 cm, green, covered with russet stellate hairs, floriferous part 20–30 cm, simple, few-flowered,  secund, axis, bracts, pedicels and sepals covered with ochre-brown to russet stellate hairs, tomentum easily detached; lower peduncular Bra leaflike, upper lanceolate; floral Bra 23  8 mm, shorter than the pedicels, lanceolate, acute, papery, margins entire; Ped slender, 3–3.3 mm; Sep rounded-triangular, 28  10 mm, apex round; Pet linear, 70–75 mm, green; St included, 63 mm; Anth green-yellow; Sty 64 mm; Sti green; Fr and Se not described. Related to P. ferruginea, but differing by the unbranched inflorescence with half-erect to horizontally spreading flowers. P. coerulea Lindley (Bot. Reg. 26: t. 11 + text, 1840). Type: Chile (Lambert s.n. [[lecto — icono]: l.c. t. 11]). — Lit: Zizka & al. (2013). Distr: Chile (Coquimbo, Valparaiso, Libertador O’Higgins, Maule, Bío Bío); dry open rocky ground, 400–1200 m.  Pitcairnia coerulea (Lindley) Bentham (1883). [2] Polycarpic, forming dense groups, 1–2 m tall at flowering time; caudex erect or prostrate with time, branched; L sheath 2–8.5  1–7 cm, fleshy, suborbicular, dark castaneous, L lamina 11–62  0.4–3.5 cm, both faces with appressed cinereous scales, margins laxly serrate with red-brown antrorse uncinate Sp 3–5 mm long; Inf peduncle exceeding the leaves, 0.7–0.8 m, red, tomentulose-lepidote, floriferous part 1–1.2 m, laxly branched, cylindrical, red, white tomentose-lepidote when young; peduncular Bra leaf-like, lower lanceolate-ovate, with filiform or laminate apex, upper ovate with apiculate apex; primary Bra broadly ovate, with attenuate apex, 5–6 cm, much shorter than the subtended branch,

Puya BROMELIACEAE

margins in the upper part serrulate; Br spreading, laxly or sub-laxly flowered, 20–30  4–5 cm, with a long or short distinct stipe 5–20 cm long; floral Bra ovate, 5–44  1–15 mm, shorter than the pedicel, tip apiculate or acute, margins entire; Ped 10–20 mm; Sep lanceolate-triangular to oblong-ovate, 12–24  10 mm, rounded and apiculate; Pet dark blue, 50 mm, erect and forming a tube; St exposed; Anth yellow; Sty exserted, green; Fr ovoid or ellipsoid, shorter than the sepals. A beautiful and variable species that is divisible into 4 varieties: P. coerulea var. coerulea — Distr: C Chile (Coquimbo, Valparaiso, Libertador O’Higgins: Coastal cordillera); stony semi-arid areas, (20–) 500–2000 m. I: Zizka & al. (2013: 392). Incl. Pourretia coerulea Miers (1826) (nom. inval., ICN Art. 38.1a). [2] Plants 2–2.5 m tall at flowering time; L sheath fleshy, 4.5  5.3–13 cm, L lamina 36–62  1.5–3.5 cm; Inf white tomentose-lepidote; Fl  upwardly secund; floral Bra 10–44  4–15 mm, acuminate or apiculate, exceeding the pedicel and reaching the lower of the sepals; Ped 3–15  1.5–2.5 mm. P. coerulea var. intermedia (L. B. Smith & Looser) L. B. Smith & Looser (Phytologia 19: 287, 1970). Type: Chile, Libertador O’Higgins (Looser 2015 [GH]). — Distr: C Chile (Valparaiso, Libertador O’Higgins, Maule); stony hills, 0–1100 m.  Puya violacea var. intermedia L. B. Smith & Looser (1935). [2] Plants 2–2.3 m tall at flowering time; L sheath fleshy, 2.6–8.5  3.4–5.7 cm, L lamina 45–47  1–1.9 cm; Inf densely and persistently white-tomentose; Fl polystichously arranged; floral Bra 12–18  3–6 mm, exceeding the pedicel and shorter than the sepals; Ped 5–15  1.5–2 mm. P. coerulea var. monteroana (L. B. Smith & Looser) L. B. Smith & Looser (Phytologia 19: 287, 1970). Type: Chile, Libertador O’Higgins (Montero 26 [GH]). — Distr: C Chile (Libertador O’Higgins, Maule); 400–1200 m.

1081

 Puya violacea var. monteroana L. B. Smith & Looser (1935). [2] Plants 1–1.4 m tall at flowering time; L sheath fleshy, 2–4.7  2.3–3.3 cm, L lamina 19–48  0.8–1.1 cm; Inf glabrous at anthesis; Fl polystichously arranged; floral Bra 5–25  2–6 mm, exceeding the pedicel and shorter than the sepals; Ped 9–12  1.5–2 mm. This is the smallest variety. It is additionally characterized by having arching leaves. P. coerulea var. violacea (Brongniart) L. B. Smith & Looser (Phytologia 19: 287, 1970). Type: Ex cult. Hort. Paris (Anonymus s.n. [P]). — Distr: C Chile (Libertador O’Higgins, Maule, Bío Bío); stony areas, 100–1300 m.  Pitcairnia violacea Brongniart (1847)  Pourretia violacea (Brongniart) Linden (1853)  Puya violacea (Brongniart) Mez (1896); incl. Pourretia rubricaulis Miers (1826) (nom. inval., ICN Art. 38.1a)  Puya rubricaulis (Miers) Steudel (1841) (nom. inval., ICN Art. 38.1a); incl. Puya paniculata Philippi (1864); incl. Pitcairnia philippii Baker (1889); incl. Puya glabrata Philippi ex Baker (1889) (nom. inval., ICN Art. 36.1c). [2] Plants 1–2.7 m tall at flowering time; L sheath fleshy, 3–6.5  1.5–7 cm, L lamina 11–56  0.4–1.6 cm; Inf white-tomentulose, soon glabrous; Fl polystichously arranged; floral Bra 3–12  1–3 mm, shorter than the pedicel; Ped 10–24  1.5–2 mm. This variety is characterized by the long pedicels. P. densiflora Harms (Notizbl. Bot. Gart. Berlin-Dahlem 10: 791, 1929). Type: Peru, Cuzco (Herrera 1954 [B, F [photo]]). — Distr: SE Peru (Cuzco, Arequipa); on rocks and rocky slopes, under shrubs and cacti, 2000–3000 m. [1] Polycarpic, forming dense clusters, 1–1.5 m tall at flowering time; L arching, sheath fleshy, broadly ovate, 4–5 cm, densely serrulate, L lamina 40–60  3–4 cm, green, adaxially covered with a membrane of white scales, abaxially glabrous, margins laxly serrate, Sp slender, antrorse, red-brown, 10 mm; Inf peduncle  equalling the leaves, 0.4–0.6 m, stout, 15 mm ∅, densely and finely white stellate-tomentose, floriferous part 0.5–0.7 m, densely branched,

1082

subcylindrical-attenuate, finely white stellatetomentose including the flowers except the petals; lower peduncular Bra leaf-like, upper broadly ovate-lanceolate, long-caudate, imbricate, lepidote, margins serrate; primary Bra ovatelanceolate, 4–8  2.5–3 cm, attenuate, much shorter than the subtended branch, papery, light brown, margins spinose-serrate; Br shortly stipitate, 4–10 cm, suberect, subdensely 10flowered, green, terminal part of the main axis longer and attenuate; floral Bra 20–30  8 mm, from slightly longer than the pedicels to 3 as long, oblong-lanceolate, aristate, green, margins entire; Ped slender, 15 mm; Sep lanceolate, 25–30  5 mm, aristate, green, glabrous; Pet narrowly oblong, 40–50 mm, subobtuse, green-bluish; St exserted; Anth green, falcate, 40–50 mm; Sty exserted; Sti yellow. P. dyckioides (Baker) Mez (in A. & C. de Candolle, Monogr. Phan. 9: 486, 1896). Type: Bolivia, Chuquisaca (Weddell 3739 [P, GH [photo]]). — Lit: Gómez & Grau (2009: with ill.). Distr: S Bolivia (Chuquisaca, Tarija), NW Argentina (Jujuy, Salta, Tucumán, Catamarca); Yungas vegetation and grassy mountain slopes, between rocks, 1500–3500 m. I: Grau & al. (2010).  Pitcairnia dyckioides Baker (1889). [2] Polycarpic, forming small clusters, 0.8–1 m tall at flowering time, stem branched, to 40–50 cm; L sheath fleshy, broadly ovate, 3–5 cm, dark castaneous, L lamina 60–80  1.5–2 cm, green, adaxially glabrous, abaxially covered with pale appressed scales, margins serrate with antrorse uncinate Sp 3 mm long; Inf peduncle shorter than the leaves, 20–30 cm, stout, furfuraceous, floriferous part 40–80 cm, subdensely branched, cylindrical in outline, axes furfuraceous; lower peduncular Bra leaf-like, upper broadly ovate, acuminate or linear, imbricate, lepidote, margins serrulate; primary Bra ovate, acuminate, 4.5–6 cm, much shorter than the subtended branch, pink, margins coarsely serrulate; Br shortly stipitate, 10–20 cm, erect or divergent, subdensely few-flowered, pink; floral Bra 20  8 mm, longer than the pedicels, ovate-elliptic, apiculate, pink, margins entire or serrulate; Fl tubular; Ped slender, 5–7 mm; Sep ovate-elliptic, 12  6 mm,

J. M. Manzanares

obtuse, pink, glabrous; Pet oblong, 40 mm, green-bluish; St included, 35 mm; Anth orange; Sty included, 35 mm; Sti yellow. A very attractive species, esp. at flowering time, due to the pinkish inflorescences with contrasting dark blue-green flowers with orange anthers. After flowering, 1–2 offsets are formed. P. eryngioides André (Énum. Bromél., 5, 1888). Type: Ecuador, Loja? (André 4542 [K, K [photo]]). — Lit: Manzanares (2011: 233). Distr: Ecuador (Azuay, Loja); open slopes and shrubby Páramo vegetation, 2700–3100 m. I: Oliva-Esteve (2002: 231); Manzanares (2005: 309–310). [2] Polycarpic, forming small dense groups from a very short fleshy stem, 30–40 cm tall at flowering time; L sheath orbicular, 2  3 cm, chestnut-brown, margins entire, L lamina 15–20  1–1.5 cm, both faces glabrous, margins serrate with yellow-brown Sp 2–3 mm long; Inf peduncle exceeding the leaves, 20–25 cm, completely covered by the bracts, lepidote, red, floriferous part 7–10 cm, with 7–14 branches, globose or cylindrical in outline, dense, glabrous, reddish to violet; peduncular Bra ovate, with attenuate tip, red to violet, sparsely lepidote, margins serrate; primary Bra ovate to orbicular, acuminate, 1.5  0.9 cm, almost completely covering the subtended branch, glabrous, margins serrate; Br reduced or aborted, 2- to 3-flowered, fasciculate, 1  1.5 cm, glabrous; floral Bra 7  5 mm, ovate, cuspidate,  shorter than the sepals, red, covering the pedicel, margins serrulate; Ped only 1 mm; Sep ovate, 15  8 mm, acute, glabrous; Pet dark blue, 20 mm; St included; Anth yellow. The stems remain hidden by the predominant grass of the Páramos. P. eryngioides can be confused with P. exigua, but differs by forming groups with an above-ground stem and branched inflorescences. P. ferruginea (Ruiz & Pavón) L. B. Smith (Phytologia 16: 461, 1968). Type: Peru, Lima (Ruiz & Pavón s.n. [BM]). — Distr: Ecuador (Bolívar, Chimborazo, Morona Santiago, Loja, Zamora Chinchipe), Peru (Amazonas, Cajamarca, La Libertad, Huánuco, Pasco, Lima, Junín, Cusco, Apurimac, Puno), Bolivia (La Paz,

Puya BROMELIACEAE

Fig. 3 Puya ferruginea (Peru: Arequipa). (Copyright: I. Treviño)

Cochabamba); dry rocky slopes in the Andes and Ecuadorian Amazonia, 1800–3800 m. I: Rauh (1990: 435); Manzanares (2005: 319–320); Wisnev (2016: 172–174). – Fig. 3.  Pitcairnia ferruginea Ruiz & Pavón (1802)  Pourretia ferruginea (Ruiz & Pavón) Sprengel (1825)  Hepetis ferruginea (Ruiz & Pavón) Mez (1896); incl. Pitcairnia asterotricha Poeppig & Endlicher (1838); incl. Puya grandiflora Hooker (1861); incl. Pitcairnia consimilis Baker (1881)  Hepetis consimilis (Baker) Mez (1896); incl. Puya echinotricha André (1888)  Pitcairnia echinotricha (André) Baker (1889); incl. Pitcairnia weberbaueri Mez (1904); incl. Pitcairnia viridis Mez (1919); incl. Pitcairnia cotahuasiana Harms (1929); incl. Pitcairnia herrerae Harms (1929); incl. Pitcairnia imperialis Harms (1929); incl. Pitcairnia latibracteata Harms (1929); incl. Pitcairnia laresiana Harms (1930). [2] Monocarpic or polycarpic, 2–5 m tall at flowering time, solitary or forming dense groups, stem branched, procumbent or ascending; L sheath fleshy, broadly orbicular, 15  10 cm,

1083

white, glabrous, L lamina 80–100  5 cm, green, adaxially glabrous, abaxially grey-lepidote, margins serrate with antrorse Sp 15 mm long; Inf all parts and including pedicels and sepals with ferruginous indumentum, peduncle exceeding the leaves, 1 m, stout, floriferous part 1.5–3 m, laxly branched, pyramidal in outline; lower peduncular Bra leaf-like, upper shorter than the internodes, basal part ovate and with serrate margins, acuminate towards the tip and serrulate; primary Bra ovate, acuminate, 3–5  2 cm, shorter than the stipe, margins entire; Br 50–80 cm, with a distinct stipe 3–11 cm long, spreading, laxly 20- to 30-flowered; floral Bra 20  11 mm, ovate, acute, shorter than the pedicel, margins entire; Fl tubular, secundly arranged and pendulous after anthesis; Ped slender, 25–40 mm; Sep oblong, 45–50  5–8 mm, obtuse; Pet linear, 58–70 mm, initially violet at the start of anthesis changing to green to cream; St included, 50–60 mm; Sty included, 55–65 mm. The plants of the populations of Ecuadorian Amazonia are monocarpic, while those of the Andean populations of Ecuador, Peru and Bolivia from dry rocky slopes are polycarpic. According to Smith & Downs (1974), there is considerable variation between populations. Common to all forms is the dense ferruginous indumentum of all parts of the inflorescence and flowers except the petals. P. gilmartiniae G. S. Varadarajan & A. R. Flores (J. Bromeliad Soc. 40(4): 161–163, ills., 1990). Type: Chile, Coquimbo (Varadarajan & al. 1481 [GH, MO, WS]). — Lit: Zizka & al. (2013: 398). Distr: C Chile (Coquimbo: N of La Serena); coastal scrub, 50–520 m; critically endangered. [1] Polycarpic, forming dense masses, 1.5 m tall at flowering time, caulescent, stem prostrate, branched, 30–50 cm, covered with dry and persistent leaf sheaths; L sheath fleshy, 4–4.5 cm wide, ovate, dark castaneous, densely lepidote, L lamina 50–60  2.5 cm, arching, towards the base densely covered with cinereous white scales, both faces densely cinereous-lepidote (more densely along the margins), margins serrate with uncinate antrorse and retrorse Sp 10 mm long; Inf

1084

peduncle exceeding the leaves, 0.45 m, stout, green, persistently lanate esp. close to the bracts, fertile part 0.6–1 m, branched, with 20 branches and a long terminal part of the main axis, ellipsoid or ovoid in outline, green, sparsely tomentoselepidote; lower peduncular Bra leaf-like, upper broadly lanceolate, acuminate, reflexed, brown, papyraceous, persistently lanate, margins entire; primary Bra broadly ovate-lanceolate, acuminate, covering from nearly to > of the subtended branch, coriaceous, strongly veined, persistently lanate, brown, margin entire; Br 30–35 cm, stipitate for 2 cm, spreading, densely flowered in the lower , sterile distally with reduced bracts; floral Bra broadly ovate, 30 mm, acuminate, as long as the sepals, green in the lower part, brown above, sparsely lanate; Ped slender, 10 mm, tomentose-lepidote; Sep elliptic to lanceolate, 15–20  11 mm, obtuse, yellow-green, sparsely lanate at the apex; Pet yellow-green, 50 mm; St included; Fil yellow-green; Anth yellow; Sty included, as long as or  longer than the stamens; Sti green. Already in the protologue, a probable hybrid origin with P. chilensis and P. boliviensis as possible parents was discussed, and such an origin is supported in the molecular study of Jabaily & Sytsma (2010). The plants are very ornamental due to the dense greyish-white tomentum that covers the young leaves completely, and becomes restricted to the areas along the leaf margins. P. glomerifera Mez & Sodiro (Bull. Herb. Boissier, sér. 2, 4: 630, 1904). Type: Ecuador (Sodiro P-2 [B]). — Distr: N-C to S-C Ecuador (Pichincha, Tungurahua, Chimborazo, Azuay); very common in desert and semi-desert vegetation of the inter-Andean valleys, 2800–3900 m. I: Gilmartin (1972: Fig. 5); Oliva-Esteve (2002: 231); Manzanares (2005: 327). – Fig. 4. [2] Polycarpic, forming dense groups, 2–3 m tall at flowering time; L sheath fleshy, orbicular, 10  8 cm, margins entire, L lamina 50–80  2–3.5 cm, adaxially glabrous, abaxially densely lepidote, margins serrate with castaneous-brown antrorse and retrorse Sp 13 mm long; Inf peduncle exceeding the leaves, 0.7–1 m, floriferous part 0.5–0.8 m, dense, cylindrical, lanate with brown

J. M. Manzanares

Fig. 4 Puya glomerifera (Ecuador: Pichincha, around Quito). (Copyright: J. Manzanares)

indumentum; lower peduncular Bra leaf-like, upper remote, 5–9  2.5 cm; primary Bra ovate, 5–7  2 cm, with triangular apex,  shorter than the subtended branch, margins serrulate; Br globose, 3–5  4.5 cm, densely 6- to 9-flowered, stipe nearly none; floral Bra obovate, 30  25 mm, rounded or apiculate at the tip, shorter than the sepals, margins entire; Ped stout, 7 mm; Sep obovate, 25  15 mm, rounded and apiculate at the apex; Pet cream, 50 mm; St exposed; Anth yellow; Sty included. Forms dense groups and commonly used by native people as fence and to delineate borders. Flowering plants form a veritable “nectar oasis” with many visiting hummingbirds. P. grafii Rauh (Trop. subtrop. Pfl.-welt 52: 5–11, ills., 1985). Type: Venezuela, Amazonia (Graf s.n. in BG Heidelberg 64663 [HEID]). — Lit: Rauh (1985); Oliva-Esteve & Steyermark (1987); both with ills. Distr: Venezuela (Amazonia); grassy rocky slopes, 1200 m. I: Rauh (1990: 275, fig. 356).

Puya BROMELIACEAE

[2] Polycarpic, with long runners that form new plants, 80 cm tall at flowering time, stem base forming a thick pseudobulb of 6  5 cm; L sheath fleshy, conspicuous, covering the pseudobulb, white, 3.5–4  2 cm, densely lepidote in the upper , L lamina 20–30  0.5–1 cm, spreading to recurved, green, canaliculate, both faces densely silvery-white lepidote when young, upper face soon becoming glabrous, margins laxly serrate with antrorse uncinate Sp 2 mm long; Inf peduncle erect, 70 cm, greenish, with white-woolly indumentum, floriferous part 10–20 cm, simple, racemose, laxly cylindrical, with up to 15 flowers; lower Bra leaf-like, upper lax, much longer than the internodes, long triangular, reddish-green, lepidote; floral Bra 35  8 mm, lanceolate, acute, longer than the pedicel but shorter than the sepals, white-woolly lepidote, margins entire; Ped thick, 8–10 mm, densely lepidote; Sep long lanceolate, 35  8 mm, acute, green, densely grey-lepidote; Pet pale yellow with wine-red lines at the margin, 60  10 mm, outer face along the midrib with white stellate scales; St included; Anth white; Sti white. The plants form a pseudobulb at the base of the stem which is densely covered by the leaf sheaths. Due to the small size very suitable for cultivation in xerophyte gardens. P. gutteana W. Weber (Cat. Herb. Lips. Pl. Peruv. II: 42, ill., 1984). Type: Peru, Cusco (Gutte & Gutte 3308 [LZ]). — Distr: Peru (Cusco: Fundo Bellavista); hanging from rocks, 3200 m; known only from the fragmentary type. [2] Plants 70–80 cm tall at flowering time; L sheath fleshy, ovate, margins serrulate, L lamina 40–50  3 cm, narrowly triangular with a longacuminate apex, adaxially densely white-lepidote, abaxially glabrous, margins densely serrate with dark brown antrorse Sp 10 mm long; Inf peduncle shorter than or equalling the leaves, 30 cm, scatteredly white-tomentose, floriferous part 25  10 cm, dense, cylindrical, densely lepidote; lower peduncular Bra leaf-like, upper not imbricate but longer than the internodes, 8  2.5 cm, ovate-lanceolate, white-tomentose, margins densely serrate with Sp 2–3 mm long; primary Bra similar to the peduncular bracts, slightly longer than or as

1085

long as the subtended branch, margins serrate; Br suberect, 4–8 cm, to 4-flowered, subdensely tomentose-lepidote, stipe nearly none; floral Bra lanceolate-ovate, 23  8 mm, acute, much shorter than the sepals, margins minutely serrulate; Ped slender, 10 mm, tomentose-lepidote; Sep narrowly lanceolate, 35  8 mm, long-acuminate, the adaxial pair obtusely carinate with fleshy midvein area; Pet green-violet, 63 mm, lanceolate; St exposed; Anth yellow; Sty exposed. Related to P. densiflora, which is also native to the Cusco region, and similar in many details (indumentum, densely serrate leaf margins with antrorse spines 10 mm long, exposed stamens and style), but differing in the peduncle (longer than the leaves in P. densiflora, as long as or shorter than the leaves in P. gutteana). P. harmsii (A. Castellanos) A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 37: 497, 1933). Type: Argentina, Catamarca (Schreiter s.n. [LIL 34524]). — Distr: N Argentina (Catamarca, Tucumán); ravines of arid mountain slopes in the Monte - Prepuna ecotone, 2500 m. I: Gómez & Grau (2009: 200). – Fig. 5.  Puya spathacea var. harmsii A. Castellanos (1929). [1] Polycarpic, forming dense circular groups, 2 m tall or more at flowering time; L sheath fleshy, broadly ovate, 7 cm, brown and serrulate towards the apex, lower part white with entire margins, L lamina 100  3 cm, recurving, acuminate to pungent, both faces covered with a membrane of white appressed scales, margins laxly serrate, Sp antrorse, uncinate, red-brown, 4 mm; Inf peduncle equal or shorter than the leaves, 1.2 m, branched with at least 10 branches, lax, purple, appressedly pubescent with fine white stellate trichomes; lower peduncular Bra leaf-like, triangular with a long leaf-like lamina, 4–7  2.5 cm, margins serrate; primary Bra triangular, 6  5 cm, veined, apex acute, margins serrulate; Br subspreading, linear, 18 cm, laxly 10- to 18-flowered; floral Bra lanceolate, 14–20  7 mm, acute,  shorter than the pedicels, margins entire; Fl divergent to spreading; Ped curved, slender, 15–17 mm; Sep triangular, 18–20  10 mm, acute, carinate, sulcate when dry, thick;

1086

Fig. 5 Puya harmsii (Argentina, Tucumán, Tafi del Valle, S of Mollar, 2000 m). (Copyright: G. Rivera)

Pet black, appearing velvety with a pale blue tinge, 25 mm; St included, 18 mm; Anth greenolive, 5 mm; Sty included, 20 mm; Fr globose, 13  10 mm. P. harry-lutheri Gouda (J. Bromeliad Soc. 62 (6): 255, 2013). Type: Venezuela, Amazonas (Anonymus s.n. in BG Utrecht 2005GR01559 [L, VEN]). — Distr: Venezuela (Amazonas: Savana near Puerto Ayacucho near the Colombian border). [2] Polycarpic, acaulescent, 1.4 m tall at flowering time; Ros 30 cm tall; L spreadingarching, sheath fleshy, broadly obovate, pale brown and castaneous, 2–2.5  5 cm, whitish with closely appressed scales, L lamina 50–60  2–2.5 cm, channelled, both faces very densely lepidote to partly glabrescent with closely appressed, pruinose, white scales, margins laxly serrate with narrow, dark brown, antrorse Sp 1–3 mm long; Inf peduncle erect, exceeding the leaves, 45 cm,

J. M. Manzanares

greenish and brownish at the nodes, glabrous, floriferous part 65 cm, simple or with a rudimentary branch at the base, laxly cylindrical, 40- to 50-flowered, greenish to brownish, glabrous; lower peduncular Bra leaf-like, upper lax, attenuate, all exceeding the internodes but so narrow to expose part of the peduncle, brownish-red and green, margins serrulate; floral Bra 11.5  8 mm, ovate, obtuse, shorter than the pedicels and shorter than the sepals, diverging from the pedicel, green and partly black, glabrous; Ped thick, 18 mm, fleshy and cuneate towards the ovary; Sep ovatelanceolate, 40  11 mm, obtuse,  cucullate at the very apex, green tinged reddish, glabrous; Pet obovate-lanceolate, 68  14 mm, brown and adaxially green-veined to wholly green towards the base; St included but longer than the style, 60 mm; Anth cream-green; Sti cream. The original habitat remained unknown, and the plant was described from cultivated material at the Utrecht Botanical Garden, who obtained the plants from the Dutch Corn. Bak Nursery. The species has nocturnal flowers (otherwise in the genus only known from P. wrightii) that are probably pollinated by bats. P. hofstenii Mez (Repert. Spec. Nov. Regni Veg. 3: 8, 1906). Type: Argentina, Jujuy (Hofsten 1710 [B, F [photo]]). — Distr: S Bolivia (Potosí), NW Argentina (Jujuy); dry rocky slopes, 3400–3500 m. I: Gómez & Grau (2009: 200). Incl. Puya hauthalii Mez (1919). [1] Polycarpic, forming dense groups, 1 m tall or more at flowering time; L sheath fleshy, ovate, 6 cm, serrulate towards the apex, L lamina 50  5 cm, recurving, pungent, cinereous-lepidote esp. abaxially, margins laxly serrate, Sp antrorse, 5 mm; Inf peduncle shorter than the leaves, 0.7 m, branched with up to 11 branches, lax, red, glabrous; lower peduncular Bra leaf-like, the upper ones ovate, acute, 4–7 cm,  equalling the sterile branch base, margins entire; primary Bra ovate, 4  3.8 cm, acute, margins entire, red-pink; Br spreading to suberect, linear, 20 cm, laxly 12to 20-flowered; floral Bra ovate-elliptic, 15–18  12 mm, acute,  equalling the pedicels, margins entire; Fl suberect to deflexed; Ped slender, 15–18 mm; Sep lanceolate, 26  9 mm,

Puya BROMELIACEAE

rounded and apiculate, carinate at the base; Pet green-lavender, 30 mm; St included, 35 mm; Anth yellow, 4 mm; Sty exserted, 37 mm. Related to P. harmsii but distinguished by leaves green adaxially and densely lepidote abaxially (vs. adaxially and abaxially covered with a membrane of white appressed scales) and the pink glabrous inflorescence (vs. purple, appressed-pubescent with fine white stellate trichomes), as well as the green-lavender flowers with included stamens and styles (vs. velvetblack with a pale blue tinge and and exserted stamens and style). P. hoxeyi Janeba (Cact. Succ. J. (US) 89(4): 178–181, ills. (pp. 176–182), 2017). Type: Peru, Moquegua (Hoxey & Loayza Hoxey 1275.01 [HUSA]). — Distr: S Peru (Moquegua: N of Ilo); W- and NW-facing slopes of coastal loma hills, 700–900 m. [1] Polycarpic, to 4 m tall at flowering time, forming dense clumps to 2 m ∅; L numerous, sheath not described, lamina to 100 cm, to 6 cm wide at the base, spreading, narrowly triangular tapering into a sharp tip, glabrous, upper face green, slightly glossy, lower face green with white longitudinal lines, margin serrate, Sp brown-tipped from pale base, 5 mm long and retrorse near the leaf base, higher up antrorse, to 2 mm long near the leaf tip, 15–20 mm apart; Inf 1.5–2.5 m, floriferous in the upper to with 20–100 branches, finely white-tomentose; lower peduncular Bra leaflike but smaller, to 30  5 cm, upper peduncular Bra to 4  3 cm, softly papery, first green but rapidly drying and becoming brownish; Br 15–18 cm, spreading to slightly ascending, up to 30-flowered, terminal part sterile or floriferous; floral Bra 10  3 mm, acuminate; Fl tubular-funnelshaped, 45  15–20 mm; Ped 7 mm, tomentose; Sep 25  7 mm, greenish-yellow; Pet 40  15 mm, yellow-greenish, with faint longitudinal veins and outer face with greenish midvein area; St included but well visible in the throat; Fil 20 mm, greenish-cream; Anth 5 mm, greenish, pollen orange; Ov conical, 8  3–5 mm; Sty 25 mm; Sti lobes 2 mm, greenish-yellow, connivent.

1087

Compared with P. boliviensis and P. chilensis. The taxon is only known from 2 restricted populations. In both, 90% of all individuals are dead, and probably 20 rosettes, 14 cm tall at flowering time; Ros stemless, with a fleshy bulbous base, bulb 2.5  1.5 cm ∅; L sheath fleshy, conspicuous, broadly ovate, 1.5  2 cm, light brown on both faces, upper part lustrous dark brown esp. beneath, membranous, veined, margin near the transition to the lamina serrate, above sparsely brown-lepidote, L lamina narrowly triangular, acuminate, 6–8  0.2–0.3 cm, both faces lepidote, adaxially soon glabrous, margins with light brown retrorse Sp 1–2 mm long, 5 mm apart and almost opposite; Inf peduncle erect, 6–9 cm, brown, densely lepidote, floriferous part simple, shortly cylindrical, 5  2.5 cm,  densely spirostichously 5- to 15-flowered; peduncular Bra densely imbricate, erect, the basal ones subfoliate, with filamentous and serrate lamina, the middle and upper ones ovate-lanceolate, acuminate, 1.5–2 cm, margins finely serrate, veined, with a conspicuous midrib, abaxially reddish; floral Bra erect, ovate, acuminate, 20  10 mm, rose-red, membranous when dry, veined, sparsely lepidote, margins inconspicuously serrate; Ped almost absent and Fl subsessile; Sep narrowly triangular, acuminate, 11–12  2–3 mm, free to the base, with hyaline margin, rose-reddish, sparsely lepidote, margins apically inconspicuously serrate, carinate; Pet erect, tongue-shaped, 16–17 mm, red-violet, darker after anthesis; St included; Sty included; Fr 15 mm.

Puya BROMELIACEAE

Ideally suited for small gardens of succulents due to the small size, and outstanding because of the fleshy bulbous rosette base. P. raimondii Harms (Notizbl. Bot. Gart. Berlin-Dahlem 10: 213, 1928). Type: Peru, Ancash (Weberbauer 2955 [B, F [photo]]). — Lit: Salazar Castillo & al. (2012). Distr: Peru (La Libertad, Ancash, Lima, Junín, Huancavelica, Ayacucho, Apurimac, Cusco, Moquegua, Puno), Bolivia (La Paz, Cochabamba); rocky slopes and páramos, 2400–4200 ( 4460) m. I: Rauh (1990: 271–273, 397); Baensch (1994: 153); OlivaEsteve (2000: 409–410); Grau & al. (2010). – Fig. 9. Incl. Pourretia gigantea Raimondi (1874). [1] Monocarpic, 8.3–9.5 ( 15) m tall at flowering time; Ros solitary, caulescent, trunk 1–2 m, covered by old leaves; L very numerous, sheath fleshy, ample, 15–22  16 cm, L lamina 100–125  9 cm, stiff, adaxially green, abaxially white-lepidote, margins laxly serrate with antrorse and retrorse uncinate dark brown Sp 10–15 mm

Fig. 9 Puya raimondii (Peru, Arequipa, Cotahuasi, Lauripampa). (Copyright: I. Treviño)

1093

long; Inf peduncle exceeding the leaves, 0.6–0.9 m, very stout and 20–40 cm ∅, brown, floriferous part 4–5 ( 8) m, densely branched, cylindrical in outline, green, white-lanate; lower peduncular Bra leaf-like, upper ovate, attenuate towards the tip, reflexed, brown, lepidote; primary Bra broadly ovate, apiculate, much shorter than the subtended branch, subglabrescent, green with brown apex, margins entire; Br shortly stipitate, 20–30 cm, spreading or reflexed, densely flowered in the lower part, sterile distally with reduced bracts; floral Bra 50–60 mm, ovate-lanceolate, acuminate, longer than the sepals, green, subglabrescent; Ped stout, 15 mm, lepidote; Sep lanceolate, acute, 40 mm, green; Pet elliptic, white, 60–80 mm; St included; Anth yellow; Sty included, longer than the stamens; Sti green; Fr globose-ovoid. This massive plant is the queen of the genus, and by far the largest-growing taxon of the family. The massive inflorescence produces thousands of flowers whose abundant nectar attracts numerous birds, both specialized hummingbirds and more generalist passerine birds (Salinas & al. 2007, Hornung-Leoni & al. 2013). The life-span of a plant is estimated to be in the range 40–100 years, and 28 year old plants can reach a height of 3.36 m (Salazar Castillo & al. 2012). The total reproductive output is estimated to be 12 million seeds (Salazar Castillo & al. 2012). The species is a protected plant in Peru, and is commonly regarded as being threatened or in danger of extinction. Population-level reports (summarized by Salazar Castillo & al. (2012)) give a mixed image, with some populations showing a steep decline in plant numbers (attributed to various causes, including burning of plants to regenerate pastures and stop animals being trapped or hurt by the spiny leaves), while others appear to be stable or increase. P. retrorsa Gilmartin (Phytologia 57: 455, 1985). Type: Ecuador, Tungurahua (Gilmartin 1103 [US]). — Distr: Ecuador (Pichincha, Cotopaxi, Tungurahua, Chimborazo); shrubby Páramo vegetation, and as living fences in the inter-Andean valleys, in ravines, 2800–3400 m. I: Manzanares (2005: 347–348).

1094

[2] Polycarpic, forming dense groups, 2–3 m tall at flowering time, stem 30–40 cm ∅; L sheath fleshy, orbicular, 10  9 cm, margins entire, L lamina 40–60  2.5–3 cm, adaxially glabrous, abaxially densely lepidote, margins serrate with chestnut-brown Sp 6–8 mm long, retrorse at the base and antrorse in the upper part; Inf peduncle exceeding the leaves, 1–1.5 m, floriferous part 0.4–0.8 m, dense, cylindrical in outline, with 80–120 branches, tomentose with brown indumentum; lower peduncular Bra leaf-like, upper remote, 10–13  2.5 cm, lamina pendent; primary Bra orbicular, 4–8  5–6 cm, lower with attenuate apex, upper with acuminate apex, at anthesis shorter than the subtended branch, margins serrulate in the upper part; Br fasciculate, 3–9  2–4 cm, densely 9- to 15-flowered, stipe nearly none; floral Bra elliptic to obovate, 30–35  10–15 mm, with rounded and acuminate apex, exceeding the sepals, margins sparsely serrate; Ped stout, 3 mm; Sep elliptic, with rounded apex, 20  11 mm; Pet cream to green, 40 mm; St exposed; Anth yellow; Sty included. The plants form dense groups united by branched stems. Growth is moderately slow: A shoot branch of 50 cm takes 7 years to grow, and the thickly branched groups, with arms to 150 cm long (including branches) need 25 to 35 years to form (Manzanares 2005). P. robinfosteri G. S. Varadarajan & H. Luther (Selbyana 16(2): 235–238, ills., 1995). Type: Peru, Pasco (Foster 9062 [MO, SEL]). — Distr: Peru (Pasco: Cordillera Yanachaga); in seepage area, 2700–2800 m. [1] Polycarpic, solitary or mat-forming, 25–30 cm tall at flowering time; Ros stemless, forming a globose pseudobulb 2–3 cm ∅; L sheath fleshy, 1–1.2  0.8–1.2 cm, blackishbrown, inflated, L lamina very narrowly triangular to linear, attenuate, pungent, 6–10  0.75 cm, involute, subappressedly white-lepidote esp. abaxially, margins serrulate towards the base but entire towards the apex, Sp filiform, brown, antrorse, 6–15 mm; Inf peduncle erect, 15–18  0.5 cm, white-floccose, floriferous part simple, 4–7 cm, green, often subsecund, 5- to 10flowered; lower peduncular Bra subfoliaceous,

J. M. Manzanares

upper ones ovate, margins serrate, equalling to exceeding the internodes; floral Bra ovate, 10–17  9–14 mm, acute to attenuate, margins entire to inconspicuously and minutely serrulate, veined, lustrous, castaneous; Ped straight to curved-ascending, 6–13 mm, sparsely floccose; Sep oblong-lanceolate, 10–13 mm, broadly acute to retuse, veined, subcarinate, lustrous, castaneous; Pet 16–20 mm, green; St and Sty deeply included; Fr 8–10 mm. P. roezlii E. Morren (Belgique Hort. 35: 80, 1885). Type: Peru (‘Andes’) (Roezl s.n. [LG, GH [photo]]). — Distr: Peru (Lima); on rocks and sandy ground, 2200–2800 m. Incl. Pitcairnia megastachya Baker (1889); incl. Puya pectinata L. B. Smith (1932). [1] Polycarpic, forming groups, caulescent with short and stout ascending stem, 0.5–1.2 m tall at flowering time; L sheath fleshy, broadly ovate, merging with the lamina, dark, lustrous, abaxially  lepidote, L lamina narrowly triangular, attenuate to a pungent apex, recurving, 100  3 cm, adaxially glabrous, abaxially covered beneath with pale appressed scales, margins laxly serrate, Sp antrorse, uncinate, 8 mm; Inf peduncle exceeding the leaves, erect, stout, floriferous part laxly and amply 1 branched, 50 cm, sordidly pruinose-lepidote; lower peduncular Bra foliaceous with long lamina, dense, margins coarsely serrate; primary Bra broadly ovate, 3 cm, margins serrate with Sp 4 mm long, subcoriaceous, the lowest with narrow decurved lamina; Br 15 cm, scorpioid-decurved, flowers to the apex, lowest branches 15- to 25-flowered, subdense to lax at anthesis; floral Bra broadly ovate, acute, 10 mm, margins serrulate, equalling or shorter than the sepals, strongly convex; Ped very short and stout; Sep lanceolate, 16–18 mm, attenuate, mucronulate; Pet deep blue or purple, 28 mm; St included; Anth linear, 5 mm; Sty included; Fr subglobose, 10 mm. Strongly succulent and comparable to Ochagavia (Zizka & al. 2002). P. roldanii Betancur & Callejas (Caldasia 19: 78, ill., 1997). Type: Colombia, Antioquia (Callejas 11158 [HUA, COL, SEL]). — Distr:

Puya BROMELIACEAE

Colombia (Antioquia: Belmira, Río Chico); shrubby Páramo vegetation, 2400–3110 m. [2] Polycarpic, forming very dense cushions, 0.7–1 m tall at flowering time, stem short, prostrate, covered by old leaves; L sheath fleshy, oblong, 3–4  1.8–2.5 cm, margins serrate at the top, dark coffee-brown towards the top, with brown indumentum, L lamina 28–34  1–1.3 cm, triangular, with attenuate apex, adaxially glabrous and mottled with wine-red, abaxially lepidote with grey indumentum becoming denser towards the apex, margins with wine-red lines, serrate with dark antrorse and retrorse Sp 3–5 mm long; Inf peduncle exceeding the leaves, 30–50 cm; floriferous part 12–16 cm, simple, cylindrical in outline, at anthesis lax, 16- to 23-flowered, tomentose; lower peduncular Bra leaf-like, green-yellow, lepidote, upper ovate-elliptic, 5–7  2.4–3.1 cm, both faces lepidote with brown indumentum, margins minutely serrate; floral Bra 40–45  15–20 mm, ovate, acute, longer than the sepals, adaxially pubescent, abaxially tomentose, margins entire; Ped 8–16 mm, ascending, lepidote with brown indumentum; Sep oblong, truncate, 25–30  7–8 mm,  fleshy, adaxially glabrous, abaxially tomentose; Pet green-blue, spatulate, 60–68 mm,  fleshy; St included, 44–47 mm; Anth yellow; Sty as long as the stamens, included. Forming dense groups and easily recognizable by the leaves, which are all ascending-erect and mottled with wine-red. The young fleshy inflorescences are often eaten by the cattle of the local farmers. P. roseana L. B. Smith (Phytologia 7: 421, t. 1: fig. 13, 1961). Type: Ecuador, Loja (Rose 23210a [US]). — Lit: Gilmartin (1972). Distr: Ecuador (Loja); shrubby vegetation, in ravines, and as living fences, 2400–2600 m. I: Manzanares (2005: 361–362). [2] Polycarpic, forming dense groups, 1.5–2 m tall at flowering time, stem 30–50 cm, branched, ascending; L sheath fleshy, orbicular, 8  8.5 cm, margins entire, white, L lamina 50  2–2.5 cm, adaxially glabrous, abaxially with brown indumentum, margins serrate with chestnutbrown Sp 4 mm long; Inf peduncle exceeding the leaves, 0.6–0.8 cm, lepidote, floriferous part

1095

0.5–1.5 m, dense, cylindrical in outline, with 20–25 branches, with ferruginous indumentum; lower peduncular Bra leaf-like, upper triangular, remote, 5–15  1.5 cm, apex attenuate and pungent, margins serrate; primary Bra 2.5  1.5 cm, with an orbicular sheath, margins  serrate, apex long attenuate, shorter than the subtended branch during anthesis; Br 19–30  3.7 cm, at an angle of 90 with the axis, subdensely 20- to 25-flowered, with a distinct sterile base 3 cm long; floral Bra ovate, 10  11 mm, with apiculate apex, as long as or longer than the pedicel but shorter than the sepals, adaxially glabrous, abaxially with ferruginous indumentum, margins entire; Ped stout, 7–9 mm; Sep oblong, obtuse, 20–22  12 mm, fleshy, adaxially glabrous, abaxially with ferruginous indumentum; Pet bluish, 30 mm; St exposed; Anth yellow; Sty included; Fr 20 mm. The inflorescence architecture is very variable and depends on the plant’s vegetative size. Inflorescences vary between 0.5 and 1.5 m long, with branches 19–30 cm long. Short branches have densely polystichously arranged flowers, long branches have subdensely or occasionally laxly arranged flowers. P. sehuencasensis R. Vásquez & al. (Bromelie 2007(3): 150–155, ills., 2007). Type: Bolivia, Cochabamba (Vásquez & Lara 5279 [LPB, BOLV, MO, SEL]). — Distr: Bolivia (Cochabamba); slopes in cloud forest, 2050–2100 m. [2] Polycarpic, forming dense groups, branched, 2 m tall at flowering time; L sheath orbicular, 5–6  8–9 cm, white towards the base, dark chestnutbrown at the apex, L lamina 50–70  5–8 cm, recurving, adaxially glabrous, abaxially densely cinereous-lepidote, margins serrate with dark brown antrorse flat Sp 8 mm long; Inf peduncle exceeding the leaves, 0.6–0.8 m, lepidote with white stellate scales but then glabrous, floriferous part 1–1.2 m, lax, pyramidal in outline, paniculate with 8 branches; lower peduncular Bra leaflike, upper oblong-elliptic, longer than the internodes, 5–20 cm, margins entire near the base, densely serrate towards the apex; primary Bra triangular, spreading, 10 cm, as long as the sterile base of the subtended branch, green, margins densely serrate with antrorse and retrorse spines

1096

near the pungent apex; Br 38–45 cm, pendent, subdense, densely covered with white stellate scales, stipe distinct, 10 cm; floral Bra triangular, 50  20 mm, attenuate, longer than the pedicel, densely white-stellate, margins serrate towards the apex with antrorse and retrorse spines; Ped stout, 12 mm, densely white-stellate; Sep triangular, attenuate, 40  8 mm, densely covered by the white-stellate indumentum; Pet blue and green at the base, 56 mm, apex recurved; St exserted, 70 mm, blue; Anth yellow; Sty as long as the stamens; Fr 20 mm. P. sodiroana Mez (Bull. Herb. Boissier, sér. 2, 4: 630, 1904). Type: Ecuador, Pichincha? (Sodiro P-5 [B]). — Lit: Distr: N Ecuador (Pichincha, Azuay); arid inter-Andean areas, ravines, and also as living fences, 2800–2900 m. I: Manzanares (2005: 357). Incl. Puya gummifera Mez & Sodiro (1904). [2] Polycarpic, forming dense groups, 2–2.5 m tall at flowering time, stem 25 cm ∅, branched; L sheath fleshy, ovate, 4.5  5 cm, white above and chestnut-brown below, with entire margins, L lamina 40–60  2.5–3.5 cm, adaxially glabrous, abaxially densely lepidote, margins serrate with brown Sp 6–10 mm long, retrorse in the lower part and antrorse in the upper part; Inf peduncle exceeding the leaves, 1 m, lepidote, not completely covered by the bracts, floriferous part 0.7–0.9 m, lax or sublax, cylindrical in outline, with 25–30 branches, covered with cinereous indumentum; lower peduncular Bra leaf-like, upper triangular, pendent, 5–6  2.5 cm, apex attenuate, margins serrate; primary Bra ovate to triangular, 5–6  2–2.5 cm, as long as or exceeding the stipe and shorter than the subtended branch, margins  serrate; Br ellipsoid in outline, 10–18  3.5–4 cm, at an angle of 90 with the axis, densely 15- to 25-flowered, with a distinct stipe 2–4 cm long; lower floral Bra orbicular, upper ovate, 15–20  13 mm, acute or acuminate, reaching the middle of the sepals and longer than the pedicel, adaxially glabrous, abaxially with cinereous-lanate indumentum, margins entire; Ped stout, 9 mm, lanate; Sep tongue-shaped, with rounded and retuse apex, 24  9 mm, adaxially glabrous, abaxially lanate; Pet green,

J. M. Manzanares

cream or violet, 40 mm; St exposed; Anth yellow; Sty included. Sometimes growing sympatrically with P. aequatorialis, with which it easily hybridizes (see P. pichinchae). The flower colour varies from green to cream and violet, depending on the population, probably indicating some degree of introgression from P. aequatorialis. The flowers are intensely visited by several species of hummingbirds, including the Giant Hummingbird of the Andes, Patagona gigas. P. tillii Manzanares (Jewels Jungle Bromeliac. Ecuador II, Pitcairnioideae, 344–345, ills., 2005). Type: Ecuador, Pichincha (Manzanares 5738 [QCNE, MO, WU]). — Distr: Ecuador (Pichincha: W Cordillera); cloud forest, humid shrubby mountain sides, 2300–3400 m. [2] Polycarpic, forming dense groups, 2–2.5 m tall at flowering time, stem 40 cm ∅, branched; L sheath fleshy, orbicular, 6  6 cm, glabrous, white above and chestnut-brown below, margins entire, L lamina 50–60  3.5 cm, adaxially glabrous, abaxially densely lepidote, margins serrate with brown antrorse Sp 10 mm long; Inf peduncle exceeding the leaves, 1.2 m, lepidote, not completely covered by the bracts, floriferous part 0.7 m, dense, cylindrical in outline, with 25–30 branches, brown or green-brown, densely lepidote; lower peduncular Bra leaf-like, the upper with an elliptic sheath, lamina triangular, pendent, 9–15  2.5 cm, brown, margins serrate; primary Bra with ovate sheath and triangular lamina, pendent or spreading, 10  6 cm, as long as or longer than the subtended branch but not hiding it, margins  serrate; Br globose or ellipsoid in outline, 5  2 cm, at an angle of 90 with the main axis, densely 6- to 9-flowered, stipe nearly none, green, tomentose; floral Bra ovate, 35  18 mm, with acute or attenuate apex, hiding the sepals, adaxially glabrous, abaxially brown-tomentose, margins entire; Ped 4 mm; Sep elliptic, obtuse, 23  13 mm, adaxially glabrous, abaxially tomentose; Pet green, cream or pink, 50 mm; St exposed; Anth yellow; Sty included. One of the few Ecuadorian species from the cloud forest zone. Its rosettes are usually covered by small shrubs or lichens. The majority of the

Puya BROMELIACEAE

plants has pink flowers, which are pollinated by a variety of hummingbird species. P. tuberosa Mez (in A. & C. de Candolle, Monogr. Phan. 9: 483, 1896). Type: Bolivia (‘Peru’) (Haenke s.n. [M]). — Lit: Vásquez & Ibisch (2007: with ills.). Distr: Bolivia (Tarija, Chuquisaca, Santa Cruz); grassy-rocky slopes, 1000–1800 m. I: Rauh (1990: 276: Fig. 359, as P. serranoensis vars.). Incl. Puya hromadnikii Rauh (1983); incl. Puya serranoensis Rauh (1983); incl. Puya vallograndensis Rauh (1983); incl. Puya serranoensis var. brevispica Rauh (1985); incl. Puya vallograndensis var. simplex Rauh (1985); incl. Puya pachyphylla R. Vásquez & Ibisch (2007). [2] Polycarpic, forming dense groups, 30–50 cm tall at flowering time (incl. the very short erect rhizome); L sheath suborbicular, fleshy, forming a globose bulb 3 cm ∅, blackish-brown, L lamina 15–18  0.2–0.35 ( 1.2) cm, linear, flexuose, green, adaxially glabrous, abaxially covered with pale narrow spreading scales, margins laxly serrate with antrorse and retrorse Sp 1–2 mm long; Inf peduncle erect, 10–15 cm, as long as or shorter than the leaves, white-flocculose, floriferous part 20–30 cm, simple or with a few short branches at the base, cylindrical, laxly 12- to 20-flowered, white-flocculose; lower peduncular Bra leaf-like, ovate with filiform lamina, palelepidote; primary Bra ovate, acuminate, 2 cm, shorter than the subtended branch, minutely serrulate; Br short, 2- to 5-flowered; floral Bra 4–7 mm, ovate, acute, red, shorter than the sepals, longer or shorter than the pedicel; Ped 4–7 mm, erect, whiteflocculose; Fl secund, tubular; Sep lanceolateoblong, acute, 7–9  6 mm, white-flocculose; Pet blue-violet, erect, 30 mm; St included; Sty included. The leaves superficially resemble a bunch of grass. The plants have a basal erect rhizome that stores water and carbohydrates, to allow to survive prolonged periods without rain. P. tuberosa is a widespread and variable species, and several local forms that were described as independent taxa were recently synonymized (Vásquez & Ibisch 2007). Populations spanning a continuum of small plants with filiform leaves to larger plants with triangular leaves are known.

1097

P. venusta Philippi (Anales Univ. Chile 91: 613, 1895). Type [lecto]: Chile, Región IV, Coquimbo (Philippi 940 [B]). — Lit: Zizka & al. (2013: with ill.). Distr: Chile (Coquimbo, Valparaíso); coastal rocks and sand, 5–250 ( 750) m. I: Oliva-Esteve (2002: 238); Wisnev (2016: 160).  Pitcairnia venusta (Philippi) Baker (1889); incl. Pitcairnia sphaerocephala Baker (1889); incl. Puya coquimbensis Mez (1896); incl. Puya gaudichaudii Mez (1896). [2] Polycarpic, forming dense compact groups, 1.2–1.7 m tall at flowering time, stem to >40 cm; L sheath fleshy, suborbicular, L lamina 40–50  2.5–3 cm, thickly succulent, both faces appressedlepidote and canescent, margins serrate with green antrorse and retrorse uncinate Sp 5–7 mm long; Inf peduncle far exceeding the leaves, 0.7–1 m, floriferous part 0.5–0.7 m, dense, cylindrical, simple and strobilate or compound with several spicate branches, white-tomentose; lower peduncular Bra leaf-like, upper triangular-ovate, 5–10  2.5–3 cm, acuminate, deep red-violet, margins laciniate-serrate; primary Bra ovate, 5–7  3 cm, acuminate, shorter than the subtended branch; Br spicate, 5–8 cm, dense, with a stipe 4–6 cm long, end part of the inflorescence cylindrical, 12 cm; floral Bra elliptic, 30–35  20–25 mm, with a rounded and acuminate apex, longer than the sepals, densely imbricate, tomentose, margins entire or at the base sparsely serrate; Ped stout, 7 mm, tomentose; Sep oblong, obtuse, 15–20  10 mm; Pet violet, 35–40 mm; St exposed; Anth yellow; Sty included, green. P. vestita André (Énum. Bromél., 5, 1888). Type: Ecuador, Pichincha (André 3739 [K]). — Lit: Gilmartin (1972). Distr: Ecuador (Pichincha, Cotopaxi, Bolívar, Chimborazo); among shrubs and in Páramo vegetation, 2800–3900 m. I: Manzanares (2005: 311–312). [2] Polycarpic, forming dense clusters, 1.5–2 m tall at flowering time, stem 25 cm ∅; L sheath fleshy, orbicular, 10  8 cm, glabrous, chestnutbrown, margins entire, L lamina 35  2.4 cm, adaxially glabrous, abaxially densely lepidote, margins serrate with bright brown antrorse Sp 6 mm long; Inf peduncle exceeding the leaves, 0.6–0.8 m, floriferous part 0.2–0.4 m, dense,

1098

J. M. Manzanares

cylindrical or ellipsoid in outline, with 15–20 branches, with ferruginous indumentum; lower peduncular Bra leaf-like, with orbicular sheath and triangular lamina, 7–15  1.2 cm, lamina pendent, densely lepidote, margins serrate; primary Bra ovate, 7–10  4 cm, attenuate or long acuminate, margins  serrulate, lower bracts exceeding the subtended branch, upper equalling or  shorter than the subtended branch, adaxially sparsely lepidote, abaxially with ferruginous indumentum; Br reduced, 3–5  1.4 cm, densely 3- to 5-flowered, stipe nearly none; floral Bra ovate, 30–35  21 mm, with acute navicular apex, hiding the sepals, margins entire; Ped nearly none, 1 mm; Sep ovate, obtuse, 20–25  11 mm, adaxially glabrous, abaxially with ferruginous indumentum; Pet greenish, 45 mm; St exposed; Anth yellow; Sty included. Related to P. glomerifera and P. retrorsa, but most clearly differing from both by the antrorse leaf margin spines (both antrorse and retrorse in P. glomerifera, antrorse in the distal part and retrorse in the basal part in P. retrorsa).

Incl. Puya flora Spegazzini (1896). [2] Monocarpic and Ros solitary, 50 cm tall at flowering time; L sheath fleshy, ovate, 6  4 cm, L lamina 20  1–2 cm, adaxially glabrous, abaxially appressedly cinereous-lepidote, margins laxly serrate with uncinate mostly retrorse Sp 4 mm long; Inf peduncle exceeding the leaves, 20–30 cm, stout, floriferous part 20–30 cm, simple, dense, ellipsoid in outline, strobilate, compact, pink; peduncular Bra densely imbricate making the peduncle appear nearly as thick as the floriferous part, elliptic, pink, apex reflexed, margins serrate; floral Bra 45  15 mm, ellipticoblong, longer than the pedicel and sepals, exceeding the flowers but reflexed, pink, margins of the lower ones  serrulate; Ped obconical, stout, 8 mm; Sep elliptic, acute, 35 mm, lanate at the base; Pet blue or violet, oblanceolate, 40 mm; St included; Anth 7 mm, yellow; Fr subpyramidal, 25–35 mm; Se irregularly triangular, 3.5 mm. Related to P. bravoi, from which it differs by having spiny leaf margins and a simple inflorescence.

P. weberiana E. Morren ex Mez (in A. & C. de Candolle, Monogr. Phan. 9: 492, 1896). Type: Argentina, Salta (Weber s.n. in BG Liège s.n. [G, GH [photo]]). — Lit: Gómez & Grau (2009: with ill.). Distr: Argentina (Salta, Tucumán); among rocks, 3600–4000 m.

P. weddelliana (Baker) Mez (in A. & C. de Candolle, Monogr. Phan. 9: 475, 1896). Type: Bolivia, Tarija (Weddell 4001 [P, GH [photo]]). — Distr: Bolivia (Chuquisaca, Tarija); shrubby rocky slopes, 1800–3000 m. – Fig. 10.  Pitcairnia weddelliana Baker (1889).

Fig. 10 Puya weddelliana (Bolivia, Chuquisaca, Nor Cinti, Camargo). (Copyright: S. Janke)

Puya BROMELIACEAE

[1] Polycarpic, forming dense groups, 4–5 m tall at flowering time, stem prostrate, woody, simple or branched, to 1–1.5 m; L sheath fleshy, broad, L lamina 90–120  4–5 cm, linear-triangular, adaxially glabrous, abaxially densely greylepidote, margins laxly serrate with retrorse curved Sp 5 mm long, dark brown; Inf peduncle exceeding the leaves, 1–1.5 m, stout, green, soon glabrous, fertile part 1.5–2.5 m, densely branched, with 60–90 branches, cylindrical in outline, bluish, sparsely tomentose-lepidote; lower peduncular Bra leaf-like, upper triangular, attenuate towards the tip, reflexed, lepidote; primary Bra triangular-ovate, attenuate towards the tip, much shorter than the subtended branch, lepidote, bluish; Br shortly stipitate, 30–40 cm, spreading, densely flowered in the lower , sterile distally with reduced bracts; floral Bra 20–30 mm, lanceolate, acute, shorter than the sepals, bluish, lepidote, margins entire; Ped slender, 10 mm, green,  tomentose; Sep oblong, broadly obtuse, 18  10 mm, lower part green, bluish above, first whitetomentose, becoming glabrous; Pet oblong, obtuse, bright blue, 40–50 mm; St included; Anth yellow; Sty included, longer than the stamens; Sti yellow. Undoubtedly the most spectacular species of Puya when flowering. The inflorescence of 1–1.5 m is filled with bright blue flowers, overtopped by the sterile cinereous top of the branches. P. yakespala A. Castellanos (in Descole, Gen. Sp. Pl. Argent. 3: 213, t. 55, 124d-f, 1945). Type: Argentina, Salta (Reynaga s.n. [LIL 34523]). — Lit: Gómez & Grau (2009: with ill.). Distr: Argentina (N Salta: Santa Victoria); grassy rocky slopes and ravines, 3400–4000 m. I: Grau & al. (2010). [2] Polycarpic, forming dense groups, 3–4 m tall at flowering time, stem elongate; L sheath orbicular, fleshy, L lamina 40–70  2–3.5 cm, recurved, green, adaxially glabrous, abaxially white-lepidote, margins serrate with Sp 5–6 mm long; Inf peduncle  upcurved at the base, 1–1.8 m, exceeding the leaves, lanate, floriferous part 0.5–0.9  0.2–0.3 m, simple, ellipsoid in outline, strobiliform, dense, brown-lanate becoming glabrous; lower peduncular Bra leaf-like, upper triangular, reflexed, with acuminate apex,

1099

7 cm, densely imbricate making the peduncle appear nearly as thick as the flowering part; floral Bra 10–14 mm, longer than pedicel and sepals, elliptic, acuminate, soft and often reflexed, brown, concave, membranous, the lower part lanate, margins entire; Ped 2–2.5 mm, subcylindrical; Sep triangular, acuminate, 45  12 mm, lanate; Pet yellow, oblong, revolute, 60  25 mm; St included; Anth blackish; Sty included, green; Fr ellipsoid, 30–35  15–20 mm; Se alate, 3–4  3 mm. A very decorative species when in flower, with a light brown inflorescence with outstanding yellow flowers. The peduncle is bent at the base and has bracts densely arranged as to give a rugged look.

References Baensch, U. (1994) Blühende Bromelien. Flowering Bromeliads. Nassau (BS): Tropic Beauty Publishers. Duval, L. (1990) The Bromeliads. Pacifica (US): Big Bridge Press. Gilmartin, A. J. (1972) The Bromeliaceae of Ecuador. Lehre (DE): Verlag von J. Cramer. Givnish, T. J. [& al. 2007], Millan, K. C., Berry, P. E. & Sytsma, K. J. (2007) Phylogeny, adaptive radiation, and historical biogeography of Bromeliaceae inferred from ndhF sequence data. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution: Poales. Aliso 23: 3–26. https://doi.org/10.5642/ aliso.20072301.04 Givnish, T. J. [& al. 2011], Barfuss, M. H. J., Ee, B. van, Riina, R., Schulte, K., Horres, R., Gonsiska, P. A., Jabaily, R. S., Crayn, D. M., Smith, J. A. C., Winter, K., Brown, G. K., Evans, T. M., Holst, B. K., Luther, H., Till, W., Zizka, G., Berry, P. E. & Sytsma, K. J. (2011) Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: Insights from an eight-locus plastid phylogeny. Amer. J. Bot. 98(5): 872–895, ills., maps. https://doi.org/10.3732/ajb.1000059. Gómez, S. & Grau, A. (2009) Las especies de Puya (Bromeliaceae) en la Argentina. Bol. Soc. Argent. Bot. 44(1–2): 175–208, ills., key. http://www.biodiver sitylibrary.org/part/112523# Gouda, E. J. [& al. 2012+], Butcher, D. & Gouda, C. S. (2012+) Encyclopaedia of Bromeliads, Version 3.1 (2012). Utrecht (NL): University Botanic Gardens, Utrecht. http://bromeliad.nl/encyclopedia/ Grau, A. [& al. 2010], Gómez Romero, S. E. & Aráoz, E. (2010) Puyas andinas. Ci. Hoy 20(118): 8–15, ills. http://www.cienciahoy.org.ar/ch/ln/hoy118/puyas.pdf Hornung-Leoni, C. T. [& al. 2013], González-Gómez, P. L. & Troncoso, A. J. (2013) Morphology, nectar characteristics and avian pollinators in five Andean Puya

1100 species (Bromeliaceae). Acta Oecol. 51: 54–61, ills. https://doi.org/10.1016/j.actao.2013.05.010. Jabaily, R. S. & Sytsma, K. J. (2010) Phylogenetics of Puya (Bromeliaceae): Placement, major lineages, and evolution of Chilean species. Amer. J. Bot. 97(2): 337–356, ills., map. https://doi.org/10.3732/ajb.0900107. Jabaily, R. S. & Sytsma, K. J. (2013) Historical biogeography and life-history evolution of Andean Puya (Bromeliaceae). Bot. J. Linn. Soc. 171(1): 201–224, ills. https://doi.org/10.1111/j.1095-8339.2012.01307.x. Manzanares, J. M. (2005) Jewels of the jungle. Bromeliaceae of Ecuador. Part II. Pitcairnioideae. Quito (EC): Published by the author. Manzanares, J. M. (2011) Bromeliaceae. In: León-Yánez, S. & al. (eds.): Libro rojo de las plantas endémicas del Ecuador. Ed. 2; pp. 214–245. Quito (EC): Herbario QCA, Pontifica Universidad Católica del Ecuador. Miller, G. A. (1994) Functional significance of inflorescence pubescence in tropical alpine species of Puya. In Rundell, P. W. & al. (eds.), Tropical alpine environments: Plant form and function; pp. 195–213. Cambridge (GB): Cambridge University Press. Oliva-Esteve, F. (2000) Bromeliads. Caracas (VE): Armitano Editores. Oliva-Esteve, F. (2002) Bromeliaceae III. Caracas (VE): Producciones Oliva Esteve. Oliva-Esteve, F. & Steyermark, J. (1987) Bromeliaceaes [sic!] of Venezuela. Caracas (VE): Armitano Editores. Quezada, I. M. [& al. 2014], Zotz, G. & Gianoli, E. (2014) Latitudinal variation in the degree of Crassulacean Acid Metabolism in Puya chilensis. Pl. Biol. (Stuttgart) 16: 848–852, map, ill., diags. https://doi.org/10.1111/ plb.12181. Rauh, W. (1966) The Bromeliads of the Peruvian Andes. Bromeliad Soc. Bull. 16(2): 28–39, ills. http://journal. bsi.org/V16/2/ Rauh, W. (1985) Puya grafii, a new dwarf Puya from southern Venezuela. J. Bromeliad Soc. 35(1): 21–23, ills. Rauh, W. (1990) Bromelien. Ed. 3. Stuttgart (DE): Verlag Eugen Ulmer. Salazar Castillo, J. [& al. 2012], Caceres de Baldarrago, F. & Poma, I. (2012) Puya raimondii. La regina delle Ande. The Queen of the Andes. Cactus & Co. 15(4): 28–51, ills.

J. M. Manzanares Salinas, L. [& al. 2007], Arana, C. & Suni, M. (2007) El néctar de especies de Puya como recurso para picaflores Altoandinos de Ancash, Perú. Revista Peruana Biol. 14(1): 129–134, map, ills. Schulte, K. & Zizka, G. (2008) Multi locus plastid phylogeny of Bromelioideae (Bromeliaceae) and the taxonomic utility of petal appendages and pollen characters. Candollea 63(2): 209–225. http://www.ville-ge.ch/cjb/publications_ candollea_632.php Schulte, K. [& al. 2009], Barfuss, M. H. & Zizka, G. (2009) Phylogeny of Bromelioideae (Bromeliaceae) inferred from nuclear and plastid DNA loci reveals the evolution of the tank habit within the subfamily. Molec. Phylogen. Evol. 51(2): 327–339. https://doi.org/10.1016/j.ympev. 2009.02.003 Smith, L. B. & Downs, R. J. (1974) Flora Neotropica. Monograph No. 14 [Bromeliaceae]. Part 1: Pitcairnioideae. New York (US): Hafner Press & New York Botanical Garden. http://www.jstor.org/stable/4393694 Subils, R. (2009) Bromeliaceae, Bromeliáceas. In: Kiesling, R. (ed.): Flora de San Juan, IV: 340–363, ills., keys. Mendoza (AR): Zeta Editores. Varadarajan, G. S. (1989) Genus Puya Molina (Pitcairnioideae): Relocation of several rare species and some preliminary remarks on geographic distributions and species divergence (continued). J. Bromeliad Soc. 39(1): 3–7, ills. Vásquez, R. C. & Ibisch, P. L. (2007) Puya pachyphylla sp. nov. (Bromeliaceae) and new synonyms of Puya tuberosa from Bolivia. J. Bromeliad Soc. 57(3): 102–111, ills. Wisnev, M. (2016) Some rarely seen Puya at the Huntington Botanical Gardens. J. Bromeliad Soc. 63(3–6): 159–179, ills. Zizka, G. [& al. 2002], Trumpler, K. & Zöllner, O. (2002) Revision of the genus Ochagavia (Bromeliaceae, Bromelioideae). Willdenowia 32(2): 331–350, ills., map, key. https://www.jstor.org/stable/3997641 Zizka, G. [& al. 2013], Schneider, J. V., Schulte, K. & Novoa, P. (2013) Taxonomic revision of the Chilean Puya species (Puyoideae, Bromeliaceae), with special notes on the Puya alpestris-Puya berteroniana species complex. Brittonia 65(4): 387–407, ills., key. https:// doi.org/10.1007/s12228-012-9290-9.

Sincoraea BROMELIACEAE U. Eggli and E. J. Gouda

Sincoraea Ule (Bot. Jahrb. Syst. 42: 191, 1909). Type: Sincoraea amoena Ule [typification by inference; this is the only taxon included in the genus]. — Bromelioideae — Lit: Louzada & Wanderley (2010: monograph, as Orthophytum); Louzada & Wanderley (2017: synopsis, key). Distr: E Brazil (Bahia, Minas Gerais: mostly Serra do Espinhaço Range). Perennial terrestrial saxicolous rosette plants, stoloniferous; Ros sessile or at most with a short stem covered by leaf sheaths; L tightly congested, (very) numerous, straight, arching or somewhat falcate, sheaths imbricate, distinct from the lamina, triangular, lepidote to glabrous, serrate, L lamina lineartriangular to narrowly triangular, (sub-) coriaceous to stiffly succulent, flat or canaliculate, (sparsely) lepidote on both faces or rarely adaxially glabrous, apex often a pungent spine, margins serrate with laxly to densely arranged spines; Inf sessile, simple or compound, inner rosette leaves and peduncular bracts becoming bright pink or red or red with a white or green base at the start of flowering and changing again to green or grey-green after anthesis; peduncular Bra gradually diminishing in size, outer

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] E. J. Gouda Curator University Botanic Gardens, Utrecht, The Netherlands e-mail: [email protected]

similar to the foliage leaves; floral Bra green or red, with pungent tip and serrate to serrulate margins; Fl sessile, hermaphrodite; Sep free, (sub-) erect, asymmetrical (rarely symmetrical), acute, acuminate, mucronate, with glandular trichomes (except S. hatschbachii); Pet free or forming an epigynous tube, suberect or upper parts spreading, white or greenish, 3–4.8  longer than wide, spatulate, obtuse, with 2 callosities laterally to the filaments, appendages sacciform, lacerate or digitate; St included; Fil filiform, unequal, outer free, inner adnate to the petals; Anth dorsifixed, obtuse; Ov inferior; Sty long-filiform; Sti  erect, 3-parted; Fr ovoid capsules with persistent sepals, 5–10  6–8 mm, greenish or white; Se ovate, striate, 1–2  0.6–1.5 mm, 30–100 per fruit. Sincoraea numbers 11 species, many of them  distinctly succulent. Until recently, the taxa here classified were treated as part of Orthophytum s.l. The widely circumscribed Orthophytum was found to be polyphyletic in several recent molecular studies (Louzada & al. 2014, Silvestro & al. 2014, Evans & al. 2015, Leme & al. 2017), where the “amoenum clade” (i.e. the species now segregated as Sincoraea) was found as sister to Lapanthus (not considered to include succulents), Cryptanthus and Orthophytum s.s. (see there for a fuller discussion). Recognizing the “amoenum clade” at generic level was first proposed in the unpublished thesis of Louzada (2012). Sincoraea differs from Orthophytum s.s. in its acaulescent rosettes with sessile, congested, simple or compound inflorescences, and consistently

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_95

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white flowers. At flowering time, Sincoraea species are notable for the highly coloured (red, pink, in some species with white or green base) inner rosette leaves and peduncular bracts, and the rosette as a whole can be regarded as a highly conspicuous pseudanthium. The colouration largely disappears again after anthesis is completed. Succulence: Many taxa grow in xeric habitats, often in association with cacti and other succulents. Horres & Zizka (1995) found a degree of succulence similar to that of many bona-fide succulents in S. burle-marxii. This and other succulent species have thickly fleshy, rigid leaves with considerable amounts of water storage tissue, while the other species have  coriaceous to leathery leaves. Succulence is very much a matter of degree, and only the most succulent taxa are covered in this handbook. Horticulture: Species of Sincoraea are only occasionally seen in cultivation. They easily grow in collections of succulents and xerophytes, and offer a spectacular sight when flowering, but are rather uninteresting otherwise. The flowers are diurnal. Individual flowers last only a single day, but flowering of an inflorescence extends over 2–3 weeks (Herndon 2018: for the non-succulent S. heleniceae). The following intergeneric hybrids have been obtained in cultivation (the respective nothogeneric Fig. 1 Sincoraea albopicta (Eggli 1249: Brazil; Bahia, near Mucugé). (Copyright: U. Eggli)

U. Eggli and E. J. Gouda

names still reflect their traditional classification as Orthophytum): Sincoraea  Aechmea = Orthomea, Sincoraea  Neoglaziovia = Orthoglaziovia, Sincoraea  Neoregelia = Neophytum, and Sincoraea  Cryptanthus = Orthotanthus (see separate entries for all of them in this handbook). S. albopicta (Philcox) Louzada & Wanderley (J. Bromeliad Soc. 66(1): 10, ill. (p. 11), 2017). Type: Brazil, Bahia (Storr 122 [CEPEC, K]). — Lit: Louzada & Wanderley (2010: 3–5, 14, with ills., as Orthophytum). Distr: E Brazil (Bahia: Chapada Diamantina: Mucugé, Andaraí); rock outcrops in grassland of Campo Rupestre vegetation, 700–1500 m. – Fig. 1.  Orthophytum albopictum Philcox (1985). Ros acaulescent or with a stem to 3  2.5–3.5 cm, offsetting with short stolons; L numerous, flatly spreading or slightly arched, coriaceous, sheaths broadly triangular, 0.6–3  1.1–3.2 cm, both faces glabrous or lepidote abaxially near the tip, margins serrate, L lamina linear-triangular, 8–24  0.8–2 cm, flat, green to greenish-red, basal part red at flowering time and adaxially with a white-lanate base, otherwise adaxially glabrous, abaxially densely lepidote, margins sublaxly serrate, Sp antrorse, 0.4–1.2 ( 2) mm; Inf sessile, compound and head-like, many-flowered; primary Bra deltoid-lanceolate, 1.7–3.8  1.6–1.7 cm, the

Sincoraea BROMELIACEAE

outer transitional to the inner rosette leaves and red with white-lanate base, the inner entirely white-lanate, all with serrate margins; floral Bra triangular,  15  10 mm, mucronulate, asymmetrical, ecarinate, green, lepidote, serrate; Fl tubular with slightly spreading limb; Sep narrowly triangular,  12–14  4–5 mm, mucronulate, asymmetrical, green, carinate, lepidote, entire; Pet linearspatulate,  18  4 mm, obtuse, white, basally with simple hairs, with 2 lacerate sacciform appendages  6 mm above the base, and with 2 conspicuous elongate callosities; outer Fil 9 mm, inner 3 mm; Anth 4 mm, yellow, not apiculate; Ov trigonous; Sty  1.3 mm; Fr and Se unknown. S. burle-marxii (L. B. Smith & Read) Louzada & Wanderley (J. Bromeliad Soc. 66(1): 10, ill. (p. 11), 2017). Type: Brazil, Bahia (Burle-Marx s.n. in Bogner 1311 [US]). — Lit: Louzada & Wanderley (2010: 7–9, 14, with ills., as Orthophytum). Distr: E Brazil (Bahia: Lençóis, Morro do Chapéu); rocky outcrops in grassland, usually in full sun, 900–1400 m. I: Braun & Esteves Pereira (2006: 161); Roguenant & al. (2016: 609); both as Orthophytum; Moonen (2018). – Fig. 2.  Orthophytum burle-marxii L. B. Smith & Read (1979); incl. Orthophytum burle-marxii var. seabrae Rauh (1985); incl. Orthophytum roseum Leme (2010). Ros acaulescent, stoloniferous, with short stems to 2.5  4 cm; L numerous, flatly spreading or arching, sheath broadly triangular or ovateFig. 2 Sincoraea burlemarxii (Supthut 8935: Brazil; Bahia, near Andaraí). (Copyright: U. Eggli)

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triangular, 1–3  1–4 cm, white-greenish, glabrous or rarely sparsely lepidote, serrate, L lamina lineartriangular, 12.8–53  1.5–2.5 cm, attenuate, strongly coriaceous, red or green, cinereous, adaxially lepidote except the glabrous base, abaxially densely lepidote, margins serrate, Sp antrorse, 1–2 mm; Inf sessile, compound and head-like; outer primary Bra transitional to the inner rosette leaves that become red-flushed during flowering time, the inner triangular or triangularlanceolate, 2.2–5.5  1–6 cm, red, coriaceous, sparsely lepidote, serrate; floral Bra some rudimentary, otherwise triangular, 14–48  8–37 mm, mucronulate, subcoriaceous, red, ecarinate, glabrous, serrate; Fl tubular with slightly spreading limb, with an epigynous tube  2 mm long; Sep narrowly triangular, 20  4–6 mm, acuminate or mucronulate, asymmetrical, red, carinate, sparsely lepidote, entire; Pet linear-spatulate, 18–25  6 mm, subacute, white, glabrous, with 2 sacciform lacerate appendages  7 mm above the base and conspicuous oblong callosities; outer Fil  14 mm, inner Fil  0.5 mm; Anth  4.5 mm, yellow; Ov trigonous, not further described; Fr and Se unknown. Similar to the non-succulent S. heleniceae with hardly thickened leaves and a green-coloured zone around the inflorescence (vs. red in S. burlemarxii). S. hatschbachii (Leme) Louzada & Wanderley (J. Bromeliad Soc. 66(1): 13, ill. (p. 12), 2017).

1104

Type: Brazil, Bahia (Hatschbach & Barbosa 56,827 [MBM]). — Lit: Louzada & Wanderley (2010: 9–10, 14, with ills., as Orthophytum). Distr: E Brazil (Bahia: Abaíra, Rio de Contas); rock outcrops, in full sun or in the shade near streams, 1300 m.  Orthophytum hatschbachii Leme (1995). Ros subacaulescent, stem to 3–4.2  3 cm, stoloniferous; L  30, horizontally spreading, sheaths broadly triangular to broadly ovate, 0.5–3  0.8–4 cm, greenish-white, glabrous, serrate, L lamina linear-triangular, 6.5–33  0.7–1.8 cm, stiffly coriaceous, slightly canaliculate, red with greenish-white base, adaxially glabrous or lepidote, abaxially sparsely lepidote, margins strongly serrate, Sp straight to antrorse, (1.7–) 2–3.5 mm,  15 mm apart; Inf sessile, simple, head-like, many-flowered; outer primary Bra transitional to the inner rosette leaves that become red-flushed during flowering time, not further described; floral Bra triangular, 20–25  9–23 mm, symmetrical or asymmetrical, coriaceous, green, glabrous, carinate, serrate, rudimentary floral bracts absent; Fl narrowly tubular, with a 2.5 mm long epigynous tube; Sep narrowly ovate, 12  3 mm, asymmetrical, mucronulate, green, carinate, glabrous, entire; Pet linear-spatulate, 17  4 mm, obtuse, erect, with only the very tips spreading, white, glabrous, with 2 sacciform lacerate appendages 4 mm above the base and 2 conspicuous callosities; outer Fil  10 mm, inner Fil  2.5 mm and adnate to the petals; Anth 3 mm, yellow, apiculate; Ov trigonous; Sty 10 mm; Fr and Se unknown. Similar to S. burle-marxii and the nonsucculent S. heleniceae. S. navioides (L. B. Smith) Louzada & Wanderley (J. Bromeliad Soc. 66(1): 14, 2017). Type: Brazil, Bahia (Foster & Foster 90 [GH, R, SP, US]). — Lit: Louzada & Wanderley (2010: 17–18, 25, with ills., as Orthophytum). Distr: E Brazil (Bahia: Jacobina); shaded rocks in dry forests.  Cryptanthopsis navioides L. B. Smith (1940)  Orthophytum navioides (L. B. Smith) L. B. Smith (1955).

U. Eggli and E. J. Gouda

Ros subacaulescent, stoloniferous, with a short stem 2  1.5 cm; L numerous, flatly spreading or arching, sheath triangular,  1  1.5 cm, white, glabrous, serrate, L lamina linear-triangular, 8–23  0.4–0.7 cm, attenuate, coriaceous, flat or slightly canaliculate, green, innermost reddish, both faces sparsely lepidote, margins serrate, Sp antrorse or retrorse, 1–1.5 mm; Inf sessile, simple and headlike, many-flowered; outer primary Bra transitional to the inner rosette leaves that become red-flushed during flowering time, not further described; floral Bra triangular,  20  10 mm, subcoriaceous, carinate, lepidote, serrate, rudimentary floral bracts absent; Fl narrowly tubular with spreading limb, with an epigynous tube  1.5 mm; Sep narrowly triangular, 20–30  5 mm, acuminate, green, with glandular trichomes, carinate, entire; Pet linear-spatulate, 25–30  4 mm, subacute, white, glabrous, with 2 sacciform lacerate appendages  3 mm above the base and 2 narrowly oblong callosities  as long as the filaments; outer Fil  20 mm, free part of inner Fil  9 mm; Anth yellow, 2–2.3 mm; Ov trigonous; Sty 20 mm; Sti capitate; Fr and Se unknown. This species is one of the most extreme xerophytes of the genus (Rauh 1990). It shows similarities with the hardly succulent S. mucugensis, which differs by having much shorter leaves and shorter flowers. S. navioides has been successfully crossed with Aechmea (see Orthomea) and Cryptanthus (see Orthotanthus).

References Braun, P. J. & Esteves Pereira, E. (2006) Succulent and xeromorphic Bromeliads of Brazil. Part 3. Encholirium maximum, Orthophytum burle-marxii and Dyckia goehringii. Cact. Succ. J. (US) 78(4): 160–164, ills. https://doi.org/10.2985/0007-9367(2006)78[160:SAX BOB]2.0.CO;2. Evans, T. M. [& al. 2015], Jabaily, R. S., Gelli de Faria, A. P., Oliveira F. de Sousa, L. de, Wendt, T. & Brown, G. K. (2015) Phylogenetic relationships in Bromeliaceae subfamily Bromelioideae based on chloroplast DNA sequence data. Syst. Bot. 40(1): 116–128. https://doi. org/10.1600/036364415X686413. Herndon, A. (2018) Flowering behavior of some Sincoraea species in cultivation. 1. Sincoraea heleniceae. J. Bromeliad Soc. 67(1): 50–60, ills.

Sincoraea BROMELIACEAE Horres, R. & Zizka, G. (1995) Untersuchungen zur Blattsukkulenz bei Bromeliaceae. Beitr. Biol. Pfl. 69(1): 43–76, ills. Leme, E. M. C. [& al. 2017], Heller, S., Zizka, G. & Halbritter, H. (2017) New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny. Phytotaxa 318(1): 1–88, ills., maps, key. https:// doi.org/10.11646/phytotaxa.318.1.1 Louzada, R. B. (2012) Revisão taxonômica e filogenia de Orthophytum (Bromeliaceae, Bromelioideae). São Paulo (BR): Universidade de São Paulo, unpublished Ph.D. thesis. Louzada, R. B. & Wanderley, M. G. L. (2010) Revision of Orthophytum (Bromeliaceae): The species with sessile inflorescences. Phytotaxa 13: 1–26, ills., key. Louzada, R. B. & Wanderley, M. G. L. (2017) Reestablishment of Sincoraea (Bromeliaceae). J. Bromeliad Soc. 66(1): 6–19, ills., key.

1105 Louzada, R. B. [& al. 2014], Schulte, K., Wanderley, M. G. L., Silvestro, D., Zizka, G., Barfuss, M. H. & PalmaSilva, C. (2014) Molecular phylogeny of the Brazilian endemic genus Orthophytum (Bromelioideae, Bromeliaceae) and its implications on morphological character evolution. Molec. Phylogen. Evol. 77: 54–64. https://doi.org/10.1016/j.ympev.2014.03.007 Moonen, J. (2018) Sincoraea burle-marxii — eine terrestrische Schönheit aus Bahia, Brasilien/a terrestrial beauty from Bahia, Brazil. Bromelie 2018(1): 4-6, ills. Rauh, W. (1990) Bromelien. Ed. 3. Stuttgart (DE): Verlag Eugen Ulmer. Roguenant, A. [& al. 2016], Lecoufle, M. & Raynal-Roques, A. (2016) Les Broméliacées. Approche panoramique d’une grand famille “américaine”. Paris (FR): Belin. Silvestro, D. [& al. 2014], Zizka, G. & Schulte, K. (2014) Disentangling the effects of key innovations on the diversification of Bromelioideae (Bromeliaceae). Evolution 68(1): 163–175. https://doi.org/10.1111/evo.12236

Tillandsia BROMELIACEAE E. J. Gouda

Tillandsia Linné (Spec. Pl. [ed. 1], 286, 1753). Type: Tillandsia utriculata Linné. — Tillandsioideae — Tillandsieae — Lit: Rauh (1970: ill. synopsis); Smith & Downs (1977: 665–1069, Fl. Neotropica); Rauh (1990: ill. synopsis); Shimizu (1992: ill. synopsis); Roguenant (2001: ill. synopsis); Isley (2009: ill. synopsis); Magalhães & Mariath (2012: seed anatomy); Donadío & al. (2015: morphological phylogeny subgen. Diaphoranthema); Barfuss & al. (2016: phylogeny/classification Tillandsioideae); Granados Mendoza & al. (2017: phylogeny). Distr: S USA to Chile and Argentina. Etym: For Elias Erici Tillandz (also Tillands or Til-Landz, later Tillander) (1640–1693), Swedishborn Finish physician and botanist at the Academy of Turku, and author of the first flora of Finland. Incl. Renealmia Linné (1753) (nomen rejiciendum, Art. 56.1). Type: Renealmia paniculata Linné. Incl. Caraguata Adanson (1763) (nom. illeg., Art. 52.1). Type: Tillandsia utriculata Linné. Incl. Bonapartea Ruiz & Pavón (1802). Type: Bonapartea juncea Ruiz & Pavón. Incl. Misandra Dietrich (1819) (nom. illeg., Art. 52.1). Type: Bonapartea juncea Ruiz & Pavón.

E. J. Gouda (*) Curator University Botanic Gardens, Utrecht, The Netherlands e-mail: [email protected]

Incl. Acanthospora Sprengel (1825) (nom. illeg., Art. 52.1). Type: Bonapartea juncea Ruiz & Pavón. Incl. Dendropogon Rafinesque (1825). Type: Renealmia usneoides Linné. Incl. Amalia Endlicher (1837) (nom. inval., ICN Art. 36.1a). Type: not typified. Incl. Buonapartea Sweet (1839) (nom. inval., Art. 61.1?). Type: Bonapartea juncea Ruiz & Pavón. Incl. Strepsia Nuttall ex Steudel (1841) (nom. illeg., Art. 52.1). Type: Renealmia usneoides Linné. Incl. Allardtia A. Dietrich (1852). Type: Allardtia cyanea A. Dietrich. Incl. Anoplophytum Beer (1854). Type: Tillandsia stricta Solander. Incl. Diaphoranthema Beer (1854). Type: Renealmia recurvata Linné. Incl. Platystachys K. Koch (1854). Type: Allardtia cyanea A. Dietrich. Incl. Phytarrhiza Visiani (1855). Type: Tillandsia duratii Visiani. Incl. Pityrophyllum Beer (1857). Type: Tillandsia erubescens Schlechtendal. Incl. Wallisia E. Morren (1870). Type: Tillandsia hamaleana E. Morren. Incl. Viridantha Espejo (2002). Type: Tillandsia plumosa Baker. Acaulescent or sometimes caulescent herbs of very variable habit; L rosulate or fasciculate or distributed along the stem, polystichous or less

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_96

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often distichous, entire, L lamina tongue-shaped to narrowly triangular or linear, leaf-scales symmetric to asymmetric and extended on one side; Inf usually with a distinct peduncle; floriferous Inf part various, usually consisting of distichously flowered spikes or sometimes reduced to a single polystichously flowered spike by the reduction of the spikes to single flowers, or rarely the whole inflorescence reduced to a single flower; floral Bra conspicuous to minute; Fl bisexual,  perigynous, seemingly pedicellate but in fact with an expanded receptacle looking like a pedicel; Sep convolute, usually symmetrical, free or equally or posteriorly joined; Pet free, usually naked; St of various lengths relative to petals and pistil; Ov superior, glabrous, ovules usually many, caudate; Fr septicidal capsules; Se narrowly cylindrical or fusiform, with a basal, plumose, straight, white appendage. Tillandsia is the largest genus in the family, with 690 species (Gouda & al. 2012+). The genera Tillandsia and Vriesea are traditionally distinguished from each other by the absence (in the first) or presence (in the latter) of ligulae on the claw of the petals. This lead to the transfer of numerous species from Tillandsia to Vriesea, resulting in a growing group of Tillandsia-like Vrieseas. The character has been found to be marginally useful, however, to distinguish natural groups within the family (Brown & Terry 1992, Schulte & Zizka 2008). Moreover, in at least one species (T. incurva Griseb.), ligulae are present in some populations, but absent from others (Gouda 1987). Recently, this group has been transferred back to Tillandsia (Grant 1993). According to Magalhães & Mariath (2012), the structure of the plumose seed appendages could provide a better character to separate Tillandsia from Vriesea, since seeds of the former have a double superimposed parachute-like plumose appendage derived from the exo- and mesostesta, as well as from the endotesta, while Vriesea seeds have a single such structure without participation of the endotesta. The derivation of the plumose appendage from the testa was probably first described by Morra & al. (2002). In the last monograph of the family (Smith & Downs 1977), 7 subgenera were recognized.

E. J. Gouda

Subgenus Pseudo-catopsis was subsequently elevated to genus level as Racinaea by Smith & Spencer (1993). Recent molecular DNA work by Barfuss & al. (2016) supports this move and shows that the traditional subgeneric classification is not fully supported. As a consequence, subgenus Aerobia (Mez 1896) was resurrected for 50 species that were previously placed either in subgen. Allardtia or subgenus Anoplophytum. The revised subgeneric classification is as follows: [1] Subgen. Allardtia (A. Dietrich) Baker 1888: St included, exceeding the claw of the petals; Sty slender, much longer than the ovary; Fil straight. — 200 species, 12 of them succulent. [2] Subgen. Anoplophytum (Beer) Baker 1887: St included, equalling the claw of the petals; Sty slender, much longer than the ovary; Fil strongly plicate. — 104 species, 13 of them succulent. [3] Subgen. Phytarrhiza (Visiani) Baker 1887: Pet blades broad, conspicuous; St deeply included; Sty short and stout; L lamina flat or terete. — 39 species, 7 of them succulent. [4] Subgen. Diaphoranthema (Beer) Baker 1878: Pet blades narrow, inconspicuous; St deeply included; Sty short and stout; L lamina terete, densely cinereous-lepidote. — 30 mostly tiny to small species, 22 of them succulent. Donadío & al. (2015) have recently published a phylogeny of the subgenus based on morphology. [5] Subgen. Tillandsia: St exserted, exceeding the petals; Pet erect at anthesis or nearly so, relatively firm; L lamina narrowly triangular in most species. — 246 species, 29 of them succulent. [6] Subgen. Pseudalcantarea Mez 1934: St exserted, exceeding the petals; Pet spreading at anthesis, rapidly becoming flaccid; L lamina tongue-shapped to narrowly triangular. — 5 species, none of them succulent. [7] Subgen. Aerobia Mez 1896: St included, often emerging from the throat of the corolla; Sty slender, many times longer than the ovary; Fil straight or plicate; Pet spreading at anthesis, often with distinct blade; L lamina narrowly triangular. — 50 species, most of

Tillandsia BROMELIACEAE

them succulent, but not treated here for space reasons. Succulence: Most subgenera contain species that can be variously described as possessing succulent or semi-succulent features at least to some degree, i.e. showing considerable amounts of water-storing tissues in the leaves. Most mesophytic species have funnel-shaped rosettes with inflated leaf sheaths compounding water, and these are — though superficially appearing to have water-storage capacity — not succulent. All xerophytic species are succulent to some extent, but only 83 species are selected here because they have obviously fleshy leaves. They are all from semi-arid to arid areas, where they grow as epiphytes on cacti and small trees, or are saxicolous on rocky surfaces. In the coastal deserts of Peru and Chile, some species grow on the sand dunes in the so-called Loma vegetation belt, without any other vegetation. Pollination: Many species, esp. from the largest subgen. Tillandsia, with tubular corolla and exserted stamens and stigma are pollinated by hummingbirds. Birds are attracted by the brightly coloured bracts (often red and yellow) and scentless flowers. Flowers are available over a longer period of time and individual flowers are open for only one or a few days. This is ideal for “trap-line” pollination by birds, where a bird flies a certain route, probably several times a day, from one plant to another. Another group of species, from subgen. Phytarrhiza, have open flowers with broad coloured petal blades, and are often strongly fragrant. The bracts are mostly uncoloured and do not have a role in attracting pollinators. The petals have a narrow claw that together form a narrow tube-like structure, accessible only to the long and very slender tongues of moths and butterflies. Most species of Tillandsia are self-incompatible and are dependent on pollinating animals for reproduction. The flowers of monocarpic species, however, are autogamous when not pollinated. Also the small-flowering plants from subgen. Diaphoranthema are (partly) autogamous, which explains the pronounced variety of forms within a species and within an area. The flowers of some

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species of this group are fragrant, e.g. T. usneoides, fragrant during the night only, and probably pollinated by small moths according to Gouda (1995). Horticulture: Tillandsia is the subject of considerable horticultural interest. Esp. the xerophytic species are frequently seen in cultivation, no doubt in part due to the availability of synoptical literature aimed at hobby collectors, such as Rauh (1990). Several profusely illustrated accounts (e.g. Isley (1987), Isley (2009) or Shimizu (1992)) have further added to the popularity of the genus. T. aizoides Mez (in A. & C. de Candolle, Monogr. Phan. 9: 866, 1896). Type: Argentina, La Rioja (Hieronymus & Niederlein 850 [B, CORD]). — Distr: SW and S Bolivia (Potosí, Tarija), Argentina (widespread); epiphytic on twigs, or sometimes saxicolous, 120–2800 m. [4] Plant small, long-caulescent, forming dense clusters or few-branched; stem to 5 cm, many from a single point; L densely rosulate, usually 1 ( 2) cm  2 mm; L sheaths 0.5 cm, broadly ovate, scarious, with broad nerveless margins, 4to many-veined, glabrous except at the distal end, distinct from the lamina; L lamina stout, rigid, densely cinereous-lepidote, erect to somewhat divergent, sometimes slightly secund, angularsubulate, convex below, with a furrow adaxially, apex pungent; Inf 1-flowered, often with a reduced flower at the apex, peduncle naked except for 1 or 2 bracts below the flower or with a single bract midway, from very short to 2 cm, angled, glabrous; upper peduncular Bra like the floral bracts, elliptic, often carinate, sulcate and strongly veined, glabrous, acute; floral Bra erect, shorter than the sepals, 5.5  3.5 mm, elliptic, membranous, at least 3-veined, ecarinate, glabrous, green, broadly rounded or subapiculate; Sep 7–8  3 mm, elliptic or obovate-oblong, scarious, 5- to 7-veined, all connate for 1–2.5 mm, ecarinate, green tinged red, glabrous, broadly rounded or subapiculate; Pet slightly longer than the sepals, spatulate, 13  3 mm, cream-coloured with purple-red speckles, brownish when dry, blade obscure or distinct and then suborbicular; St all of equal length, 6 mm, deeply included, exceeding the style; Fil flaccid; Anth basifixed, 1.4 mm; Ov

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Fig. 1 Tillandsia albertiana. (Copyright: E. J. Gouda)

4 mm, subovoid; Sty slender, shorter than the ovary; Sti capitellate. T. albertiana Vervoorst (Bromeliad Soc. Bull. 19: 121, 128, ills., 1969). Type: Argentina, Salta (Vervoorst 7255 [LIL, US]). — Distr: NE Argentina (Salta); lithophytic on rocks and cliffs, altitude not recorded. – Fig. 1. [2] Plant short- to long-caulescent, forming dense clusters or laxly pulvinate; stem much branched, 3 mm ∅; L distichous; L sheaths imbricate, making the stem appear 6 mm thick, 2–2.5  1.4 cm, ovate, clasping the stem with overlapping margins, with broad membranous margins, veined or slightly rugose, densely appressedlepidote abaxially and glabrous adaxially except at the distal end, green or brown, tapering into the lamina; L lamina linear-triangular, fleshy, at the base with extending trichomes at the margins, 5–9 ( 17)  0.4–0.8 cm, dark green flushed with red or red-striated at the base, appressed-lepidote, divergent to less often spreading, veined at the base, strongly canaliculate, convex or involutesubulate when dry, apex abruptly obtuse; Inf 1-flowered (with a short extended rachis), erect; peduncle 2 cm, concealed by the leaves; peduncular Bra 2, excl. lamina 1.8  0.85 cm, the sheathing part glabrous except abaxially towards the apex, with foliaceous lamina, obtuse; floral Bra (excl. acumen) 17–20 ( 23)  9 mm, ovate-

oblong, clasping the flower with slightly overlapping margins, margins hyaline, veined, glabrous at the base and adaxially to subdensely appressedlepidote abaxially, green or with reddish apical margins, rounded and fleshy-acuminate; Fl subsessile; Sep 17–24  4–6 mm, narrowly oblong, slightly contracted in the centre, thin, with membranous margins, faintly veined, but distinctly veined when dry, free, ecarinate, green, tinged wine-red at the apex, glabrous or with a few trichomes abaxially, rounded; Pet claw narrow with a broad suborbicular blade or spatulate, 28–44  to 11 mm, margins slightly crenulate, wine-red to bright red, rounded or obtuse; St all of equal length, 17–22 mm, included and longer than the style; Fil ribbon-like, flat at the base but upwards fleshy, not plicate, green towards the apex; Anth subbasifixed, 4 mm, linear, apex obtuse, yellow; Ov 5  2.5 mm, triangularovoid (broadest near the base) to prismatic, gradually contracted into the style; Sty slender, 8–17 mm; Sti linear, erect, weakly conduplicate. T. andicola Gillies ex Baker (J. Bot. 16: 239, 1878). Type: Argentina, Mendoza (Gillies s.n. [K, CGE, GH]). — Distr: Argentina (Catamarca, Buenos Aires, San Juan, Mendoza, Río Negro); epiphytic, altitude not recorded. [4] Plant flowering to 30 cm tall; stem to >20 cm, branched esp. near the apex; L lax, weakly rosulate

Tillandsia BROMELIACEAE

or distichous, 4 ( 6) cm, cinereous-lepidote with subappressed basally produced scales; L sheaths densely imbricate, 5 mm apart on the stem, making it appear 3–4 mm ∅, much broader than the lamina, suborbicular, occasionally with glabrous broad veinless margins, ciliate with long scales, several-veined, usually lepidote except at the extreme base or wholly glabrous in age; L lamina 2 mm wide, recurved, veined when dry, subulate, apex acuminate, pungent; Inf 1- or 2-flowered (then internode 8 mm long and rachis extended behind the terminal flower), densely lepidote; peduncle 0–6 cm, bractless or with 1 bract, densely lepidote; peduncular Bra lanceolate, involute, lepidote, acuminate; floral Bra exceeding the sepals in the first flower to shorter than the sepals in the second one, to 14  7 mm, ovate to lanceolate, clasping and closely enveloping the flower, veined, ecarinate, 2 as long as the internodes, densely appressedlepidote, cinereous to green, acuminate or apiculate; Fl subsessile; Sep 10–12  3.5 mm, lanceolate-oblong, with broad hyaline margins, veined, adaxial ones connate for 6 mm, the others less connate, adaxial ones with a strong midvein, green, lepidote from the middle to the apex, apiculate; Pet tongue-shaped, 14  3 mm, pale white, the blade narrow, rounded; St deeply included, exceeding the style; Anth basifixed, 1.5 mm, red-orange; Ov 3.5 mm, ovoid, contracted into the style; Sty stout, 3 mm; Sti shortly lobed. T. angulosa Mez (in A. & C. de Candolle, Monogr. Phan. 9: 868, 1896). Type: Argentina, La Rioja (Hieronymus & Niederlein 851 [B, CORD, F [photo]]). — Distr: C Argentina (La Rioja, San Juan, Mendoza); epiphytic in xeric environments; 1160–1200 m. [4] Plant small, pulvinate, stem to 3 cm, richly branched at the base; L weakly polystichous or somewhat distichous (to spirally distichous), to 1.5  0.2 cm, coarsely densely cinereous-lepidote; L sheaths suborbicular, thin, with broad nerveless margins, many-veined, densely lepidote to glabrous at the base; L lamina subtriangular, spreading to recurved, subulate, angled, lower canaliculate or grooved, apex mucronate; Inf 1(rarely 2–) flowered; peduncle almost none or

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short and concealed by the leaves, with a single bract at the base or bractless; peduncular Bra ovate, involute, prominently veined, densely lepidote, acute; floral Bra slightly to distinctly shorter than the sepals, 12  7 mm, ovate, clasping the flower, 5- to 9-veined, sparsely lepidote, acute; Sep 11  4 mm, triangular-ovate, 7to 9-veined, almost free or evenly shortly connate, lepidote; Pet elliptic, 10–13  3.5 mm, brown to yellow, obtuse; St included, slightly exceeding the style with part of the anthers; Fil 4–5  0.2 mm, whitish; Anth basifixed, 3  to 0.5 mm; Ov 3.5  2–3 mm, obovoid-pyriform, gradually contracted into the style; Sty slender, 1.5 mm; Sti disc-shaped, shortly lobed. T. araujei Mez (in Martius, Fl. Bras. 3(3): 600, t. 112, fig. 2, 1894). Type [lecto]: Brazil, Rio de Janeiro (Glaziou 16457 [B, B [photo], P]). — Distr: SE & S Brazil (Rio de Janeiro, São Paulo, Rio Grande do Sul), Paraguay; saxicolous or very rarely epiphytic, 0–850 m. – Fig. 2. Incl. Tillandsia araujei var. minima E. Pereira & I. A. Penna (1980). [2] Plant caulescent, flowering 15–30 cm tall; stem simple or few-branched, often curved; L densely polystichous along the stem, 3–7 cm; L sheaths short, broadly triangular, glabrous at the base, white; L lamina rigid, appressed-lepidote, all upwardly secund, keeled, angled-subulate, trichomes with a brown centre, apex attenuate and pungent; Inf simple; peduncle completely covered by the bracts or partly visible, erect or ascending, much exceeding the leaves, slender, glabrous, exposed part red; peduncular Bra with thick linear lepidote lamina, imbricate, elliptic or obovate, membranous, rose-coloured, Inf spikes sublax, 3–5 cm, 5- to 12-flowered; floral Bra exceeding the sepals, 20–23  10–12 mm, ovate, membranous, ecarinate, glabrous except the apex in the lower ones, dark pink-red, broadly acute or the lower ones apiculate; Fl sessile, polystichous, erect to slightly divergent, 30 mm; Sep (12–) 15–17  4.5 mm, narrowly ovate to lanceolate, with broad hyaline margins, adaxial ones highly connate for 11 mm, adaxial ones carinate, rosecoloured with green base, acute; Pet spatulate, 20–32  8 mm, white, blade tapering into the

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Fig. 2 Tillandsia araujei. (Copyright: E. J. Gouda)

Fig. 3 Tillandsia argentina. (Copyright: E. J. Gouda)

cuneate claw, spreading, rounded or obtuse; St 17 mm, included; Fil inflated towards the apex, flaccid, lightly plicate near the apex to strongly plicate in the middle; Anth basifixed, 2 mm, yellow; Ov 3  2 mm, obovoid (to pyriform), abruptly contracted into the style; Sty slender, 16 mm, exceeding the stamens; Sti weakly conduplicate, erect. There is some variation in succulence between different populations; the small, short-leaved form (var. minima) is more succulent than the larger ones.

Incl. Tillandsia unca Baker (1887) (nom. illeg., Art. 53.1). [2] Plant distinctly short-stemmed, flowering 7–13 cm tall, rosette solitary or few-branched; L densely rosulate, the outer greatly reduced, 5.5–10 ( 13) cm; L sheaths 8–11  8–16 mm, broadly triangular-ovate, thin, lustrous, at least the lower glabrous, passing almost imperceptibly into the lamina; L lamina very narrowly triangular, linear, rigid, 3–6 mm wide, appressed-lepidote, divergent and often secundly curved upward, often pulvinate, canaliculate, obtusely keeled below or angular-subulate, apex abruptly acute and pungent; Inf always simple, shorter than to exceeding the leaves; peduncle hidden by the inner leaves and bracts, erect or ascending, shorter than the leaves, 0–6 cm, glabrous; lowest peduncular Bra foliaceous, imbricate, lanceolate, chartaceous, strongly nerved, glabrous or sparsely pale-lepidote near the apex, stramineous, acuminate; Inf spikes

T. argentina C. H. Wright (Bull. Misc. Inform. Kew 1907: 60, 1907). Type: Argentina, Córdoba (Stuckert s.n. [K, GH [photo]]). — Distr: S Bolivia (Chuquisaca, Tarija), N Argentina (Catamarca, Córdoba, Jujuy, Salta, Tucumán, La Rioja); saxicolous or epiphytic in dry habitats, 450–1950 m. – Fig. 3.

Tillandsia BROMELIACEAE

(2–) 3.5–4.5  (0.7–) 1–1.5 cm, broadly (ob-) lanceolate, strongly complanate, (2- to) 4- to 7flowered, rachis strongly 4-angled, nearly straight, glabrous, partly exposed; floral Bra imbricate, much exceeding the sepals, 19–25  7–10 mm, lanceolate or triangular-ovate, subcoriaceous, even or somewhat veined, sublustrous, ecarinate or carinate towards the apex, so narrow as to expose the rachis in part, glabrous throughout, rose-coloured or green, acuminate; Fl subsessile; Sep 12–18  4–5.5 mm, narrowly ovate or elliptic, subequal, free, adaxial ones carinate, green with reddish apex, glabrous, apiculate or broadly obtuse; Pet linear or subspatulate, 30–34  6–7.5 mm, margins entire, bright rose-red except at the base, blade scarcely distinct, divergent to strongly recurved, obtuse or rounded-emarginate; St 20 mm, included; Fil broadly ribbon-like, flaccid, strongly plicate near the apex, white; Anth basifixed, 4–5 mm, linear, cream-coloured to yellow; Ov 2.5 mm, subprismatic or subglobose, abruptly contracted into the style; Sty slender, 19–24 mm, exceeding the stamens; Sti spreading or recurved, conduplicate. T. arhiza Mez (in A. & C. de Candolle, Monogr. Phan. 9: 855, 1896). Type: Paraguay, Paraguarí (Balansa 4747 [P, GH [photo]]). — Distr: E Brazil (Minas Gerais), S Paraguay (Paraguari, Guairá); saxicolous, 540 m. Incl. Tillandsia rupestris Mez (1896)  Tillandsia arhiza var. rupestris (Mez) Hassler (1919); incl. Tillandsia rupestris var. pendens Chodat & Vischer (1916) (nom. inval., ICN Art. 32.1c); incl. Tillandsia breweri hort. ex D. Butcher (1996) (nom. inval., ICN Art. 29.1). [3] Plant stout, caulescent; stem to 60 cm; L spirally arranged, 20 cm, white-tomentose-lepidote; L sheaths elongate, broadly elliptic, amplexicaul, glabrous inside and below the middle outside and densely lepidote upwards; L lamina very narrowly triangular, 7–9 mm wide, densely ferrugineous or cinereous -lepidote or tomentoselepidote, erect or divergent and recurved towards the apex, canaliculate or involute-subulate above the middle, apex filiform attenuate; Inf often simple or depauperate-compound, distichous; peduncle erect, exceeding the leaves, slender, glabrous;

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peduncular Bra erect, equalling or exceeding the internodes, sublanceolate, tubular-involute, densely lepidote, acute, apiculate; primary Bra like the peduncular bracts, shorter than the spikes, lepidote; Inf spikes densely flowered, to 7.5 cm, 1 cm wide, lanceolate, (single/principal one) to linear, 6- to 12-flowered, rachis angled, slightly flexuous; floral Bra erect, subimbricate, from distinctly shorter than to slightly exceeding the sepals, 15 mm, narrowly elliptic, clasping the flower, chartaceous, veined, glabrous or the lowest sparsely lepidote, apiculate; Sep 12  5 mm, elliptic, coriaceous, even, subevenly short-connate for 3 mm, glabrous, acute, obtuse; Pet claw narrow with suborbicular blade, 23 mm long, violet; St deeply included, exceeding the style; Anth linear, apex acute; Ov elongate, prismatic. T. ariza-juliae L. B. Smith & J. Jiménez Almonte (Phytologia 6: 433, t. 1, figs. 1–2, 1959). Type: Dominican Republic, La Vega (Ariza Julia s.n. [US, Herb. J. J. Jiménez]). — Distr: Dominican Republic (Santiago Rodríguez, La Vega, Santiago Rodríguez); epiphyte in pine forest or on shrubs, 200–900 m. [5] Plant stemless, flowering 19 cm tall; L 12, densely cinereous-lepidote; L sheaths forming an ovoid pseudobulb 3–4 cm long, orbicular, greatly inflated, green tinged with purple near the margin, abruptly contracted into the lamina; L lamina 3–4 as long as the sheath, erect to convergent, 9  20 cm  2–4 mm, involutesubulate and nearly straight, narrowly canaliculate, apex attenuate; Inf simple or rarely branched; peduncle erect; lowest peduncular Bra subfoliaceous, imbricate, the uppermost broadly ovate, subchartaceous, apiculate; primary Bra shorter than the axillary branches; Inf spikes rather laxly to densely flowered, 8 cm, lanceolate, complanate, 2to 5- (to 8–) flowered, acute, rachis slightly alate, slender; floral Bra imbricate at anthesis, 30 mm, ovate, subchartaceous, veined, ecarinate, >3 as long as the internodes, densely appressed-lepidote, roseate, acute; Fl subsessile; Sep 20 mm, oblong, adaxial ones shortly connate, lepidote, apiculate; Pet forming an erect tube, size unknown, purple; St exserted.

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T. baileyi Rose ex Small (Fl. Southeast. US, 246, 1903). Type: USA, Texas (Bailey “26” [226] [US]). — Distr: S USA (Texas); epiphytic in scattered oak forest, 0–1080 m. [5] Plant stemless, flowering 20–40 cm tall, forming dense clusters; L several, to 40 cm, densely appressed-cinereous lepidote throughout; L sheaths forming a pseudobulb 2–5 cm long, ample, ovate, often conspicuously ciliate-lepidote, merging into the lamina; L lamina linear, at the base 5 mm wide, contorted or involute-subulate, apex attenuate; Inf simple, shorter than to as long as the leaves; peduncle nearly fully concealed by the bracts, erect or ascending, shorter than the leaves, often flexuous, internodes 2–4 mm, upwards decreasing in size, glabrous to cinereous-lepidote upwards at least near the nodes, green; peduncular Bra foliaceous but smaller than the leaves, suberect, few (4), basal part (excluding lamina) slightly or not amplexicaulous,  as long as the internodes, upper ones 2.3–2.7  1.1 cm, broadly ovate, abaxially densely pale-lepidote, green, shortly and weakly acuminate; Inf spikes, often ascending, (4–) 8–10  0.8–1.2 cm, linear, complanate, subdensely 6- to 17-flowered; floral Bra erect, loosely imbricate, exceeding the sepals, 20–23  8–10 mm, narrowly ovate-triangular, evenly incurved, closely enveloping the flowers, subchartaceous, margins membranous and with extended scales, smooth or veined, ecarinate, narrow and exposing the rachis in part, 1 mm, obovoid (to pyriform), abruptly contracted into the style; Sty stout, shorter than to about as long as the ovary; Sti capitellate, lobes short, concave and erect. A very variable species including many forms, some of them more succulent than others. Infraspecific taxa are no longer recognized (Till 1989, 1991), with the exception of fa. virescens, which is recognized as separate species, T. virescens. T. virescens is very similar, however, intermediate forms occur and separation is probably not warranted. T. caput-medusae E. Morren (Belgique Hort. 30: 90, 1880). Type: K [Morren icono]. — Distr: Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, W & C Panama; epiphytic, 15–2400 m. Incl. Tillandsia langlassei Poisson & Menet (1908). [5] Plant stemless, flowering 15–25 ( 40) cm tall; L often exceeding the inflorescence, densely subappressed-lepidote; L sheaths forming an ovoid pseudobulb, (broadly) ovate to triangularovate, strongly inflated, with membranous margins, adaxially densely ferruginous-lepidote with dark-centered scales, brown to dark brown inside, upwards with purple-red margins, merging into the lamina; L lamina linear-triangular, 10–25 cm, to 1.5 cm wide, involute-subulate or usually much contorted, apex attenuate; Inf simple or digitately composed of 2–6 spikes; peduncle erect, ascending, slender; peduncular Bra leaf-like, densely imbricate; primary Bra usually smaller than the floral bracts, broadly ovate, with a little lamina or without, lepidote; Inf spikes bearing several sterile bracts at the base, suberect to spreading, often curved, to 18 cm, linear-lanceolate, 6- to 12-flowered, acute, rachis angled, slender, nearly straight, glabrous; floral Bra suberect or divergent, imbricate, shorter than to slightly exceeding the sepals, 12–15 ( 20)  8–9 mm, ovate-lanceolate, incurved esp. at the apex, chartaceous, even or veined when dry, lustrous, carinate but sometimes obscurely so (not thickened), exposing most of the rachis, barely more than 2 as long as the internodes, nearly or quite glabrous or obscurely sparsely lepidote on both faces, red or pink or sometimes green, rounded

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or broadly obtuse; Fl subsessile; Sep incurved especially at the apex, 13–14  5 mm, oblong, thin-coriaceous to chartaceous, veined only when dry, lustrous, adaxial ones connate for 5–6.5 mm, adaxial ones sharply carinate, glabrous or very sparsely lepidote adaxially only, rounded or broadly obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 28–44  7 mm with narrow base only 2 mm wide, violet (rarely white), with cuneate claw, rounded and then abruptly obtuse; St unequal in length, 30 mm exserted; Fil 53–57 mm, thin and slender and flattened at the base and terete at the distal end; Anth dorsifixed just below the middle, (fresh) 6  1 mm, linear, base bilobed, apex apiculate; Ov 4  2 mm, ovoid, contracted into the style; Sty exceeding the stamens. T. castellanii L. B. Smith (Contr. Gray Herb. 104: 80, t. 3, figs. 17–19, 1934). Type: Argentina, Córdoba (Castellanos s.n. in BA 1576 [GH, BA]). — Distr: NW & C Argentina; epiphytic, 1000–3000 m. [4] Plant flowering to 15 cm tall, forming dense clusters; stem simple, few-branched, many from a single point; L dense, distichous, to 4–5 mm; L sheaths broadly ovate, strongly veined, densely lepidote with a ciliate margin of elongate scales; L lamina sublinear, 2–3 mm wide, spreading,

Fig. 4 Tillandsia chiapensis. (Copyright: E. J. Gouda)

E. J. Gouda

recurved, subulate with pungent apex, cinereouslepidote with spreading scales; Inf 1- to 2-flowered; peduncle bractless, pseudo-axillary, evident, to 3 cm, slender, glabrous; floral Bra shorter than the sepals, (5.5–) 6.5–9.5 ( 10.5)  (5.5–) 5.8–6.8 ( 7) mm, ovate, strongly 9- to 11-veined, glabrous or upper part lepidote, broadly acute or short-laminate; Fl subsessile, not fragrant; Sep 7.5–8.5 ( 9)  2.9–3.3 ( 3.5) mm, obovatelanceolate, strongly 7- to 9-veined, evenly shortconnate or abaxial one free, glabrous, rounded or obtuse; Pet ovate-lanceolate, 7–8 ( 10)  1.5–2 mm, veined, yellowish, rounded or rarely obtuse; St deeply included, exceeding the style; Fil 3–3.7  0.2 mm; Anth basifixed, 2.1–2.2  0.3 mm, whitish; Ov obovoid (to pyriform), 3.2–3.4 mm, abruptly contracted into the style; Sty stout, 1.6–1.7 mm, as wide as the ovary; Sti obscurely lobed. T. chiapensis C. S. Gardner (Selbyana 2(4): 338, ill., 1978). Type: Mexico, Chiapas (Gardner 211 [SEL]). — Distr: SE Mexico (Chiapas); saxicolous, 600–1000 m. – Fig. 4. [5] Plant stemless, offsetting from the base, eventually forming clumps; L forming a spreading rosette, numerous, 30 cm; L sheaths 4.5 cm wide, suborbicular, often brown-lepidote in the distal part, pale castaneous; L lamina narrowly

Tillandsia BROMELIACEAE

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Fig. 5 Tillandsia circinnatoides. (Copyright: E. J. Gouda)

triangular, fleshy, 2.5 cm wide at mid-blade, coarsely and densely cinereous-lepidote, curved to recurved, strongly canaliculate or the margins involute towards the base, apex attenuate, trichomes 0.5 mm long, suborbicular; Inf simple,  equalling or exceeding the leaves; peduncle short and concealed by the leaves and bracts; peduncular Bra polystichous, the lowest leaflike, densely imbricate, tinged or wholly rosecoloured; spikes suberect or becoming decumbent after flowering, lanceolate; floral Bra distichous, densely imbricate, >2 as long as the sepals, 40  26 mm, ovate-elliptic, incurved at the apex, finely veined when dry, ecarinate or slightly so at the apex, concealing the rachis, 3–4 as long as the internodes, subdensely or densely lepidote, pink, acute or obtuse; Fl distichous, suberect (contiguous with the bracts in the lower part); Sep incurved, 15  7.5 mm, elliptic or obovate, coriaceous esp. at the base, veined when dry, free or nearly so, adaxial ones carinate, acute or obtuse; Pet forming an erect narrow tube clasping the filaments, with revolute apex, narrowly tongueshaped, 75 mm, 8.5 mm wide in the uppermost , 3 mm wide at the base, blue-violet, obtuse; St in 2 whorls of unequal length, 75–80 mm, shorter than the style; Fil subterete towards the apex, purple; Anth dorsifixed at from the base, 3.5  1 mm, oblong-ellipsoid, apex obtuse; Ov narrowly ovoid, merging into the style; Sty slender; Sti conduplicate-spiral, white.

T. circinnatoides Matuda (Cact. Succ. J. (US) 45: 187, figs. 4, 4a, 5, 1973). Type: Mexico, Guerrero (Matuda 38432 [MEXU, US]). — Distr: S & E Mexico, NW Costa Rica; epiphytic on cacti, trees, and bushes in dry areas, 600–2200 m. – Fig. 5. [5] Plant stemless, flowering 10–20 cm tall, often forming dense clusters; L few, the outer ones reduced and without lamina, the others 7–16 cm, appearing veined by trichomes in longitudinal rows, covered with appressed cinereous scales; L sheaths forming an elongate pseudobulb, large, ovate, merging into the lamina; L lamina narrowly triangular, erect or curved, grooved, involute-subulate; Inf simple; peduncle erect, 7.5 cm, concealed by the leaves; lowest peduncular Bra leaf-like, the upper like the floral bracts or slightly larger, densely imbricate, acuminate; spikes curved or erect, 5–10 cm, 1.4–2 cm wide, lanceolate to linear, complanate, attenuate, rachis hidden or (slightly) partly exposed; floral Bra densely imbricate, exceeding the sepals, 20–30  9.5 mm, elliptic, thin, finely veined when dry, scarcely or not at all carinate towards the apex, 3–4 as long as the internodes, densely cinereous-lepidote except the base and margins and less dense on the inside, pale roseate, broadly acute or apiculate; Fl subsessile, contiguous with each other and/or the rachis; Sep 19–25  4–6 mm, lanceolate, abaxial one chartaceous and adaxial ones with a rigid subcoriaceous keel, subfree or adaxial ones connate for up to 5 mm, adaxial ones

1120

carinate from base to apex, lepidote with few large scales abaxially, or glabrescent; Pet forming an erect narrow tube clasping the filaments with revolute apex, tongue-shaped, 40–45  5 mm, pale blue to dark violet in the upper , rounded; St long-exserted; Fil twisting at the base, flat and exposed part slightly inflated and (sub-)terete, bluish in the upper ; Ov ovoid, abruptly contracted into the style; Sty slender, exceeding the stamens; Sti conduplicate-spiralized, longpapillose. T. colganii Ehlers (J. Bromeliad Soc. 47: 103, ill., 1997). Type: Bolivia, Santa Cruz (Ehlers EB951002 [WU]). — Distr: E Bolivia (Santa Cruz); saxicolous, 2700–3200 m. [2] Plant short stemmed or with a rhizomatous stem, flowering 10–15 cm tall, often branched; L rosulate, 12–14, 3–7 cm long (outer ones reduced with short acuminate lamina), coriaceous to subsucculent, margins with asymmetric trichomes, covered with appressed cinereous scales, greengrey looking striped; L sheaths erect or suberect, 2–3  1–1.8 cm, ovate, passing almost imperceptibly into the lamina; L lamina narrowly triangular, 1–1.5 cm wide at the base, sometimes slightly secund or spreading or recurved, abaxially with a median keel and strongly veined, subulate or canaliculate, apex attenuate; Inf simple, exceeding the leaves; peduncle completely covered by bracts, erect, 0–2 cm and concealed by the leaves, stout, lepidote; lowest peduncular Bra almost leaf-like, imbricate, the apical ones like the floral bracts, becoming glabrous, red, attenuate or acute; spikes with a sterile bract at the base, 3–4 cm, 5–8 ( 10) mm wide, lanceolate, subterete, 2- or 3-flowered, acute, rachis 4-angled, nearly straight, glabrous, hidden at anthesis, green; floral Bra imbricate, exceeding the sepals, 16–25 ( 28)  8–10 mm, ovate-triangular, submembranous, adaxially veined, ecarinate, glabrous or the apical 4–6 mm finely grey-lepidote, sublustrous winered, acute or cuspidate; Fl sessile, distichous, not fragrant; Sep 14–16  4.5–6 mm, narrowly elliptic, membranous, thickened along the keels, free, adaxial ones carinate, greenish with pink tips, acute; Pet erect and throat open, 36–40  5 mm, at the base 3 mm wide, margins entire, violet to

E. J. Gouda

white at the base, blade spreading to recurved, obtuse; St just visible in the throat of the corolla; Fil 25–30  0.5 mm, flat, thin towards the base, once transversely plicate or straight; Anth basifixed, 4–5 mm, linear, apex acute, yellow; Ov 4  2.5 mm at the base, ovoid; Sty surpassing the stamens, 25–30 mm, 1 mm wide at the base, tapering to 0.7 mm at the apex; Sti 4  3 mm, exserted for 2–12 mm, narrow, spreading, papillose, yellowish. T. copanensis Rauh & Rutschmann (J. Bromeliad Soc. 38: 7–9, ill., 1988). Type: Honduras, Copán (Kamm s.n. in BG HEID 68087 [HEID]). — Distr: W Honduras (Copán); saxicolous on rock cliffs and epiphytic on pine trees, elevation not recorded. – Fig. 6. [5] Plant stemless, flowering to 80 cm tall; L numerous; L sheaths 9  6 cm, narrowly ovate, brown-lepidote on both faces, distinct from the lamina; L lamina very narrowly triangular, somewhat succulent, to 35  4 cm at the base, pale green, abaxially densely white-lepidote (trichomes in rows with a green centre), recurved, canaliculate, apex attenuate and pungent; Inf mostly laxly 1 branched or rarely simple, composed of up to 6 spikes, often decurved and then ascending, lax, to 30 cm (including the terminal spike), 20 ( 30) cm wide; peduncle curved, shorter than the leaves, 15 mm ∅; lower peduncular Bra somewhat leaflike, the upper ones like the primary bracts, imbricate, spike axes angled, 10 mm ∅, glabrous, bright carmine-red; primary Bra triangular, the upper ones to 9  3 cm, acute; spikes 20  2 cm, shortly stipitate for 1.5–3 cm, with 1 or 2 sterile bracts at the base that are sharply keeled, sub-pendulous and then ascending or curved, linear-lanceolate, complanate, acute; floral Bra imbricate, >2 as long as the sepals, 47–50  20–23 mm, elliptic, strongly convex, bluntly subcarinate towards the apex and ecarinate below, densely lepidote becoming glabrous at the base, pale green to cinereous-white at the apex, acute or obtuse; Fl with a stout pedicel-like receptacle, contiguous with each other and/or the rachis; Sep 22  5–8.5 mm, oblong-lanceolate, fleshy at the base, with broad hyaline margins and apex, finely veined adaxially, evenly short-connate for

Tillandsia BROMELIACEAE

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Fig. 6 Tillandsia copanensis. (Copyright: E. J. Gouda)

1–2 mm, adaxial ones inconspicuously bluntly carinate, green, glabrous on both faces, obtuse; Pet forming an erect narrow tube clasping the filaments, with revolute apex, tongue-shaped, to 66  7.5 mm, broadest at from the apex, base 4 mm wide, dark violet with hyaline margins and apex, covered part white, blade rounded-emarginate; St in 2 whorls of unequal length, 72 and 78 mm, exserted; Fil exposed part slightly inflated and (sub-) terete, violet; Anth dorsifixed at from the base, 4  1 mm, elliptic in outline, apex rounded, yellowish-brown; Ov 7  3 mm, attenuate from near the base, gradually contracted into the style; Sty slender, to 73 mm, exceeding the stamens; Sti 3 mm, conduplicate-spiral.

3-flowered, rachis lepidote; floral Bra equalling or shorter than the sepals, (8.2–) 11.8  5.4–6.8 mm, ovate, 11- to 12-veined, lepidote, with a very short lamina; Fl subsessile, fragrant; Sep 8.2–10.2  2.4–2.6 mm, lanceolate, 5- to 7-veined, adaxial ones connate for 4–6 mm and slightly carinate, abaxial one connate for 1 mm, green, glabrescent; Pet tongue-shaped, 14.2–15.4  2.7–3.1 mm, brown-violet to black-violet or rarely ochraceous, blade recurved, moderately broadened, tip rounded; St included; Fil 4.4–5.1 mm, white, 1-nerved; Anth basifixed, 1.9–2.3  0.3 mm, golden-yellow; Ov 2–2.3  1.5 mm, narrowly ovoid; Sty slender, 4–4.6 mm, included.

T. cotagaitensis L. Hromadnik (Pl. Syst. Evol. 147: 285–285, 1984). Type: Bolivia, Potosí (Hromadnik & Hromadnik 5093 [WU]). — Distr: SW Bolivia (Potosí); epiphytic on Oreocereus in association with T. caliginosa, 3000 m. [4] Plant caulescent, flowering to 14 cm tall, branched; L densely distichous, divergent to recurving; L sheaths to 1  0.8 cm, amplexicaul, scarious, with broad veinless margins, veined or lustrous, glabrous, becoming greenish; L lamina flexible, to 4  0.2–0.3 cm, densely tomentoselepidote, subulate, apex subulate-acuminate; Inf simple; peduncle 2–3 cm, 1 mm ∅, densely lepidote; spike dense, 2 cm, distichously 2- to

T. crocata (E. Morren) Baker (J. Bot. 25: 214, 1887). Type: Brazil, Rio de Janeiro (Lietze s.n. [K [icono: Morren drawing], GH [photo]]). — Distr: SE & S Brazil (Rio de Janeiro, Paraná, Rio Grande do Sul), NE Bolivia (La Paz), NE Argentina (Entre Ríos), SW Uruguay (Soriano); saxicolous, 875–2650 m.  Phytarrhiza crocata E. Morren (1880). [3] Plant short-stemmed, leafy stem part often shorter than the leaves, flowering 15–35 cm tall, forming dense clusters; stem to 20 cm, L distichous, 10–30 cm, densely white-tomentose-lepidote with asymmetric elongated scales; L sheaths broadly ovate, glabrous except for the upper

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E. J. Gouda

Fig. 7 Tillandsia curvifolia. (Copyright: E. J. Gouda)

outside; L lamina linear, 2–5 mm wide, spreading and recurved, involute-subulate, apex long-attenuate; Inf always simple; peduncle bractless or with a single leaf-like bract at the base, erect or nearly so, 5–15 cm, slightly >1 mm ∅, slender, retrorsetomentose like the leaves; spike densely 2- to 6-flowered, 1–4 cm (excl. petals or capsules), lanceolate, elliptic, acute, with a reduced flower at the apex; floral Bra imbricate,  equalling the sepals, 9–10 ( 20) (including blade like apex)  6.5–7.5 mm, ovate or elliptic, 2–5 as long as the internodes, densely tomentose-lepidote abaxially only, green, tinged purple-red, acuminate, the lowest with a thick blade-like apex; Fl with a 1 mm pedicel-like receptacle or subsessile, distichous, fragrant; Sep 10  4.5 mm, elliptic or sublanceolate, thin, with veinless broad hyaline margins, faintly veined in the centre, unequally short-connate for 1–2 mm, adaxial ones obscurely carinate esp. towards the apex, lower green and then stramineous speckled purple-red, densely appressed-lepidote abaxially on the centre ridge towards the apex, obtuse or acutish; Pet claw narrow with suborbicular blade or spatulate with elliptic blade, tapering into the cuneate claw, 19–20 mm, blade 6–8 mm wide, spreading, obtuse, margins slightly crenulate, bright yellow (paler towards the base), claw 1 mm wide at the base to 3 mm distally; St 7 mm, deeply included, much exceeding the style; Fil slender-attenuate

from near the base, flaccid, straight; Anth basifixed, 1.5 mm, apex broadly obtuse, orangeyellow; Ov 3  2 mm, obovoid (pyriform), abruptly contracted into the style; Sty 1.5 mm; Sti erect, shortly lobed. T. curvifolia (Ehlers & Rauh) Ehlers (Bromelie Sonderheft 6: 126–134, ills., 2009). Type: Mexico, Guanajuato (Ehlers M850201 [WU]). — Distr: C Mexico (Guanajuato); mostly saxicolous, sporadically epiphytic, 1650–3000 m. – Fig. 7.  Tillandsia tortilis ssp. curvifolia Ehlers & Rauh (1990)  Viridantha curvifolia (Ehlers & Rauh) Lopez-Ferrari & Espejo (2009). [1] Plant stemless or short-stemmed, flowering 5–15 cm tall; L spirally arranged, 8–12, 4–16 cm, white-tomentose-lepidote; L sheaths 1.5  1.2 cm, ovate, somewhat succulent, with translucent margins, densely tomentose and glabrous only at the base, conspicuously offset from the lamina; L lamina narrowly triangular, succulent esp. at the base, to 14 cm, 7 mm wide at the base, mostly upwardly secund or erect, silvery-white, subulate and canaliculate, margins involute, apex filiformattenuate, adaxially densely appressed-lepidote, abaxially tomentose-lepidote, trichomes eccentric; Inf simple or very rarely with a smaller lateral spike, erect, shorter than or slightly exceeding the leaves; peduncle concealed within the leafy rosette or extended and hidden by the inner leaves

Tillandsia BROMELIACEAE

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Fig. 8 Tillandsia diaguitensis. (Copyright: E. J. Gouda)

and bracts, to 5 ( 14) cm, 2 mm ∅, glabrous or sparsely lepidote; lowest peduncular Bra leaflike, upper ones like the floral bracts, erect, few (3–5 only), imbricate, 3–5 as long as the internodes, with setaceous lamina to 7 cm; spikes with a sterile bract at the base, 2–3  to 1 cm, (narrowly) ellipsoid, terete, (1- to) 2- to 5-flowered, rachis angled, stout, densely lepidote, hidden; floral Bra erect, densely imbricate, slightly exceeding the sepals, 10–20  10 mm, ovate to lanceolate, thin, margins hyaline, finely veined or distinctly veined towards the apex, 2–3 as long as the internodes, glabrous or sparsely lepidote, roseate or sometimes green, acuminate; Fl polystichous or subdistichous; Sep to 15  4 mm, lanceolateoblong, membranous, with hyaline margins, free, adaxial ones distinctly carinate, glabrescent or sparsely lepidote at the apex; Pet for most of the length tubular-erect, tongue-shaped, 20–27  4 mm, green with white base, blade apex recurved, rounded; St all of equal length, deeply included; Fil to 15 mm, filiform, flat, white; Anth basifixed, 3–5 mm, sagittate, pale brown; Ov 4  2 mm; Sty slender, 6 mm; Sti lobes short, erect or somewhat divergent at maturity. T. diaguitensis A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 36: 55, t. 10, 1929). Type: Argentina, Jujuy (Castillon 7224 [BA]). — Distr: N Argentina (Jujuy, Formosa,

Salta, Tucumán), C Paraguay; epiphytic, 90– 2050 m. – Fig. 8. [2] Plant long caulescent, slender; stem 60 cm, simple or few-branched, 5 mm ∅; L laxly polystichous along the stem, 8–10 cm, densely cinereous-lepidote (furfuraceous); L sheaths 1.3 cm wide, elliptic, amplexicaul, imbricate, making the stem appear 7–10 mm thick; L lamina narrowly triangular, 0.6 cm wide, erect to recurved, strongly veined at least when dry, canaliculate, apex attenuate; Inf always simple; peduncle completely covered by the bracts, conspicuous, to 8 cm, 3 mm ∅; peduncular Bra imbricate, elliptic, thin, strongly veined, lepidote towards the apex, stramineous, acute; spike dense, 4–9 cm, 1.3–1.4 cm wide, lanceolate, 3- to 6-flowered, acute, rachis slender, nearly straight, glabrous, sulcate; floral Bra imbricate to divergent at anthesis, much exceeding the sepals, 40–50  10 mm, lanceolate, chartaceous, margins scarious and veinless, finely to strongly veined when dry and waxy, ecarinate, 2–3 as long as the internodes, glabrous, roseate or stramineous, narrowly acute; Fl subsessile, fragrant; Sep 26 ( 32)  4.5–6 mm (abaxial one smaller), lanceolate-oblong, with membranous margins, veined, free, adaxial ones carinate, glabrous, acute; Pet spatulate, 60–70  13 mm, margins minutely denticulate or slightly crenulate, white or bluish, claw linear, forming a tube well beyond the sepals, blade obovate,

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rounded; St all of equal length, 38–51 mm, included and visible in the throat; Anth subbasifixed, 5–8 mm, linear; Ov 5–7 mm, tapering into the style; Sty slender, 37 mm, exceeding the stamens; Sti 3 mm, spreading, weakly conduplicate. T. dorisdaltoniae Ibisch & al. (Revista Soc. Boliv. Bot. 4(1): 45, 2003). Type: Bolivia, Cochabamba (Vargas 6355 [WU]). — Distr: C Bolivia (Cochabamba); epiphyte, 950–1050 m. [2] Plant short-stemmed, flowering to 20 cm tall or more, forming dense clusters; stem richly branched at the base; L forming a dense rosette, numerous, erect to divergent, exceeding or about equalling the inflorescence; L sheaths to 1.5  1 cm, abaxially lepidote, conspicuously offset from the lamina; L lamina rigid, to 10 cm, 0.6–0.7 cm wide at the base, erect or divergent, subulate, apex attenuate, acute, intensely green, sparsely lepidote, adaxially glabrescent towards the apex; Inf simple; peduncle erect, much shorter than the leaves, to 3 cm, slender, green; peduncular Bra imbricate, upper ones reduced to ovate sheaths, lepidote to glabrous on the sheath, whitish, the lower ones with leaf-like lamina; spike erect, 4–5 cm, complanate, densely 8-flowered, rachis completely covered by the bracts at anthesis but slightly exposed when dry as the margins Fig. 9 Tillandsia dorotheae. (Copyright: E. J. Gouda)

E. J. Gouda

of the bracts bend outwards, internodes 3.5 mm; floral Bra erect, distichous, densely imbricate, exceeding the sepals, 12–14 ( 20)  5–8 mm (2.5–3 mm wide in lateral view), ovate, faintly veined towards the apex esp. when dry, ecarinate, covering > of the flowers, glabrous, bright red, acute; Fl subsessile; Sep 11  3.5 mm, membranous, completely free, subadaxial ones alatecarinate, whitish, glabrous, subobtuse; Pet suberect, tongue-shaped, 16  3 mm, white, blade slightly divergent, apex rounded; St all of equal length, 12 mm, included; Fil 8.5 mm, flat, slightly plicate in the upper ; Anth subbasifixed, 4 mm; Ov 2  1.2 mm, ovoid; Sty slender, 9 mm; Sti 0.5 mm, erect, papillose. T. dorotheae Rauh pro sp. (Trop. subtrop. Pfl.-welt 60: 59, ill., 1987). Type: Argentina, Salta (BG Heidelberg 31306 [HEID, MO, WU]). — Distr: NE Argentina (Salta); saxicolous in bare lands, in crevices and almost inaccessible crags and bluffs, altitude not recorded. – Fig. 9. [2] Plant  caulescent, forming dense clusters; stem 5–10 cm, laxly branched at the base; L densely spirally arranged but sometimes showing a tendency to distichous, erect to spreading; L sheaths distinct, to 1.5  1 cm, elongate, with membranous margins, base glabrous, upper densely cinereous-lepidote, whitish or

Tillandsia BROMELIACEAE

occasionally pinkish-violet; L lamina very narrowly triangular, rigid, 8–10  0.3 cm, cinereous-green, densely lepidote, veined and distinctly rugose at the base, adaxially deeply and narrowly canaliculate but not keeled, apex abruptly obtuse; Inf simple, decurved, shorter than the leaves; peduncle mostly decurved, sometimes erect, short and concealed by the leaves, 4–6 cm, 2 mm ∅, terete, green; peduncular Bra few (3–4), the lower leaf-like, the upper 1 or 2 with shorter lamina, densely imbricate; spike decurved, sublax or subdensely flowering, to 3  0.8 cm, (1- to) 2(to 4–) flowered, with a reduced flower at the apex, rachis flat on one side, slender, flexuous, glabrous; floral Bra erect, slightly shorter than the sepals, 20  5–8 mm, lanceolate, coriaceous, margins hyaline, shiny at the base, strongly veined, carinate towards the apex, lower part glabrous, scatteredly white-lepidote upwards, green to wine-red, acuminate or fleshy-apiculate; Fl subsessile, distichous; Sep 19–22  4 mm, lanceolate, thin, even, free, adaxial ones carinate, green and flushed red, acuminate or abruptly acute; Pet spatulate, 33–35  6–7 mm (base 2.5 mm wide) reddish or dark pink, blade obscure, recurved and often twisting, acuminate; St (9–) 13–17 mm, deeply included, shorter than the style; Fil ribbon-like, not plicate; Anth dorsifixed near the base, 5–6 mm, yellow; Ov 5  2.5 mm, cylindrical, abruptly contracted into the style; Sty slender, 15 mm; Sti erect, weakly conduplicate. Considered to be the hybrid T. argentina  T. albertiana. T. duratii Visiani (Nuovi Saggi Imp. Regia. Accad. Sci. Padova 5: 271, t. 29, 1840). Type: PAD?; [icono]: l.c. t. 29. — Distr: Brazil, N Peru, Bolivia, Argentina, Paraguay, NE & NW Uruguay; epiphyte or saxicolous, 80–2750 m.  Phytarrhiza duratii (Visiani) Visiani (1854)  Anoplophytum duratii (Visiani) Beer (1856); incl. Tillandsia floribunda Durat ex Visiani (1840) (nom. illeg., ICN Art. 53.1); incl. Tillandsia circinalis Grisebach (1874)  Phytarrhiza circinalis (Grisebach) E. Morren ex Baker (1889); incl. Tillandsia gigantea Ruchinger (1876) (nom. inval., ICN Art. 32.1c); incl. Tillandsia decomposita Baker (1889); incl.

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Tillandsia weddellii Baker (1889); incl. Tillandsia revoluta Burbidge ex Baker (1889) (nom. inval., ICN Art. 32.1c); incl. Tillandsia tomentosa N. E. Brown (1894); incl. Tillandsia confusa Hassler (1919)  Tillandsia duratii var. confusa (Hassler) L. B. Smith (1968); incl. Tillandsia confusa var. minor Hassler (1919); incl. Tillandsia confusa var. saxatilis Hassler (1919)  Tillandsia duratii var. saxatilis (Hassler) L. B. Smith (1968). [3] Plant caulescent, flowering 20–100 cm tall or more; stem to 30 cm, simple, stout, curved; L laxly polystichous, densely cinereously lepidote with subappressed coarse scales; L sheaths 2 cm, broadly ovate, distinct from the lamina; L lamina very narrowly triangular, rigid, thick, 15–20 ( 40)  1–2 cm, involute-subulate to often contorted towards the apex that is then twining around tree branchlets, apex pungent; Inf 1 branched or 2 branched in the lower part, 6–60 cm; peduncle erect, elongate, stout, glabrous; lowest or all peduncular Bra almost leaf-like, densely imbricate, elliptic, densely cinereouslepidote, the upper apiculate; primary Bra erect, like the peduncular bracts, enveloping the bracteate sterile bases of the axillary branches; Br erect or spreading or recurving, lower Inf spikes longstipitate for up to 10 cm, spikes complanate, 4.5 cm long, 1.7 cm wide, with several sterile bracts at the base, lanceolate or linear; floral Bra erect, slightly longer or shorter than the sepals, to 17  8 mm, ovate to oblong, even, ecarinate, subdensely to densely lepidote, greenish but soon stramineous, obtuse; Fl subsessile, fragrant; Sep to 14 mm, ovate, elliptic, coriaceous at the base, with membranous margins and apex, even, evenly or unequally short-connate for 2 mm, ecarinate, green or brown, glabrous, subobtuse; Pet 25–33 mm, claw narrow with suborbicular blade, margins slightly crenulate, usually (purple-) blue-speckled, claw linear, white, blade subrhombic to orbicular, spreading; St all of nearly equal length, 11 mm, deeply included but exceeding the style; Fil thin but not flaccid, not plicate; Anth basifixed, 2.5 mm, green; Ov 7.5 mm, bottle-shaped, ribbed; Sty short; Sti shortly lobed. This is a very varible species growing in a range of habitats. It can grow on rock faces mixed with T. streptocarpa, looking very different in habit,

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Fig. 10 Tillandsia ehlersiana. (Copyright: E. J. Gouda)

but in other habitats intermediate forms occur. Intermediate forms between these 2 species, but also with T. paleacea, are common and are difficult to place under one of the 3 species, mainly because the flowers and spikes are not very different if at all from each other. The infraspecific taxa recognized by some authors are not accepted here. T. ehlersiana Rauh (J. Bromeliad Soc. 34(4): 166–169, ills., 1984). Type: Mexico, Chiapas (Ehlers 83/404 [HEID]). — Distr: SE Mexico (Chiapas); on steep granitic rocks in deciduous forest, 700 m. – Fig. 10. [5] Plant stemless, flowering to 20 cm tall, forming clumps; L many, exceeding the inflorescence, silver-grey; L sheaths forming a pseudobulb to 8  10 cm, individual sheaths to 7  6–7 cm, suborbicular, strongly inflated, densely furfuraceously lepidote, adaxially light leatherbrown, abaxially whitish, distinct from the lamina; L lamina narrowly triangular to linear, with asymmetric trichomes at the margins, above the sheath to 2.5 cm wide, both faces densely whitelepidote, upper part often recurved, strongly involute-subulate and canaliculate; Inf 1 branched, composed of 4 spikes, ovoid in outline, much shorter than the leaves; peduncle short and concealed by the leaves, to 7 cm, 1 cm ∅, terete, appressed-lepidote; peduncular Bra leaf-

like, few (3), similar to the upper rosette leaves, silver-greyish; axes compressed, 7 cm, straight; primary Bra erect or divergent with the branches, the basal ones with broadly ovate sheaths, somewhat leaf-like, 2  1 cm, exceeding the spikes or inflorescence and the upper shorter than the spikes and acute, densely furfuraceously lepidote; spikes shortly stipitate for 0.5 cm, erect or divergent, 3  1.8 cm, complanate, 2- to 3-flowered; floral Bra distichous, imbricate, much exceeding the sepals, 19–22 ( 28)  10–13 mm, triangular-ovate, membranous, margins even, carinate with a thick midvein, concealing the rachis, cinereously lepidote, brownish-red or roseate, acute or obscurely apiculate; Fl subsessile, 30–65 mm; Sep 13–19  6–7 mm, (narrowly) elliptic, membranous, even, free or shortly connate for up to 3 mm, adaxial ones carinate (sharply so when dry), greenish-red, (very) sparsely lepidote especially on the keel, acute or broadly obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, linearly tongue-shaped, (33–) 42–53  8 mm, upper blue-violet, white towards the base, obtuse; St in 2 whorls of unequal length, 57–60 mm, exserted, shorter than or as long as the style; Fil subterete towards the apex, straight, upper violet; Anth dorsifixed at from the base, 2.5 mm, blackish; Ov 5 mm, subprismatic, gradually contracted into the slender style.

Tillandsia BROMELIACEAE

T. erecta Gillies ex Baker (J. Bot. 16: 239, 1878). Type: Argentina, Mendoza (Gillies s.n. [K]). — Distr: C Bolivia (Cochabamba), Argentina (Mendoza, La Rioja); saxicolous and epiphytic, 1300–2610 m. Incl. Tillandsia rigida Gillies ex Baker (1878) (nom. inval., ICN Art. 32.1c)  Tillandsia erecta var. rigida (Gillies ex Baker) Baker (1889). [4] Plant to 11 cm tall; stem 3–6 cm, simple or few-branched; L laxly polystichous, to 5 cm, densely cinereous-lepidote; L sheaths 0.5–1 cm, broadly ovate, glabrous; L lamina very narrowly triangular, rigid and succulent, 0.4 cm wide at the base, erect to divergent, veined or obviously grooved, involute-subulate, angled towards the acuminate apex; Inf 1-flowered with the rachis extended behind the flower; peduncle to 4.5 cm, elongating in fruit, slender, strongly sulcate or strongly furrowed when dry, glabrous; peduncular Bra 0–2, 1.5 cm, elliptic, glabrous; floral Bra equalling or shorter than the sepals, (8–) 12–13.5  5.5–6 mm, triangular-ovate to ovate-lanceolate, not tapering at the base, distinctly 6- to 10-veined, glabrous or upper part sparsely lepidote, stramineous or brown-reddish in the lower part, acute or obtuse; Fl subsessile, erect, weakly fragrant; Sep 10–11.5  3–4 mm, narrowly elliptic or narrowly lanceolate-triangular, 5-veined, evenly short-connate or adaxial ones somewhat more connate, glabrous or sparsely lepidote, acute or Fig. 11 Tillandsia erici. (Copyright: E. J. Gouda)

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obtuse; Pet broadly tongue-shaped, 9–12.5  2.5–3 mm, dirty greenish-yellow or greenishpunctate, blade narrowly elliptic, rounded; St deeply included, exceeding the style; Fil 3–5  0.2 mm, whitish; Anth (sub-) basifixed, 2–2.5  0.5 mm; Ov 2 mm, obconical or subcylindrical, abruptly contracted into the style; Sty stout, only as long as the ovary; Sti capitellate, flat. T. erici Ehlers (Bromelie 1998(1): 18–21, ills., 1998). Type: Bolivia, Tarija (Haugg 11,247 [WU]). — Distr: S Bolivia (Tarija); epiphyte, 1800–2200 m. – Fig. 11. [2] Plant stemless, flowering 12–22 cm tall, in small clusters or solitary; L 12–20, forming an erect rosette to 25 cm ∅, almost erect, fleshy to coriaceous, strongly veined (prominently when dry), finely appressed-lepidote, reddish-brown; L sheaths 2.5–4  1.4–2 cm at the base, triangularovate, inconspicuously offset from the lamina; L lamina very narrowly triangular, 6–18 cm (outer ones reduced and only 10–12 cm), 0.8–1 cm wide at the base, very finely appressed-lepidote (adaxially less), margins strongly incurved, apex acute and pungent; Inf simple, shorter than the leaves; peduncle completely covered by the bracts, decurved, only 3–4 cm, 4 mm ∅; lowest peduncular Bra leaf-like, the upper like the floral bracts, densely imbricate, 3.5 cm, coriaceous, lower ones

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caudate, the upper acute; spike with 1–2 sterile bracts at the base, often  decumbent, 4–7 cm, 0.8–2 (after flowering to 3) cm wide, lanceolate or elliptic, complanate, (1–) 3- (to 7–) flowered, acuminate; rachis 4-angled, almost straight, glabrous, green; floral Bra imbricate, exceeding the sepals, 18–35  8–13 mm, narrowly ovate, rigid and coriaceous, margins broadly hyaline, veined (strongly when dry), shiny, ecarinate, concealing the rachis, glabrous, stramineous or green tinged purple, acute; Fl sessile or with a short bicarinate pedicel-like receptacle, distichous, not fragrant; Sep 18–26  6–9 mm, elliptic or oblong, membranous, with broad hyaline margins, slightly finely veined, free or shortly connate for 1–2 mm (abaxial one less if at all), adaxial ones bluntly carinate, pale green or brown tinged purplish-red, glabrous, acute; Pet spatulate, throat opening, 40–60  9–12 mm, claw 3 mm wide, margins crenateserrate and slightly undulate, bright orangeyellow, blade spreading and recurved; St included or  equalling the petal claw; Fil 25 mm, ribbonlike, filiform, thin, straight, yellowish-white, upwards yellow; Anth basifixed, 4–5  0.4 mm, linear, yellow; Ov 5  2.2 mm, slenderly prismatic; Sty slender, 30 mm; Sti lobes 14 mm, spreading or recurved, apex somewhat papillose for 4 mm, yellow. T. ferrisiana L. B. Smith (Bromeliad Soc. Bull. 10: 92, ills., 1960). Type: Mexico, Sinaloa (Ferris & Mexia 5121-A [DS]). — Distr: Mexico (Sinaloa, Baja California); epiphytic in dry woods and scrub, 630 m. [5] Plant forming dense clusters or few-branched; stem 2–6 ( 10) cm; L laxly spirally arranged, much exceeding the inflorescence, to 15 cm, densely cinereous-lepidote; L sheath 1 cm, suborbicular to narrowly ovate, densely cinereouslepidote but glabrous where overlapping; L lamina very narrowly triangular to linear, involutesubulate, apex filiform-attenuate; Inf simple, erect, to 3 cm; peduncle short and concealed by the leaves; peduncular Bra somewhat leaf-like, densely imbricate; spike dense, terete, (1- to) 2- to 3-flowered; floral Bra exceeding the sepals, 20 mm, ovate, thin, veined towards the apex,

E. J. Gouda

ecarinate, densely cinereously-lepidote, apiculate; Fl subsessile, distichous, 50 mm; Sep 12–20  6 mm, elliptic to lanceolate, membranous, smooth, free, adaxial ones carinate, appressed-lepidote or becoming glabrous, obtuse; Pet erect with recurving apex margins, 50 mm, bicoloured, the base red-violet to purple, the upper 20 mm yellowish or cream-coloured (Rauh 1979); St and Sty long exserted. T. fresnilloensis W. Weber & Ehlers (Feddes Repert. 94(9–10): 609–611, fig. 9, 1983). Type: Mexico, Zacatecas (Ehlers s.n. [HAL (Herb. Weber)]). — Distr: C Mexico (Zacatecas); ecology and altitude not recorded. – Fig. 12. [5] Plant flowering to 22 cm tall; stem richly branched at the base; L forming an erect rosette, numerous, white-lepidote; L sheaths to 2.5  2 cm, triangular to ovate, strongly wrinkled when dry, indistinctly merging into the lamina; L lamina narrowly triangular, to 9 cm, in the middle to 1 cm wide, erect and secund, strongly grooved, apex acuminate; Inf simple; peduncle to 12 cm with internodes 2–2.5 cm, 2 mm ∅, terete, smooth, with weak purplish tint; peduncular Bra erect, just exceeding the internodes, lanceolate, veined, sparsely appressed-lepidote to almost glabrous, stramineous, finely apiculate; spike 3- to 6-flowered, rachis strongly geniculate; floral Bra shorter than the sepals, 20–23  10–12 mm, ovate or lanceolate, margins broadly hyaline, faintly veined or even, ecarinate, glabrous on both faces, green and tinged red at the base, rounded and apiculate; Fl with a stout pedicel-like receptacle, spreading out, 40 mm; Sep to 25  9 mm, narrowly obovate or lanceolate, with broad hyaline margins, finely veined adaxially, connate for 2–3 mm, ecarinate but with very fleshy midvein, green, flushed red at the apex, glabrous on both faces, rounded or obscurely apiculate; Pet tongue-shaped, 42  9 mm (3 mm at the base), greenish-white or green, acute; St in 2 unequal series of 46 and 51 mm, exserted but shorter than the style; Fil fleshy throughout, slender and flattened at the base and dilated-terete at the distal end, green with paler base; Anth dorsifixed near the middle, 2.5 mm, apex and base both truncate, green; Ov

Tillandsia BROMELIACEAE

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Fig. 12 Tillandsia fresnilloensis. (Copyright: E. J. Gouda)

4 mm, ovoid, gradually contracted into the style; Sty slender, 51 mm, twisting; Sti shortly conduplicatespiral and slightly dilated, green. T. funebris A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 37: 502, 1933). Type: Argentina, Tucumán (Schreiter 1689 [BA, B, GH, LIL]). — Distr: Venezuela, Bolivia, N to E & C Argentina, Paraguay; epiphytic in dry forest, 200–2100 m. [4] Plant small, flowering rarely over 10 cm tall, forming dense clusters; stem 2–5 cm, few- to richly branched at the base but otherwise simple; L laxly polystichous, to 5 cm; L sheaths making the stem appear stout, 0.5–0.8 cm, suborbicular to reniform, densely imbricate, subcoriaceous, with broad veinless margins, many-veined, lepidote and glabrous below, distinct from the lamina; L lamina narrowly triangular, densely cinereouslepidote with small nearly symmetrical subappressed scales, divergent to reflexed, often distinctly keeled below,  contorted, triangular-subulate, strongly angled but not at all sulcate-striate, with a furrow above that does not reach the attenuate apex; Inf simple, 1- or 2-flowered, glabrous; peduncle distinct, nearly naked, erect or curved, much

exceeding the leaves, to 6 cm, 0.5 mm ∅, slender, glabrous; peduncular Bra 1–2 just below the flowers, 1.3 ( 1.7) cm, oblong-lanceolate, subcoriaceous, even or many-veined, glabrous, acute; spike rachis like the peduncle and nearly as thick, swollen at the nodes; floral Bra slightly shorter or longer than the sepals, 9–15  4.5–5 mm, like the upper peduncular bract but more ovate or elliptic, progressively smaller upwards, even, ecarinate, subacute; Fl subsessile, erect; Sep 9.5–10  3 mm, narrowly elliptic, even or finely veined (many-veined), evenly shortly connate for 2 as long as the sepals, 35–45  15–20 mm, narrowly ovate, faintly veined, ecarinate, mostly concealing the rachis, 2–3 as long as the internodes, densely lepidote except towards the base, pink or roseate, acute; Fl with a short pedicel-like receptacle, not fragrant; Sep 22–24  4–5 mm, lanceolate to narrowly oblong, subcoriaceous, with broad hyaline margins, veined, connate for 1 mm, fleshy, adaxial ones carinate at the base (sharply so when dry), green at the base but centre part red, glabrous, acutish; Pet erect, forming a tube, tongue-shaped, 50–70  9–10 mm, basally 4 mm wide and white, upper blue-violet, blade tip margins recurving, rounded; St 70–75 mm, exserted; Fil flat, thicker towards the apex but still complanate, thin and flaccid at the base, sometimes slightly twisted, white at the base and violet upwards; Anth dorsifixed at from the base, 3–4  1 mm, pale brown; Ov 7  2.5 mm, cone-shaped with flat sides, contracted into the

T. kuehhasii W. Till (Bromelie 1995(2): 33, ill., 1995). Type: Bolivia, Chuquisaca/Potosí (Kühhas & Kirschnek s.n. [WU]). — Distr: S Bolivia (Chuquisaca, Potosí); saxicolous, 3600 m. [4] Plant forming dense clusters; stem to 25 cm but usually only terminal 10 cm with living leaves, branched at several nodes along the stem; L densely distichous, distally straight, flexible, white; L sheaths 1.5 cm, broadly ovate, amplexicaul, densely lepidote, merging gradually into the lamina; L lamina subulate, (2.5–) 5–7.5 cm, 2 mm wide at the base, densely cinereouslepidote, suberect or irregularly spreading, upper face with a furrow; Inf 1-flowered, erect, 5 cm, shorter than to as long as the leaves; peduncle with a single bract at the base, partly visible, very short or slightly elongate, 2–3.5 cm, elongating after flowering, 1 mm ∅, grooved when dry, sparsely lepidote to almost glabrous towards the base; peduncular Bra involute around the peduncle with overlapping margins, cinereous-lepidote, with leaf-like lamina; floral Bra shorter than the sepals, 13  10 mm, broadly ovate to ovate-oblong, distinctly 10- to 12-veined, abaxially densely lepidote only, cinereous-green or tinged brown towards the margins, rounded and apiculate or caudate with a 2 mm long apiculus; Sep (10–) 13.5–16  3.5 mm, lanceolate, veined, adaxial ones connate for 5–7 mm, adaxial ones bluntly if at all carinate or narrowly convex, green or tinged brown-red at the apex, abaxially sparsely to subdensely appressed-lepidote towards the apex, apiculate or obtuse; Pet tongue-shaped, 15–25  4 mm, yellow tinged brown to chestnut-brown with yellow claw, apical part recurved to recoiled; St 10 mm, deeply included, much exceeding the pistil; Fil flat, attenuate from near the base, straight; Anth basifixed, slightly 4 cm, most of it leafless, simple or few-branched; L densely rosulate, mostly 2–3 ( 4) cm, densely cinereous-lepidote; L sheath 0.3 cm, only slightly broader than the lamina, subcoriaceous, with broad hyaline margins, veined, glabrous, whitish; L lamina very narrowly triangular, rigid, 0.3–0.5 cm wide, densely cinereouslepidote with  spreading scales, erect to suberect or arching-secund, subulate, apex long-attenuate but not filiform; Inf simple or very rarely forked; peduncle (nearly) completely covered by its bracts, curved, mostly much exceeding the leaves, to 10 cm but often much less, 1 mm ∅, straight, lepidote near the nodes, dark green; peduncular Bra like the floral bracts, many,  equalling the internodes, elliptic or obovate, tubularly amplexicaul, chartaceous, veined, densely lepidote, acutish; spikes to 4 cm, linear, sinuous, very much like those of Lolium, to 16-flowered but often with very few flowers or reduced to a single flower at the apex, rachis excavated next to the flowers,  flexuous, lepidote with large scales; floral Bra not at all imbricate, shorter than the sepals, 3–6 ( 8)  3.5–4 mm, (broadly) ovate, closely enveloping the flowers, thin, margins hyaline, veined, ecarinate, small and exposing most of the rachis, just exceeding the internodes, only abaxially subdensely lepidote with large green centered scales, cinereous-green, acute or apiculate; Fl with a short pedicel-like receptacle, distichous, erect, appressed to the rachis except the upper part of the sepals and petals caused by the flexuous rachis; Sep 5.5–6 ( 9)  3.5–4 mm, (oblong) lanceolate, faintly 7-veined, evenly short-connate for >1 mm, green or spotted brown-red, glabrous, acutish or obtuse; Pet 10 mm, claw sublinear, yellow, blade narrow, spreading or recurved, obtuse or narrowly rounded;

E. J. Gouda

St 4 mm, deeply included, exceeding the style with much of the anthers; Fil filamentous, thin; Anth basifixed, 1.5 mm, yellow; Ov shortly cylindrical, obovoid or pyriform, abruptly contracted into the style; Sty stout, shorter than the ovary. T. lydiae Ehlers (Bromelie 2000(2): 55–59, ills., 2000). Type: Mexico, Guerrero (Köhres & Köhres s.n. [MEXU, WU]). — Distr: S Mexico (Guerrero); epiphytic, 350–1000 m. [5] Plant flowering 20–30 cm tall; Ros 3–4 cm ∅; L 5–8, forming a bulbous rosette, 25–30 cm, the outer greatly reduced, very thick and succulent, stiffly leathery, margins very strongly grooved, densely appressed-lepidote, silver-grey but lustrous; L sheaths erect, 6–10  3–4 cm, narrowly ovate, firmly pressed together, inflated, appressed-lepidote, glabrous towards the base, inside brown, outside dark green, conspicuously offset from the lamina; L lamina narrowly triangular, 10–20 cm, 2–2.5 cm wide at the base, red-brown, with striking longitudinal brown-red stripes, almost erect or slightly divergent, in dry conditions appearing grooved or strongly veined, subulate, margins involute, apex acuminate tapering to a sharp tip; Inf 1 branched or simple, composed of (1–) 2–5 spikes; peduncle with a single bract similar to the inner leaves, short and concealed by the leaves, 4 mm ∅, sparsely lepidote; primary Bra with a basal flat part like the floral bracts, 1–1.8 cm, but with an acute short lamina, densely grey-lepidote; spikes short-stipitate, stipe 2–4 cm, with 2–3 (bi-) carinate sterile bracts at the base, spreading at an angle of 45–90 , 5–12 cm, 8–10 ( 12) mm wide, lanceolate, complanate, 3- to 11-flowered, rachis with internodes 8–10 mm; floral Bra 3–5 mm shorter than the sepals, 13–15  6–8 mm, ovate, veined, towards the apex keeled, densely appressed-lepidote, pink, acute; Fl distichous, contiguous with each other and/or the rachis; Sep 20–22  6–7 mm, elliptic, with membranous margins, subfree, ecarinate or adaxial ones slightly carinate at the base, raspberry-red except the base, glabrous, acute; Pet erect, forming a tube, spatulate, 35  6 mm (base 4 mm wide), pale violet with white base, blade tip margins divergent; St in 2 whorls of unequal length, exserted; Fil

Tillandsia BROMELIACEAE

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Fig. 21 Tillandsia mauryana. (Copyright: E. J. Gouda)

34–36 mm, broadened towards the apex and 0.8 mm wide, the upper portion pale lilac tapering to the white base; Anth dorsifixed at from the base, 3  0.75 mm (5 mm before dehiscence), linear or somewhat sagittate, dark yellow; Ov 4  2 mm, cone-shaped; Sty slender, to 35 mm; Sti 1.5  1–2 mm, narrow, small, erect, lobes green. T. mauryana L. B. Smith (Contr. Gray Herb. 117: 31, t. 2, figs. 32–33, 1937). Type: Mexico, Hidalgo (Maury 5747 [GH, F]). — Distr: C Mexico (Hidalgo); saxicolous, 1200–2000 m. – Fig. 21.  Viridantha mauryana (L. B. Smith) Espejo (2002). [1] Pants often vertically on steep rockwalls, stemless or short-stemmed, flowering 5–15 cm tall; Ros 4–11  5–14 cm, usually in groups; L densely rosulate, to 30, 5–10 ( 12) cm, thickfleshy, densely cinereous-lepidote, silver-grey; L sheath 1.5–2  0.8–1.2 cm at the base, triangular, grey to brown-lepidote, glabrous for the lower 2 mm, both faces light brown, inconspicuously offset from the lamina; L lamina narrowly triangular, 4–10  0.5–1 cm wide at the base, densely cinereous-lepidote, spreading or reflexed, flat, apex attenuate; Inf composed of 3–9 spikes, longer or shorter than the leaves; peduncle short and concealed by the leaves, 1–2 ( 3.5) cm, with a

collar of bladeless acute bracts below the inflorescence; lowest peduncular Bra leaf-like, the upper like the primary bracts, erect or bent outward, densely imbricate, (2–) 5–8 cm, narrowtriangular; primary Bra like the floral bracts, lanceolate, 2.5–3  1–1.5 cm, shorter than the spikes, membranous, acuminate, densely lepidote; spikes with 1 or 2 sterile bracts at the base, erect or divergent, to 30  10 ( 15) mm, complanate and with slightly convex sides, 3- to 8-flowered, rachis hardly visible at anthesis; floral Bra densely imbricate, slightly exceeding the sepals, 13–20 ( 25)  10–15 mm, (triangular-) ovate, thin, faintly veined, finely to sharply carinate esp. at the apex, adaxially glabrous, abaxially cinereously lepidote with large grey spreading scales, roseate, acuminate; Fl subsessile, distichous, densely arranged; Sep 12–15 ( 17)  4–5 mm, narrowly ovate, membranous, evenly short-connate for 1 mm or subfree, alate, adaxial ones carinate, sparsely to subdensely lepidote, acute; Pet almost erect, linear, 14–24  3–4 mm (base 2–2.5 mm wide), dark emerald-green and yellow-green at the base, blade suberect or slightly divergent, rounded or obtuse; St all of equal length, 18–20 mm, included; Fil 12–14 mm, ribbon-like, thin and flaccid, white; Anth subbasifixed, 2.5–3  0.4 mm, linear, base bilobed, apex apiculate, yellow-olive to light brown; Ov 3  2 mm, ovoid, tapering into the

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Fig. 22 Tillandsia mitlaensis. (Copyright: E. J. Gouda)

style; Sty 11 mm, reaching or surpassing the anthers; Sti erect, slightly wider than the style, linear, lobes green. T. mitlaensis W. Weber & Ehlers (Feddes Repert. 94: 617–618, ill., 1983). Type: Mexico, Oaxaca (Ehlers s.n. [Herb. Weber 283]). — Distr: S Mexico (Oaxaca); saxicolous, 1800 m. – Fig. 22. [5] Plant flowering 16 cm tall; L suberect, the outer ones reduced, densely coarsely whitetomentose-lepidote; L sheath to 3  2 cm, ovate, indistinctly merging into the lamina; L lamina narrowly triangular,  to strongly secund, finely veined, 7–9 cm, stout, canaliculate, apex subulate, subobtuse or acute, not pungent; Inf simple; peduncle suberect, short and concealed by to  equalling the leaves, stout; peduncular Bra with short somewhat leaf-like blades, erect or slightly bent upwards, densely imbricate, enveloping the peduncle, densely lepidote; spike 7–12  1.6–1.8 cm, lanceolate, subterete to distinctly complanate, 5-flowered, attenuately acute; floral Bra suberect, densely imbricate, much exceeding the sepals, 32–35 ( 44)  16 mm, ovate-lanceolate, submembranous, margins hyaline, strongly veined, ecarinate, densely concealing the rachis, on both faces (sub-) densely lepidote, reddish, obtuse or obscurely acuminate; Fl with a short pedicel-like receptacle, distichous; Sep 23–24  5 mm, lanceolate,

membranous, with broad hyaline margins, veined, adaxial ones connate for 4–5 mm, abaxial one subfree, adaxial ones distinctly bluntly carinate, whitish-green, glabrous or abaxially sparsely lepidote with large trichomes, acute; Pet forming an erect narrow tube clasping the filaments at the apex, 50  7.5 mm, pale to dark violet, whitish below, blade tip margins recurved, roundedemarginate; St in 2 whorls of unequal length,  exserted; Fil 50–54 mm, slender and flattened at the base and dilated-terete at the distal end, white below, exserted part violet; Anth dorsifixed at from the base, 2 mm, elliptic, brownish-green; Ov 5 mm, ovoid, contracted into the style; Sty 55 mm, slender, slightly dilated upwards; Sti conduplicatespiral, unequal, 2.5 mm, whitish-green. T. mollis H. Hromadnik & W. Till (Pl. Syst. Evol. 142: 123–128, ills., 1983). Type: Bolivia, Tarija (Hromadnik 9088 [WU, B, NY, W, Herb. W. Till]). — Distr: S Bolivia (Tarija); saxicolous on S-facing cliffs, 1770–2700 m. [4] Plant flowering (9–) 16–24 ( 26) cm tall and 3.5 – 4 cm wide, forming dense clusters 40–50 cm ∅; stem simple or few-branched, internodes (0.7–) 0.8–1 cm; L distichous, spreading, (2–) 3–5 ( 6) cm, soft, flexible; L sheath amplexicaul, completely covering the stem and making it firm, 0.9–1.1  0.8–1.1 cm, broadly obovate, with translucent margins near the

Tillandsia BROMELIACEAE

middle, margins connate to each other in the lower , 9- to 10-veined, adaxially glabrous, abaxially densely lepidote with eccentric spreading scales, pale stramineous; L lamina subulate-terete, 1.9–4  0.3–0.5 cm wide at the base, silvery-white, densely lepidote, trichomes eccentric and spreading, apex subacuminate, abruptly flattened and obtuse; Inf 1-flowered, rachis thread-like extended above the flower but hidden between the bract; peduncle nearly none or very short, to 0.8 cm, soon laterally displaced after anthesis, densely lepidote; peduncular Bra 1, placed approximately beneath the floral bract, 1.2–1.3 cm, densely lepidote, short-caudate; floral Bra  equalling the sepals, 6–9  2–5.5 mm, suborbicular, 9-veined, ecarinate, shortly laminate or obtuse-apiculate; Fl sessile, distinctly terminal; Sep (5–) 6–7.5  2.5–3.5 mm, (narrowly) ovate, thin-membranous, with overlapping broad hyaline margins, (3- to) 4- to 5-veined, the outer veins joined to the midvein near the apex, subfree, ecarinate, abaxially densely lepidote except the bottom , adaxially glabrous, rounded; Pet tongue-shaped, 10–15  2–2.5 mm, base 2 mm wide, dirty yellow-brown or olive-green, blade recurved, rounded-emarginate; St all of equal length; Fil 7  0.5 mm, flat and attenuate, flaccid, straight or rarely plicate, white; Anth basifixed, 1.8–2.5  0.5 mm, apex obtuse, greenish-yellow; Ov 1.5–2  1.5 mm, ovoid to shortly cylindrical, abruptly contracted into the style; Sty 3 mm, 0.4 mm ∅; Sti short, lobes crested, capitellate, not wider than the style. T. myosura Grisebach ex Baker (J. Bot. 16: 240, 1878). Type: Argentina, Córdoba (Lorentz 122 [BM, B, G, MO, NY, P]). — Distr: S Peru (Arequipa), Bolivia, Argentina, Uruguay; epiphytic in dry habitats, 700–3200 m.  Tillandsia nappii Lorentz & Niederlein (1881) (nom. illeg., ICN Art. 52.1); incl. Tillandsia mandonii E. Morren ex Mez (1896). [4] Plant flowering to 30 cm high or sometimes more, forming dense clusters; stem to 6 ( 10) cm, simple, few-branched or richly branched at the base; L subdensely distichous, 5–17 cm, stiff; L sheaths densely imbricate and margins overlapping behind the stem for most of their length,

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making the stem to appear stout (5–8 mm ∅), 1–1.5 cm, suborbicular to reniform, normally with the upper densely lepidote like the lamina or sometimes glabrous except a fringe of narrow marginal scales, distinct from the lamina; L lamina narrowly triangular or linear, 0.3–0.5 cm wide, densely cinereous-lepidote with small basally produced subappressed to reflexed pruinose scales, strongly recurved, becoming strongly sulcate when dry,  contorted, subulate, with a furrow, apex acuminate and pungent; Inf simple; peduncle bractless or with 1 bract well removed from the flowers, erect, 4–20 cm, lepidote or glabrous; peduncular Bra lanceolate, involute, lepidote; spikes lax, to 8 cm but often much shorter, linear, (1to) 2- to 5- (to 8-) flowered, rachis slender, flexuous in the few-flowered specimens or usually geniculate in the larger ones,  lepidote; floral Bra slightly or not at all imbricate, the lower ones exceeding or equalling or the upper ones often distinctly shorter than the sepals, (13–) 17.5–30  9–9.5 mm, broadly ovate or ovatelanceolate, clasping the flower, chartaceous, many-veined, not concealing the rachis except by their extreme base, rarely >2 as long as the internodes (but much less esp. in the manyflowered specimens), cinereously lepidote, acuminate with a short tip; Fl subsessile, closely appressed to the rachis, fragrant; Sep 11–13.5  5–5.8 mm, oblong-lanceolate, thin, many-veined, evenly short-connate, adaxial ones bluntly carinate, usually sparsely lepidote or glabrescent, rounded or obtuse; Pet linear to tongue-shaped, 13 ( 30)  3 mm, at the base slightly narrowing, pale yellow; St deeply included, exceeding the style; Fil to 5.5 mm; Anth (sub-)basifixed, 3.5  0.5 mm; Ov 4  2 mm, subcylindrical, tapering into the style; Sty  as long as or shorter than the ovary; Sti almost flat and disc-shaped (gemmate). T. neglecta E. Pereira (Bradea 1: 78, t. 2, 1971). Type: Brazil, Rio de Janeiro (Sucre s.n. [HB]). — Distr: SE Brazil (Rio de Janeiro); saxicolous on granitic rock walls. – Fig. 23. [2] Plant caulescent, flowering to 20 cm tall, often forming dense clusters; stem few-branched at the base, curved; L rosulate, somewhat curved,

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polystichous, suberect or divergent, densely arranged, 22–30 mm; Sep 15–16  3 mm, lanceolate, with broad hyaline margins, connate for 2–2.5 mm, ecarinate, glabrous, narrowly acute; Pet spatulate or tongue-shaped with long cuneate base, 22–30  5 mm (base 1.5 mm wide), blue, blade slightly divergent, rounded-emarginate; St 11–16 mm, included, visible in the throat but shorter than the style; Fil slender and narrow at the distal end or ribbon-like, towards the middle slightly plicate, 17.5 mm long; Anth basifixed, 2  0.5 mm, linear, base bilobed, apex obtuse; Ov 3–4  1.5 mm, trigonous or ellipsoid, the top sterile, tapering into the style; Sty 14 mm, slender; Sti erect, linear, unequal.

Fig. 23 Tillandsia neglecta. (Copyright: E. J. Gouda)

4–7 cm, stiff and fleshy-coriaceous, subappressedlepidote on both faces, green or tinged reddish or silver-grey at the base because of spreading trichomes; L sheaths 1.5 cm wide, whitish; L lamina narrowly triangular, not or slightly secund, subulate towards the apex, apex abruptly acute and pungent; Inf simple, mostly much exceeding the leaves; peduncle completely covered by its bracts or partly visible, erect, equalling or exceeding the leaves, glabrous, green; peduncular Bra erect, laxly imbricate, much (3–4 longer than) to slightly exceeding the internodes, ovate-oblong, sparsely pale-lepidote at the margins and apex, long-acuminate; spike 3–4 cm, corymbiform, 6to 10-flowered; floral Bra equalling to much exceeding the sepals, 17  9.5 mm, ovate or elliptic, margins narrowly hyaline, darker veined, subglabrous or lepidote towards the apex (esp. the lower ones), green to reddish esp. at the margins, acutish or obscurely acuminate; Fl sessile,

T. paleacea C. Presl (Reliq. Haenk. 1: 125, 1827). Type: Chile (Haenke s.n. [PR, GH [photo]]). — Distr: C Colombia (Cauca, Huila, Tolima), Peru (widespread), NE Bolivia (La Paz), N Chile, S Paraguay; on rocks and desert sands, or on trees in dry areas, 0–3900 m. – Fig. 24. Incl. Tillandsia fusca Baker (1878); incl. Tillandsia scalarifolia Baker (1887); incl. Tillandsia chilensis Baker (1889); incl. Tillandsia schenckiana Wittmack (1889); incl. Tillandsia favillosa Mez (1906). [3] Plant variable in habit, caulescent, flowering 10–70 cm tall, often much branched; stem 10–35 cm, appearing stout because of the densely imbricate leaf sheaths, brittle; L laxly spirally arranged, densely tomentose-lepidote, silvery-grey often becoming fuscous with age; L sheaths large, broadly ovate, elliptic, glabrous, the upper clearly visible, outside lepidote; L lamina narrowly triangular, to 12 cm, 4–6 mm wide, abruptly spreading, usually slightly flexuous, for most part canaliculate to subulate in the upper part, apex attenuate; Inf simple; peduncle mostly completely covered by its bracts, erect, shorter than to much exceeding the leaves, to 15 cm or sometimes much more, slender, glabrous or nearly so; lowest peduncular Bra somewhat leaf-like with short lamina, soon erect, laxly imbricate, exceeding or about equalling the internodes, narrowly elliptic, tightly enveloping the peduncle, lepidote, apiculate; spike dense, to 5 cm, linear-lanceolate, complanate, (1- to) 2- to

Tillandsia BROMELIACEAE

Fig. 24 Tillandsia paleacea. (Copyright: E. J. Gouda)

5- (to 12-) flowered, acute, rachis angled, straight or flexuous, glabrous, partly exposed; floral Bra distichous, slightly shorter than or slightly exceeding the sepals, 13–16  6 mm, ovate or elliptic, finely veined, ecarinate, so narrow as to expose the rachis in part, 2–3 as long as the internodes, soon glabrous or in the upper part subdensely lepidote, green, sometimes tinged reddish; Fl with a short pedicel-like receptacle; Sep 10–17  5 mm, elliptic or obovate-lanceolate, with hyaline margins, free, ecarinate but slightly fleshy at the base, tinged reddish-purple, glabrous; Pet claw narrow with suborbicular blade, 20–23  10 mm, margins slightly undulate, blade blue to violet or rose, claw white, corolla sometimes with a white eye; St 6–8 mm, deeply included; Anth basifixed, 1.5 mm; Ov subcylindrical, 2.5–3.5 mm, abruptly contracted into the style; Sty stout, 1.5 mm (incl. stigma); Sti disc-shaped, gemmate.

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T. paraensis Mez (in Martius, Fl. Bras. 3(3): 586, t. 109, 1894). Type: Brazil, Pará (Sieber 68 [BR, GH [photo]]). — Distr: SE & S Colombia (Vaupés, Amazonas), E & S Venezuela (Bolívar, Amazonas), Guyana (Cuyuni-Mazaruni, Potaro-Siparuni, Upper Takutu-Upper Essequibo), E & S Suriname (Brokopondo, Sipaliwini), French Guiana (Saül, Régina, Maripasoula), Brazil (Amazonas, Mato Grosso, Rio Grande do Norte, Rondônia, Roraima, Pará, Rio de Janeiro, Bahia, Pernambuco), NE Ecuador, Peru, NE Bolivia; epiphytic in forest, 20–1120 m. Incl. Vriesea sanctae-crucis S. Moore (1895)  Tillandsia sanctae-crucis (S. Moore) Mez (1896); incl. Tillandsia juruana Ule (1907). [5] Plant stemless, flowering 12–40 cm tall; L 10–25, forming a narrowly utriculate rosette, often exceeding the inflorescence or at least the peduncle, 10–35 cm, the outer greatly reduced and sheath-like, very densely subappressed-lepidote throughout; L sheaths forming an elongate pseudobulb, 3–7  1.7–3 cm, ovate, inflatedconvex, stiff-coriaceous, with membranous margins, slightly distinct from and merging into the concolorous lamina; L lamina very narrowly triangular, succulent (fleshy) and coriaceous, 6–25 ( 33)  1.4–2 cm, suberect or curved, often finely keeled on both faces, often the margins involute towards the apex when dry, apex attenuate and abruptly pungent; Inf simple (or rarely compound of 2 spikes), 10–30 ( 40) cm; peduncle completely covered by its bracts or partly visible, often curved, short and concealed by the leaves or elongate, 8–19 cm, 2–3.5 mm ∅, sparsely lepidote or subglabrous; lowest peduncular Bra leaf-like, erect, densely imbricate to remote, ovate, chartaceous, (densely) appressedlepidote or floccose at least at the apex, the upper ones more apiculate; spikes mostly curved, 6–15 ( 20) cm, 1–1.5 cm wide, linear (-lanceolate), complanate, subdensely distichously 4- to 15-flowered, acute, rarely with >1 sterile bract at the apex, rachis sharply angled when dry,  flexuous, sparsely lepidote or glabrate, only partly hidden at anthesis but often wholly exposed when dry; floral Bra scarcely imbricate (if at all), equalling to much exceeding the sepals, 23–39 mm, (broadly) elliptic, clasping the flower,

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scarcely incurved, subchartaceous, even or veined, ecarinate, so narrow as to expose the rachis in part, 2 cm; L sheaths densely imbricate making the stem appear stout, 0.6 cm, suborbicular, scarious, with broad veinless margins, several-veined, glabrous except for the extreme apex, distinct from the lamina; L lamina narrowly triangular, 0.2 cm wide, densely cinereous-lepidote, divergent or spreading from the stem at 90 , with a central keel, with small nearly symmetrical subappressed trichomes, subulate and strongly angled, with a furrow not reaching the apex, apex attenuate and pungent; Inf 1-flowered; peduncle naked for most of its length, erect to ascending, apparently always terminal, almost none or elongate to 2 ( 4) cm, slender, strongly angled, glabrous; peduncular Bra usually 2, one at the extreme base of the peduncle, mostly hidden by the leaves, the other just below the flower, lanceolate, subcoriaceous, several-veined, even, glabrous or occasionally pale-appressed-lepidote, acute or obtuse; floral Bra much shorter than the sepals, 8  4 mm, like the upper peduncular bract but smaller, hyaline, glabrous, pale green, obtuse; Fl subsessile; Sep 9–10  4 mm, narrowly elliptic, membranous, with broad hyaline margins, even, evenly short-connate or slightly longer connate posteriorly for up to 2 mm, pale green, glabrous or sparsely lepidote, acute, obtuse; Pet tongue-shaped, 13  3 mm, yellow to ochrebrown, blade spreading or recurved at anthesis, obtuse; St 6 mm, deeply included, exceeding the

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style with most of the anthers; Anth dorsifixed near the base, >1 mm, linear, apex obtuse; Ov subprismatic, 2.5 mm, contracted into the style; Sty slender, shorter than the ovary; Sti capitellate. T. reducta L. B. Smith (Phytologia 5: 399, t. 1, figs. 11–12, 1956). Type: Peru, Cajamarca (Ferreyra 3216 [US, USM]). — Distr: NE Peru (Cajamarca); only on rocks in higher regions, 2500–2600 m; only known from the type. [1] Plant caulescent, forming dense clusters, stem >8 cm, 0.5 cm ∅ but appearing stout (0.8 cm ∅) because of the densely imbricate leaf sheaths; L persistent, densely tomentose-lepidote with fine linear spreading trichomes; L sheath 1.2–2  1.2–1.8 cm, broadly ovate, thin at the base and margins; L lamina very narrowly triangular, fleshy, spreading or slightly recurved, 4–5 ( 8) cm, subulate but adaxially flat or with a weak furrow; Inf depauperately compound, subdigitate, composed of 3–5 spikes, dense, ellipsoid, fertile part 2–3  1 cm; peduncle completely covered by its bracts, erect, 2–10 cm, 2 mm ∅, slender, lepidote at least near the nodes and in the upper part; lowest peduncular Bra almost leaflike with short lamina, the upper like the primary bracts, erect and with tightly inrolled margins, imbricate, sheath rose, axes of the inflorescence hidden and very short; primary Bra dense, suberect, ovate, to 2.2 cm, nearly as long as the spikes and covering them, thin, strongly veined and carinate, acuminate, abaxially lepidote; spikes with 1 ( 2) sterile bracts at the base, (1- to) 2- to 3-flowered (topmost spike to 5-flowered), with a reduced flower at the apex, rachis stout and short, densely lepidote, internodes 2 mm; floral Bra suberect,  equalling the sepals, 11–13  7–8 mm, ovate or triangular-ovate, incurved, closely enveloping the flowers, chartaceous, veined, scarcely or not at all carinate, so narrow as to expose the rachis in part, sparsely lepidote with few coarse white subappressed trichomes, roseate, acute or apiculate; Fl subsessile, distichous; Sep 9–11  5 mm, oblong, thin, somewhat veined, evenly shortly connate or subfree, adaxial ones carinate (faintly when fresh), rose in the upper part, glabrous, sometimes with a few trichomes at the base, broadly acute; Pet barely exceeding the stamens, tongue-

E. J. Gouda

shaped, 15–21  3–4 mm (base 1.5 mm), centre violet, apical part white (often turning purple after anthesis), blade slightly divergent and then recurved, rounded-emarginate; St 15–17 mm; Fil narrowly ribbon-like, straight; Anth dorsifixed near the base, 3–3.5  0.7 mm, lanceolate, grey-brown; Ov 3–4  1.5 mm, narrowly ovoid or subglobose, contracted into the style; Sty slender, 8–9 mm, slightly longer or shorter than the stamens; Sti spreading, shortly lobed. T. reichenbachii Baker (Handb. Bromel., 166, 1889). Type: K [icono, Reichenbach drawing]. — Distr: SE Brazil (Rio de Janeiro), Bolivia (Santa Cruz, Tarija, Cochabamba, La Paz, Chuquisaca), N Argentina, Paraguay; epiphytic in woods, 100–2500 m.  Tillandsia duratii ssp. reichenbachii (Baker) Halda (2005); incl. Tillandsia tucumanensis Mez (1896); incl. Tillandsia herzogii Wittmack (1916); incl. Tillandsia euosma Spegazzini (1917). [3] Plant caulescent, flowering to >20 cm tall, stem 2–5 cm, stout, curved; L laxly spirally arranged, to 14 cm; L sheaths suborbicular or very broadly elliptic, lustrous, glabrous except for the lepidote apex, very distinct from the lamina; L lamina very narrowly triangular, thick, to 8 cm, 0.8 cm wide, cinereous-lepidote, spreading to reflexed and often recurved at the apex, strongly canaliculate (involute-subulate when dry), slightly flexuous; Inf subdigitate or sometimes simple, composed of 2–8 spikes, to 7  3–4.5 cm; peduncle completely covered by its bracts, erect, 15 cm, 2.5 mm ∅, slender; lowest peduncular Bra somewhat leaf-like but soon with much reduced lamina, imbricate, sublanceolate, enveloping the peduncle, veined, lepidote, subacute; primary Bra like the upper peduncular bracts, much shorter than the stipe of the axillary branches, obtuse; spikes long-stipitate, stipe to 3 cm and densely bracteate, first often erect and then suddenly divergent, densely flowered, 3 cm, 0.9–1 cm wide, linear, complanate, 5- to 7-flowered; floral Bra much shorter than the sepals, 10–11  5 mm, elliptic, clasping the flower, subchartaceous, veined when dry, ecarinate, exposing most of the rachis, 3 as long as the internodes, subglabrous or sparsely lepidote, green tinged with

Tillandsia BROMELIACEAE

reddish-brown, obtuse or acutish; Fl subsessile, slightly divergent or contiguous with each other and/or the rachis; Sep 12–12.5  6 mm, elliptic or obovate, thin-coriaceous, with broad hyaline margins, even, adaxial ones or all evenly shortly connate for 1–2 mm, ecarinate, green tinged with reddish-brown, glabrous, rounded or obtuse; Pet with narrow claw and suborbicular blade, 23  13 mm (base 2.5 mm wide), white or the blade often pale blue, margins crenate-serrate, blade spreading at right angle; St  of equal length, 8 mm, deeply included; Fil ribbon-like, straight; Anth basifixed, 1.5 mm, greenish; Ov 2.5 mm; Sty stout,  as long as the ovary, shorter than the stamens; Sti disc-shaped, gemmate. T. retorta Grisebach ex Baker (J. Bot. 16: 238, 1878). Type: Argentina, Córdoba (Lorentz 125 [BM, B, GOET, M, P]). — Distr: N and C Argentina (widespread to Mendoza and La Pampa in the S); epiphytic, 600–1460 m. Incl. Tillandsia caespitosa Gillies ex Baker (1878); incl. Tillandsia nappii var. darwinii Lorentz & Niederlein (1881) (incorrect name, ICN Art. 11.4). [4] Plant flowering usually 7.5 or rarely to 15 cm tall, forming a globose mass, stem 2–8 cm, many from a single point, usually much branched; L distichous, 3–5 ( 7) cm, densely cinereous-lepidote; L sheaths imbricate, suborbicular, making the stem appear 3–5 mm thick, with broad veinless margins, veined, normally glabrous with a ciliate margin of elongate scales or occasionally in the upper part lepidote like the lamina; L lamina barely over 2 mm wide, recurved, then often towards the apex spreading again, veined when dry, subulate, apex acuminate and pungent; Inf 1- or 2-flowered, densely cinereous-lepidote; peduncle bractless, from almost none to much surpassing the leaves but usually quite short, slender, densely lepidote, reduced rachis extended from the base of the terminal flower; floral Bra erect, equalling or exceeding the sepals, 10–12 ( 15)  7 mm, triangular-ovate, veined, slightly >2 as long as the internodes, densely lepidote, acute, acuminate; Fl subsessile, erect; Sep 10–10.5  3 mm, lanceolate-oblong or narrowly obovate, thin, with broad hyaline margins, finely

1153

veined, adaxial ones connate for 3 mm, abaxial one almost free, adaxial ones carinate, sparsely lepidote, acute or obtuse; Pet erect with slightly divergent apex, tongue-shaped, 12.5  3.3 mm (base 1.3 mm wide), yellow, blade distinct, narrowly elliptic, rounded-emarginate, subtruncate; St 7 mm, deeply included, exceeding the style by of the anthers; Fil ribbon-like and tapering from base to apex, straight; Anth basifixed, 2 mm, linear; Ov 3.5  5 mm, ovoid, abruptly contracted into the style; Sty stout, 2.5 mm; Sti erect, shortly lobed. T. schatzlii Rauh (Trop. subtrop. Pfl.-welt 27: 8, 1979). Type: Mexico, Oaxaca (Rutschmann s.n. in BG Heidelberg 46304 [HEID, US]). — Distr: S Mexico (Oaxaca); steep rock walls; altitude not recorded. [5] Plant stemless or sometimes with a rhizomatous stem; flowering Ros 25  12 cm; L 10, forming a small funnel-shaped rosette, densely appressed-lepidote with trichomes with a green centre, outer leaves reduced and deltoid; L sheaths 4  3 cm, broadly ovate, wrinkled, light brownish to violet inside, inconspicuous from and merging into the lamina; L lamina narrowly triangular, slightly succulent, to 20 cm, 2 mm wide at the base, divergent and often the upper spreading, apex attenuate and subpungent; Inf simple, often subpendulous or erect; peduncle completely covered by its bracts, erect or curved, shorter than the leaves, 5 mm ∅; lower peduncular Bra with short leaf-like blade but soon bladeless, raspberry-red or rose but colour masked by the trichomes; spikes to 16  2 cm, lanceolate, slightly complanate, 8- to 10-flowered; floral Bra imbricate, much exceeding the sepals, 47–60  20 mm, (narrowly) ovate, margins with extended scales, finely veined when dry, ecarinate, densely lepidote, raspberry-red or rosy yellow-green towards the base; Fl to 80 mm (incl. stamens and style); Sep 21.5–27 ( 29)  6–7 mm, narrowly obovate, finely veined when dry, adaxial ones connate for 3–4 mm, adaxial ones bluntly carinate at the base, green, sparsely lepidote with large trichomes, obtuse or acute; Pet forming an erect narrow tube clasping the filaments at the apex, narrowly tongue-shaped with long cuneate

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base, 65–80  9 mm (base 2.5 mm wide), dark violet fading to the white base, blade tip margins recurving; St in 2 whorls of unequal length, 70–85 mm, exserted, shorter than the style; Fil slender and flattened at the base and dilated becoming terete at the distal end, violet in the upper part; Anth dorsifixed, 3 mm, apex obtuse; Ov 5  2.5 mm, ovoid, tapering or gradually contracted into the style; Sty slender, exceeding the petal tube. T. schiedeana Steudel (Nomencl. Bot. ed. 2, 2: 688, 1841). Type: Mexico, Veracruz (Schiede & Deppe 1004 [B, BM, HAL]). — Distr: USA (Missouri), Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Cuba, Jamaica, Haiti, Dominican Republic, N & C Colombia, Venezuela, NW Peru; epiphytic, 0–2250 m.  Tillandsia vestita Schlechtendal & Chamisso (1831) (nom. illeg., ICN Art. 53.1); incl. Tillandsia flavescens Martens & Galeotti (1843); incl. Tillandsia caerulea Grisebach (1865) (nom. illeg., ICN Art. 53.1); incl. Tillandsia grisebachii Baker (1887); incl. Tillandsia eggersii Baker (1889). [5] Plant caulescent, flowering usually 20 ( 40) cm tall, forming a globose mass, stem 5–20 cm, simple or few-branched; L polystichously few-ranked, varying greatly in density, to 25 cm, densely cinereous-lepidote or sometimes ferruginous-lepidote; L sheaths large, suborbicular, densely imbricate; L lamina very narrowly triangular, divergent to spreading at a right angle from the stem, adaxially often with 1 or few grooves, subulate (involute when dry) and angled, shallowly canaliculate, slightly flexuous, apex filiform-attenuate; Inf always simple; peduncle completely covered by its bracts, erect, shorter than the leaves and  as long as the spike; lowest peduncular Bra leaf-like, imbricate, nearly all with leaf-like lamina; spikes 7 cm, 0.7 cm wide, lanceolate, distichously arranged, rachis slender, glabrous; floral Bra densely imbricate, much exceeding the sepals, 27  9 mm, chartaceous and with coriaceous base, veined near the apex, sublustrous, ecarinate, concealing the rachis, 3–4 as long as the internodes, glabrous or the lower ones sparsely lepidote, red or sometimes

E. J. Gouda

yellow, obtuse; Fl sessile, distichous, 45–55 mm; Sep 14 ( 20)  3 mm, lanceolate, thin-coriaceous, even, evenly short-connate for 1 mm, adaxial ones bluntly carinate, yellow, glabrous, rounded or obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, 44  8 mm, lemonyellow, blade tip margins recurved; St in 2 whorls of unequal length, exserted; Fil slender and flattened at the base and dilated becoming terete at the distal end, green; Anth dorsifixed near the middle, apex apiculate, brownish; Ov ellipsoid; Sty shorter than or equalling the shorter stamens; Sti spreading, green. T. seleriana Mez (Bull. Herb. Boissier, sér. 2, 3: 84, 1903). Type: Mexico, sine loco (Seler 3439 [B]). — Distr: SE Mexico (Jalisco, Michoacán, Chiapas, Nayarit, Oaxaca, Veracruz), Guatemala, C & SW Honduras (Francisco Morazán, Lempira), NW El Salvador (Santa Ana), N & W Nicaragua (Estelí, Nueva Segovia); epiphytic in pine and oak woods, 270–2100 m. – Fig. 28. [5] Plant stemless, flowering 20–25 cm tall; L much shorter than to exceeding the inflorescence, outer ones reduced and sheath-like, densely cinereous-lepidote or ferruginous-lepidote; L sheaths forming an ovoid pseudobulb 7–12 cm, broadly ovate or suborbicular, whitish-cinereous-green, merging into the lamina; L lamina linear to triangular, fleshy, 1 cm wide at the base, green (masked by the trichomes but less so than on the sheaths), erect or curved or secund, involute-subulate, apex attenuate; Inf 1 branched, subdigitate or nearly so, composed of 3–6 spikes, dense, 6–10 cm; peduncle often partly visible, erect, short and concealed by the leaves or rarely elongate; peduncular Bra leaf-like, lamina shorter than or exceeding the inflorescence, erect, densely imbricate except the upper ones, with rose-coloured sheath; primary Bra broadly ovate or elliptic, the lower ones slightly shorter and the upper ones much shorter than the axillary branches, shortly laminate or apiculate, densely lepidote, rose-red; spikes sessile, 4 cm, broadly elliptic, strongly complanate, 4- to 8-flowered, rachis straight, densely lepidote, exposed; floral Bra exceeding the sepals except in the top flower and so narrow as to expose part of

Tillandsia BROMELIACEAE

Fig. 28 Tillandsia seleriana. (Copyright: E. J. Gouda)

the sepals, 20–28  9 mm, narrowly elliptic, incurved at the apex, coriaceous, margins with extended scales at the margins, slightly if at all carinate, not concealing the rachis, 4 as long as the internodes, densely cinereous-lepidote with coarse spreading scales, acute; Fl sessile, densely distichous, contiguous with each other; Sep 15.5–17  4 mm, narrowly elliptic, prominently veined, adaxial ones connate for 4–6 mm, adaxial ones distinctly carinate, green or red, glabrous, acute, obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, constricted above the ovary, linear-oblong, 35–37  6 mm (3 mm at the base), violet for most of the length, blade tip margins recurved, narrowly rounded; St in 2 whorls of unequal length, 44–46 mm, exserted; Fil slender and flattened at the base and dilated becoming terete at the distal end; Anth dorsifixed near the middle, black; Ov ovoid, 6 mm; Sty slender, 44 mm, exceeding the stamens; Sti spreading, dilated, white.

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T. subulifera Mez (Repert. Spec. Nov. Regni Veg. 16: 74, 1919). Type: Trinidad (Broadway 4200 [B, B [photo]]). — Distr: NE Nicaragua (Atlántico Norte), W Costa Rica (Puntarenas), C Panama (Panamá), Trinidad & Tobago (CouvaTabaquite-Talparo (Chaguanas)), NW Colombia (Antioquia), Venezuela (Miranda, Sucre, Táchira); epiphytic in forests and plantations, 5–980 m. [5] Plant stemless, flowering 15–19 cm tall; L few, erect, the outer greatly reduced, the inner to 18 cm, appressed-lepidote, with distinct or faint white cross-bands; L sheaths forming a cylindrical pseudobulb, inflated, about as long as the lamina, concolorous with the lamina; L lamina linear, fleshy, coriaceous, 5 mm wide at the base, strongly angled, subulate and shallowly canaliculate, apex obtuse or abruptly acute; Inf simple; peduncle erect, shorter than and largely concealed by the leaves, slender; peduncular Bra erect, imbricate, many-veined, densely lepidote; spikes erect or curved downwards, 5–12 cm, 1.5 cm wide, oblong-lanceolate or linear in outline, complanate, 4- to 12-flowered, without sterile bracts at the apex or base, rachis slender, flexuous to geniculate (when dry), appressed-lepidote, exposed, mostly not or only partly covered by the floral bracts; floral Bra erect, much shorter than the sepals, 20–25  15 mm, elliptic, incurved, margins hyaline, prominently veined,  carinate towards the apex, appressed-lepidote, (yellowish-) green or roseate, broadly acute; Fl with a short pedicel-like receptacle, distichous, suberect; Sep slightly incurved at the apex excluding the apiculus, 22–25  8 mm, narrowly elliptic or narrowly obovate, finely veined (prominently when dry), free, ecarinate, rose, cinerous-appressed-lepidote, apiculate or narrowly obtuse; Pet forming a tube slightly widening towards the apex, linearoblong, 32–34  7 mm (at the base 5 mm), rose, upper part yellow, often with rose margins and apex, narrowly rounded; St  32 mm, included and visible in the throat; Fil terete, slightly coiled halfway, rose towards the distal end; Anth dorsifixed near the middle, 3  1.8 mm, oblong, base bilobed, apex obtuse and apiculate; Ov 5.5  3 mm, ovoid, tapering form near the base, slightly contracted into the style; Sty slender, to 27 mm; Sti weakly conduplicate, spreading.

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E. J. Gouda

Fig. 29 Tillandsia tectorum. (Copyright: E. J. Gouda)

T. tectorum E. Morren (Belgique Hort. 27: 328, t. 18, 1877). Type: Peru (Wallis s.n. [BR]). — Lit: Hromadnik (2005: synopsis T. tectorumcomplex). Distr: N & S Ecuador (Azuay, El Oro, Loja, Pichincha), Peru (Amazonas, Cajamarca, Huánuco, La Libertad, Ancash, Lima); saxicolous, and cultivated on roofs, 680–3400 m. – Fig. 29. Incl. Tillandsia argentea K. Koch (1867) (nom. illeg., ICN Art. 53.1); incl. Pourretia nivosa hort. ex E. Morren (1877) (nom. inval., ICN Art. 36.1c); incl. Tillandsia saxicola Mez (1906). [1] Plant short-stemmed to long caulescent, flowering 30–100 cm tall; L densely rosulate, 12–25 cm, white-tomentose-lepidote; L sheaths 1.2–3 cm wide at the base, triangular-ovate, clasping the stem, fleshy in the upper part, densely lepidote except the inner base, distinct from the lamina; L lamina very narrowly triangular, fleshy and 0.8–1.5 cm wide at the base, silvery-white, divergent to spreading, apex filiform-attenuate, trichomes eccentric, with elongated wings; Inf compound, densely digitate with 4–6 spikes, 5–10  (2.5–) 5–8 cm; peduncle erect, much exceeding the leaves, 10–40 cm  5 mm ∅, slender, densely lepidote; lowest peduncular Bra somewhat leaf-like with setaceous lamina, sheaths imbricate, 1.5–2.5  0.8–1.2 cm, broadly ovate, membranous, veined, adaxially scatteredly

lepidote and abaxially densely lepidote; primary Bra like the upper peduncular bracts, much shorter than the spikes; spikes sessile, erect or somewhat spreading, 3–4.5  1–1.2 cm, lanceolate, strongly complanate, (3- to) 5- to 10-flowered; floral Bra densely imbricate,  equalling to exceeding the sepals, 10–17  7–11 mm, ovate, thin, veined, carinate at least at the apex, lepidote to subglabrous, red, green or roseate, acutish or apiculate; Fl subsessile, 20 mm; Sep 9–15  3–4 mm, ovatelanceolate, thin, with broad hyaline margins, free, adaxial ones carinate, green and flushed red in the upper , abaxially sparsely lepidote along the midvein or glabrous; Pet erect and forming a tube, tongue-shaped, 15–25  3–4 mm, white with a blue band in the middle section, paler blue to whitish towards the apex, becoming pink after anthesis, blade divergent, obtuse; St all of equal length, 12–17 mm, included but visible in the throat; Fil narrowly ribbon-like, flaccid, straight or plicate, white; Anth subbasifixed or dorsifixed at from the base, 2–3 mm, linear, black or brown; Ov 2.5–3  2 mm, ovoid; Sty slender, 5–15 mm, longer than the stamens; Sti with short somewhat spreading lobes. T. tenebra L. Hromadnik & W. Till (Bromelie 1991(2): 32, ill., 1991). Type: Argentina, La Rioja

Tillandsia BROMELIACEAE

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Fig. 30 Tillandsia tenebra. (Copyright: E. J. Gouda)

(Hromadnik & Hromadnik 7275 [WU, WU]). — Distr: Argentina (Catamarca, La Rioja, Buenos Aires, San Luis); saxicolous, 800–1900 m. – Fig. 30. Incl. Tillandsia myosura var. saxicola Hieronymus ex A. Castellanos (1945) (nom. inval., ICN Art. 36.1c). [4] Plant short-stemmed, flowering 4–6 cm tall, few-branched or forming clumps; L densely distichous, appearing to be equitant; L sheaths densely imbricate, 1–1.2  1.4 cm, upper part densely cinereous-lepidote like the lamina; L lamina succulent, rigid, 4–6  0.4–0.5 cm, recurved or spreading, adaxially grooved in the lower , subulate and laterally compressed, apex subulately acuminate and pungent, densely cinereouslepidote, with extending trichomes at the margins, trichomes suborbicular; Inf simple; peduncle bractless or with 1–2 bracts at the base, erect, very short to elongate, lustrous, glabrous or lepidote near the first flower, often dark brown; peduncular Bra 2–4.5 cm, tightly enveloping the peduncle, fleshy, short-caudate; spike 1- to 2(to 3-) flowered, rachis straight, lepidote, barely visible, internodes 0.25–0.4 ( 0.65) cm; floral Bra much shorter than the sepals, (7–) 10–12 ( 15)  6–7 ( 8) mm, broadly ovate or rarely triangular, margins hyaline, distinctly 14- to 16veined, the lower one densely appressed-lepidote

and the upper sparsely lepidote esp. at the base, cinereous or brown-red, apiculate; Fl subsessile, distichous, strongly fragrant; Sep (8–) 9.5–11  (3–) 3.5–4.5 mm, elliptic, thin-coriaceous at the base, 8- to 10-veined (distinctly so at the apex), unequally short-connate for 1–3 mm, adaxial ones with a strong midvein and slightly carinate near the apex, brown to red in the upper part, glabrescent or sparsely appressed-lepidote at the apex, rounded or apiculate; Pet spatulate, 12–14 ( 15)  3.5–4.2 mm (base 1.5 mm wide), fleshy, with a thick midvein or with ridges, dark brownviolet, blade broadest near the apex, tapering into the cuneate claw, recurved, rounded; St all of equal length, 7 mm, deeply included; Fil 4–5.5 mm, ribbon-like and tapering from near the base to the apex; Anth basifixed, 1.5  0.5 mm, yellow; Ov 3.4–4 ( 6)  2–2.5 ( 3) mm, angular-obconical, abruptly contracted into the style; Sty stout, 1.1–1.5 mm, 0.5–0.8 mm ∅, almost conical; Sti capitate. T. tortilis Klotzsch ex Baker (J. Bot. 25: 237, 1887). Type [lecto]: Mexico, San Luis Potosí (Parry & Palmer 872 [BM, E, GH, K, MO, P, US]). — Lit: Till (1986). Distr: C & E Mexico, Jamaica; epiphytic or saxicolous, 1830–2440 m.  Viridantha tortilis (Klotzsch ex Baker) Espejo (2002); incl. Platystachys ehrenbergii

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Beer (1856) (nom. inval., ICN Art. 36.1); incl. Platystachys tortilis Beer (1856) (nom. inval., ICN Art. 36.1); incl. Tillandsia ehrenbergii Klotzsch ex Beer (1856) (nom. inval., ICN Art. 38.1); incl. Tillandsia tortilis Klotzsch ex Beer (1856) (nom. inval., ICN Art. 38.1); incl. Platystachys ehrenbergii K. Koch (1874) (nom. inval., Art.?)  Tillandsia ehrenbergii (K. Koch) Klotzsch ex Mez (1896); incl. Tillandsia ehrenbergiana Baker (1889). [1] Plant short stemmed to distinctly caulescent, flowering 10–20 cm tall, forming dense clusters; stem 3–5 cm, simple or branched; L 15, densely rosulate, to 17 cm, densely tomentose-lepidote or villous, with fine silver-grey scales; L sheaths 2  1.2 cm, elliptic, the lower part membranous, strongly veined, densely tomentose except at the base, distinct from the lamina; L lamina 6–15  0.5–1 cm, mostly spreading to reflexed, with a distinct midvein adaxially, with involute margins, subulate towards the long-attenuate apex, trichomes with elongated wings; Inf always simple; peduncle erect or ascending, much shorter than to just exceeding the leaves, 4–8 cm, 1–2 mm ∅, strongly sulcate, glabrous; lowest peduncular Bra leaf-like, imbricate, much exceeding the internodes, lanceolate, involute, thin, strongly veined, lepidote, the upper roseate, acuminate; spike 3–4  1–1.6 cm, elliptic, strongly complanate, 3- to 10-flowered, rachis slender, nearly straight; floral Bra densely imbricate, much exceeding the sepals, 20–26  8–12 mm, lanceolate, membranous, veined at least when dry, upper distinctly carinate, concealing the rachis, 3–4 as long as the internodes, abaxially cinereously lepidote, roseate, acute or acuminate; Fl subsessile, distichous, suberect, dense; Sep (16–) 17–20  3–5 mm, lanceolate, membranous, strongly veined, subfree, adaxial ones carinate, rose but green in the lower , lepidote, acuminate; Pet erect with slightly divergent apex, linearly tongue-shaped, 25–38 ( 40)  4 mm, green, lower white, rounded; St 24–31 mm, included (reported as exserted by Mez); Fil white; Anth basifixed, 4 mm; Ov 4.5 mm, ellipsoid; Sty slender, 11–20 mm, shorter or  equalling the stamens; Sti erect to spreading at maturity, green.

E. J. Gouda

T. violaceiflora L. Hromadnik (Bromelie 2012 (2): 49, ills. (pp. 48, 50), 2012). Type: Argentina, Salta (Hromadnik 7176 [MCNS, WU]). — Distr: NE Argentina (Salta: Quebrada de las Conchas); rocky cliffs, 1500 m. [5] Plant saxicolous, nearly stemless forming a short, 3 mm thick rhizome, flowering Ros only 10–12  6–10 cm, growing in groups; L 4.5–9 cm, stiff; L sheaths 0.6–1  1.2–2 cm, trapezoid, amplexicaul, adaxially veined, glabrous, abaxially the upper lepidote like the lamina, pale brownish, slightly distinct from the lamina; L lamina narrowly triangular, fleshy, rigid, 0.7–1 cm wide at the base, divergent and the longer ones upwardly secund, finely veined, strongly canaliculate and subulate at the somewhat pungent apex, cinereous-lepidote; Inf simple, erect or somewhat decurved, 6–8 cm, equalling or exceeding the leaves; peduncle completely covered by its bracts, short and concealed by the leaves, 3–4 cm, 2 mm ∅, glabrous, white; peduncular Bra 2–3  0.7 cm, oblong, membranous, strongly veined, glabrous or sparsely pale-lepidote on the lamina, the basal ones pale green, greenish-white or whitish, acuminate; spike 4  1.2 cm, lanceolate, strongly complanate, densely 3- to 5-flowered, rachis 4-angled, slender, glabrous, partly exposed, green, internodes 5 mm; floral Bra exceeding the sepals, 30  12 mm, lanceolate, subcoriaceous, smooth but rapidly veined after anthesis, glabrous, greenish to ruby-red, apiculate; Fl sessile, distichous, erect and densely appressed to the rachis, 40 mm, not fragrant; Sep 15–18  4 mm, lanceolate, the adaxial pair nearly free, adaxial ones carinate, keels red; Pet forming a narrow tube exceeding the floral bracts, then spreading, spatulate, 40  to 5 mm (base 2 mm wide), violet, the lower white, obtuse, abaxially weakly carinate; St 25 mm, included; Fil ribbon-like, straight, white; Anth subbasifixed, 3 mm; Ov 5  2 mm, ellipsoid; Sty slender, 25 mm, exceeding the stamens; Sti reaching the throat of the corolla, the narrow lobes spreading, white. T. virescens Ruiz & Pavón (Fl. Peruv. 3: 43, t. 270: fig. b, 1802). Type: Peru, Huánuco (Ruiz & Pavón s.n. [MA?/BM?, F]). — Lit: Till (1989);

Tillandsia BROMELIACEAE

Maruska (2012); Castello & Galetto (2013). Distr: SE Brazil (Rio de Janeiro), Peru, Bolivia, N & C Chile, N to NC Argentina, C Paraguay; saxicolous or epiphytic in dry habitats, 3–4000 m.  Diaphoranthema virescens (Ruiz & Pavón) Beer (1856)  Tillandsia capillaris fa. virescens (Ruiz & Pavón) L. B. Smith (1935); incl. Tillandsia propinqua Gay (1853); incl. Tillandsia recurvata Grisebach (1874) (nom. illeg., ICN Art. 53.1); incl. Tillandsia cordobensis Hieronymus (1885)  Tillandsia capillaris fa. cordobensis (Hieronymus) L. B. Smith (1935); incl. Tillandsia stolpii Philippi (1895); incl. Tillandsia dependens Hieronymus ex Mez (1896); incl. Tillandsia dependens var. percordobensis Mez (1896)  Tillandsia dependens fa. percordobensis (Mez) A. Castellanos (1933); incl. Tillandsia dependens var. perusneoides Mez (1896) (nom. inval., ICN Art. 26.1?)  Tillandsia dependens fa. perusneoides (Mez) A. Castellanos (1933) (nom. inval., ICN Art. 26.1?); incl. Tillandsia williamsii Rusby (1910); incl. Tillandsia dependens var. sanzinii Hicken (1914)  Tillandsia virescens var. sanzinii (Hicken) A. Castellanos (1945); incl. Tillandsia tomasii Halda (2005). [4] Plant small but very variable in both size and form, flowering 2–16 cm tall, densely clumpforming; stems many from a single point, simple or branched; L distichous, mostly 1–4 ( 9) cm (rarely 100, often with long filiform Fil and basifixed basally deeply

E. Van Jaarsveld (*) Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa e-mail: [email protected] U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected]

sagittate Anth opening with longitudinal slits; Ov superior, consisting of 3 Ca, 1-locular with a single ovule and with 2 (-3) often long and feathery Sti; Fr a caryopsis (grain) consisting of a single Se usually fused with the fruit wall, usually enclosed in the dry persistent lemma and palea, indehiscent. Order: Poales Important Literature: Grass Phylogeny Working Group (2011: phylogeny); Kellogg (2015: synopsis). Distribution: Worldwide. Grasses (alternatively known by the family name Gramineae) embrace some 700 genera and 11'127 species, and they are sister to a group of restioid families, either as sister to the Ecdeiocoleaceae (Graham & al. 2006), or as sister to the Joinvilleaceae (Givnish & al. 2006). The Grass family is among the economically and ecologically most important plants worldwide, and many species such as wheat, oats, rice, corn are important food plants – wheat, rice, and maize provide >50% of all calories consumed by humans (Mabberley 2008), and many species are important elements of animal (esp. cattle) fodder. Numerous other species are of ethnobotanical importance, from building material (bamboos) to sugar (Saccharum officinarum, “sugar cane”) and spices (Cymbopogon citratus, “lemon grass”).

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_22

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Dr

123 4

Bambus.

Pooideae

Ar.

Panicoideae

MA D Chloridoid.

2000 species

Fig. 1 Summary phylogeny of Poaceae (11’127 species total, based on Grass Phylogeny Group (2012). 1 = Anomochloideae, 2 = Pharoideae, 3 = Puelioideae, 4 =

Ehrhartoideae, A = Arundinoideae, Ar. = Aristidoideae, D = Danthonioideae, Dr = Dregeochloa, M = Micrairoideae. (Copyright: U. Eggli)

Many vegetation communities are dominated by grasses, esp. in regions with periodic drought, and grasslands such as prairie and pampas or steppes cover almost 25% of the planet’s land surface. Succulence: Leaf succulence has evolved only in the small genus Dregeochloa, placed in subfamily Arundinoideae by Kellogg (2015) (Fig. 1). D. pumila co-occurs with many other succulents in South Africa (Jaarsveld 2009). Goebel (1889: 46) reports a small degree of succulence (confined to the upper hypodermis) for the leaves of Spinifex squarrosus, but the species appears to be merely xerophytic.

H. S., Roalson, E. H., Evans, T. M., Hahn, W. J., Millam, K. C., Meerow, A. W., Molvray, M., Kores, P. J., O’Brien, H. E., Hall, J. C., Kress, W. J. & Sytsma, K. J. (2006) Phylogenetic relationship of Monocots based on the highly informative plastid gene ndhF: Evidence for widespread concerted convergence. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 28–51. https://doi.org/10.5642/aliso.20062201.04 Goebel, K. (1889) Pflanzenbiologische Schilderungen. Erster Teil. Marburg (DE): N. G. Elwert’sche Verlagsbuchhandlung. https://doi.org/10.5962/bhl. title.1665 Graham, S. W. [& al. 2006], Zgurski, J. M., McPherson, M. A., Cherniawsky, D. M., Saarela, J. M., Horne, E. F. C., Smith, S. Y., Wong, W. A., O’Brien, H. E., Biron, V. L., Pires, J. C., Olmstead, R. G., Chase, M. W. & Rai, H. S. (2006) Robust inference of Monocot deep phylogeny using an expanded multigene plastid data set. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 3–21. https://doi.org/10.5642/aliso.20062201.02 Grass Phylogeny Working Group (2011) New grass phylogeny resolves deep evolutionary relationships and

References Givnish, T. J. [& al. 2006], Pires, J. C., Graham, S. W., McPherson, M. A., Prince, L. M., Patterson, T. B., Rai,

Poaceae discovers C4 origins. New Phytol. 193: 304–312. https://doi.org/10.1111/j.1469-8137.2011.03972.x Jaarsveld, E. J. van (2009) Kleinduimpie grass. Dregeochloa pumila, the only succulent grass in the world. Veld Fl. (1975+) 95(1): 19–21, ills. Kellogg, E. A. (2015) Poaceae: In: Kubitzki, K. (ed.): The families and genera of vascular plants. Vol. XIII.

1309 Flowering plants. Monocots. Cham (CH) etc: Springer International Publishing Switzerland. https://doi.org/ 10.1007/978-3-319-15332-2 Mabberley, D. J. (2008) Mabberley’s Plant-Book. A portable dictionary of plants, their classification and uses. Ed. 3. Cambridge (GB): Cambridge University Press.

Dregeochloa POACEAE E. Van Jaarsveld

Dregeochloa Conert (Senckenberg. Biol. 47(4): 335, 1966). Type: Danthonia pumila Nees. — Arundinoideae — Lit: Gibbs Russel & al. (1991); Jaarsveld (2009). Distr: Namibia, NW RSA. Etym: For Jean François Drège (1794–1881), German plant collector in S Africa 1826–1833; and Gr. ‘chloe, chloa’, young green sprouts, grass. Tufted stoloniferous or caespitose perennials with short spreading rhizomes, rooting at the nodes; L lamina linear to ovate or ovate-lanceolate, succulent or not, involute, ligule minutely ciliate; Inf racemose or paniculate with 4–12 spikelets; spikelets solitary, with 4–6 fertile florets,  laterally compressed, lanceolate, acute, disarticulated above the glumes, rachis and Ped hairy; glumes 2, 5- to 7-veined, lanceolate, without awn; incomplete florets distal, awned, lemmae ovate, 7- to 9-veined, apex with 2 acute bristle-tipped lobes, membranous, with transverse rows of minutely hairy tufts; awns solitary, geniculate; palea present, narrowly elliptic, prominently 2-veined, apex rounded to emarginate, ciliate distally, hyaline; lodicles 2, fleshy; Ov sparingly hairy; Fr small, pericarp free. A small genus of just 2 species, mainly native to the desert and semi-desert regions of the

Northern Cape and Western Cape, and adjacent Namibia. Only 1 is succulent. In an anatomical study, Ellis (1977) found weakly expressed all-cell succulence. D. pumila (Nees) Conert (Senckenberg. Biol. 47 (4): 335–343, ill., 1966). Type: RSA, Northern Cape (Drège 2562 [B, K]). — Distr: S Namibia, NW RSA (Northern Cape); Western Gariep Lowland Desert vegetation, on sandy rocky wind-blown hills of the narrow coastal belt, flowers spring to early summer.  Danthonia pumila Nees (1899). Plants with short rhizomes, tufted and forming small mats, to 7 cm tall, base of individual growths with broad scales; L lamina succulent, 10–25  1.6–3.5 mm, minutely pubescent, apex obtuse, spiny-apiculate; Inf racemose or paniculate; spikelets 6- to 10-flowered; glumes 9–13 mm, 5- to 7-veined; lemma 3–3.5 mm; central awn 4–7 mm. The species was discovered by Drège in 1830 in the Richtersveld. Difficult to cultivate away from its native habitat, and best grown in a greenhouse under controlled conditions.

E. Van Jaarsveld (*) Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_23

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References Ellis, R. P. (1977) Leaf anatomy of the South African Danthonieae (Poaceae). I. The genus Dregeochloa. Bothalia 12(2): 209–213, ills., map. https://doi.org/10. 4102/abc.v12i2.1398 Gibbs Russel, G. E. [& al. 1991], Watson, L., Koekemoer, M., Smook, L., Barker, N. P., Anderson, H. M.,

E. Van Jaarsveld Dallwith, M. J. & Leistner, O. A. (ed.) (1991) Grasses of southern Africa. Mem. Bot. Surv. South Afr. 58: 437 pp., keys. Jaarsveld, E. J. van (2009) Kleinduimpie grass. Dregeochloa pumila, the only succulent grass in the world. Veld Fl. (1975+) 95(1): 19–21, ills.

Part XVII The Family Ruscaceae

Ruscaceae U. Eggli

Including Aspidistraceae Hasskarl. Including Convallariaceae Horaninow. Including Dracaenaceae Salisbury. Including Nolinaceae Nakai. Including Ophiopogonaceae Meissner. Including Peliosanthaceae Salisbury. Including Polygonataceae Salisbury. Including Sansevieriaceae Nakai. Including Tupistraceae Schnizlein. Trees or shrubs and then often pachycaulous, deciduous herbaceous perennials, or rhizomatous herbs; L alternate, spiral or rarely distichous, sometimes reduced to scales and replaced by phylloclades (Ruscaceae s.s.), basally somewhat to distinctly amplexicaul, often in rosettes, strapshaped to ensiform, more rarely terete, or angularwiry, usually fibrous-tough and xeromorphic, sometimes slightly to distinctly succulent, margins entire or rarely with prickles, tips blunt, tapering and hardened, or fibrous, almost never sharp and pungent; Inf usually terminal but sometimes lateral, or appearing lateral due to later growth of branches, small to huge, usually richly branched and often paniculate, or elongate with condensed lateral branches or fascicles of flowers, or condensed and capitate; Fl often smallish and

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

inconspicuous, usually numerous to very numerous, actinomorphic, hermaphrodite or unisexual, white, whitish, greenish or yellowish, very rarely orange-reddish; Tep 3 + 3 (or sometimes 4), free or more commonly fused and Per tubular to urceolate; St 6, often long exserted; Ov superior, 1- or 3-locular with 1 ovule per locule; Fr few-seeded globose red to orange berries, or dry indehiscent capsules (functionally nutlets); Se globose to elongate, without true testa, pale-coloured. Order: Asparagales Important Literature: Trelease (1911: synopsis Nolineae); Bogler (1998a: synopsis Nolinaceae); Perry & Rudall (1998: synopsis Dracaenaceae s.s); Rudall & al. (2000: phylogeny); Kim & al. (2010: molecular phylogeny); Lu & Morden (2014: molecular phylogeny dracaenoid genera). Distribution: Tropical and subtropical Africa, Arabia, Asia and Australasia, arid N America. A family of 25 genera and 375 species, including several horticulturally important taxa (incl. species of Dracaena and Sansevieria), whose circumscription has changed considerably in the past. Classification and History: Traditionally, Ruscaceae embraced only 3 genera and some 10 species with reduced leaves and photosynthetic phylloclades from the Mediterranean and Asia Minor, but all recent molecular phylogenies agree that Ruscaceae s.s. and the traditional

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_24

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families Dracaenaceae, Eriospermaceae and Nolinaceae, together with the non-succulent Convallariaceae and related herbaceous perennials, form a monophyletic clade that could be recognized as a widely cirumscribed Ruscaceae s.l. as part of the core Asparagales (Rudall & al. 2000, Bogler & al. 2006, Givnish & al. 2006, Graham & al. 2006, Kim & al. 2010), Together with the Asparagaceae s.s., the clade diagnostically shows articulated pedicels (Graham & al. 2006) and (with the exception of Eriospermaceae) juicy or dry berrylike fruits (Givnish & al. 2006). Janssen & Bremer (2004) and APG (2009) opted for inclusion of the Ruscaceae in their very widely circumscribed Asparagaceae s.l. (as subfam. Nolinoideae) (Chase & al. 2009). Nyffeler & Eggli (2010) argued against such a wide circumscription of Asparagaceae, and also felt that the Eriospermaceae, identified as basal sister to the remaining Ruscaceae s.l., and characterized by their tuberous corms, herbaceous growth, capsular fruits and woolly seeds, should be recognized as separate taxon. Seberg & al. (2012) found the same general topology for the core Asparagales but stress that the clade of Ruscaceae s.l. is one of the most weakly supported parts of their phylogeny, and also Steele & al. (2012) have limited support for the relationships within Asparagaceae s.l., where Ruscaceae s.l. are found as sister of Asparagaceae s.s. The inconclusiveness of the data thus not automatically supports a wide Nolinoideae

*Convallarioideae

circumscription of Ruscaceae or Asparagaceae. Here we follow Judd & al. (2016) and Nyffeler & Eggli (2010) and recognize Ruscaceae s.l. as separate from Eriospermaceae. Ruscaceae s.l. in this circumscription represent a clade with many drought-adapted species with fibrous and therefore “tough” leaves, conspicuous to smallish to insignificant white or pale-coloured flowers, and juicy berries or indehiscent capsules. Within the expanded Ruscaceae, subfam. Nolinoideae (representing the former Nolinaceae) occupies a basal position in sister-group position to a grade of “Convallariaceae” tribes, with subfam. Ruscoideae (= Ruscaceae s.s.) and subfam. Dracaenoideae (= former Dracaenaceae) as sisters in terminal position (Fig. 1). The subfamilies with succulents are characterized as follows: • Dracaenoideae (3 genera,  150 species): Trees, shrubs, or rhizome geophytes; L in rosettes, rarely bifarious, usually fibrous, xeromorphic, not to distinctly succulent, flat or semi-terete to terete, tip at most hardend but not strongly pungent; Inf terminal but sometimes appearing lateral, appearing spicate to paniculate, sometimes capitate; Fl many to very numerous, bisexual, actinomorphic, small to conspicuous, pale-coloured (often white); Ov superior, 3-locular; Fr globose berries; Se without true testa and therefore remaining fleshy, but covered by a flimsy scarious envelope derived Dracaenoideae

R

50 species Fig. 1 Summary phylogeny of Ruscaceae based on Kim & al. (2010) (375 spp.; data from Bogler & al. (1998) and Bos (1998), and own counts). R = Ruscoideae. (Copyright: U. Eggli)

Ruscaceae

from the innermost part of the endocarp (Budweg 2015). — This subfamily was formerly recognized as separate family Dracaenaceae. The close relationship between Dracaena and Sansevieria (both in the more distant past classified as Agavaceae) was known for long, and Bos (1984) and Bos (1998) suggested that they should be treated as congeneric. According to Jankalski (2004) the two genera differ in pollen morphology, and should not be united. Preliminary observations by Budweg (2015) suggest that there might also be differences in germination and seedling root systems between the genera. The chloroplast DNA data of Lu & Morden (2014) shows that the dracaenoid genera form a monophyletic clade, and that monophyletic Sansevieria is nested within Dracaena, rendering the latter paraphyletic. For a phylogenetically correct classification, Sansevieria ought to be included in Dracaena, but this appears impractical, and unpublished results (L. E. Newton, pers. comm.) appear to support recognition of Sansevieria. In the first issue of this handbook, Cordyline was also included in the Dracaenaceae, but this genus is now part of Laxmanniaceae. • Nolinoideae (4 genera, c. 50 species): Woody pachycaul trees or shrubs, or rosette plants, or

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caudiciform, or rhizome geophytes; L in rosettes, fibrous, xeromorphic, at most very slightly succulent with fibrous (never pungent) tip; Inf terminal, often very large, paniculate; Fl usually very numerous, unisexual (and plants dioecious or polygamo-dioecious), actinomorphic, small, pale-coloured; Ov superior, 1- or 3-locular; Fr dry indehiscent capsules. — This subfamily was formerly recognized as separate family Nolinaceae. Earlier, these genera were placed in the Agavaceae. Rowley (1990) considered Beaucarnea and Calibanus to be synonymous with Nolina. However, molecular data strongly indicates that Calibanus should be included in Beaucarnea, which is closely related but distinct from Nolina. Succulence: While tough, somewhat thickish and xeromorphic leaves are not rare in the family, undisputed leaf succulence is restricted to Sansevieria. Scant parenchymatic water storage tissue in leaves is otherwise known from a handful of species such as Dasylirion wheeleri or Nolina microcarpa (Walter 1931: 99, 153). The massive pachycaulous stems of Beaucarnea and Dracaena consist of voluminous parenchymatic tissue with numerous scattered vascular bundles, and the basal swelling in Beaucarnea is the result

Key to genera with succulents 1a 1b

2a 2b 3a 3b 4a 4b 5a 5b

L usually distinctly succulent, flat to terete, margins never armed; stems short, often horizontal, subterranean and rhizomatous, or absent: L not truly succulent, flat to narrow and wiry, sometimes with marginal prickles; stems evident and often with richly branched crown, or massively pachycaulous, or as a globose caudex, or apparently absent or subterranean: Stems evident, of  even diameter over their length, branching after flowering and often forming huge crowns; Fl bisexual: Stems absent or largely subterranean, as caudex, or pachycaulous and basally swollen; Fl unisexual: L margins with sharp curved prickles: L margins without sharp prickles: Stem a large depressed to globose caudex without apparent branches; L  wiry; Fr globular: Stem absent or subterranean, or small to large and pachycaulous (globose only when young), with few to numerous branches in old plants: Stem prominent, pachycaulous and basally swollen in adult plants (globose only when young); Ov 1-celled; Fr 3-winged capsules: Stem absent, when present woody at maturity, never swollen basally and never globose when young; Ov 3-celled; Fr with 1 seed per locule but often only 1 locule developing:

Sansevieria 2

Dracaena 3 Dasylirion 4 Dasylirion (Calibanus) 5 Beaucarnea Nolina

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of secondary thickening growth due to a “monocot cambium” (Stevenson 1980, Carlquist 2012). The stems and rhizomes of Dasylirion and Nolina are merely woody with at most minimal water-storage capacity.

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 Bogler, D. J. (1998a) Nolinaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 392–397, ills., key. Berlin (DE) etc.: Springer Verlag. https://doi. org/10.1007/978-3-662-03533-7_46 Bogler, D. J. [& al. 2006], Pires, C. & Francisco-Ortega, J. (2006) Phylogeny of Agavaceae based on ndhF, rbcL, and its sequences: Implications of molecular data for classification. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 313–328. Bos, J. J. (1984) Dracaena in West Africa. Agric. Univ. Wageningen Pap. 84-1: 126 pp., ills., maps, key. Bos, J. J. (1998) Dracaenaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 238–241, ills., key. Berlin (DE) etc.: Springer Verlag. https://doi.org/ 10.1007/978-3-662-03533-7_30 Budweg, H.-G. (2015) Vom Beginn des Lebens der Sansevierien: Beobachtungen an meinen Pflanzen. Sansevieria Online 3(2): 24–34, ills. Carlquist, S. (2012) Monocot xylem revisited: New information, new paradigms. Bot. Rev. (Lancaster) 78: 87–153, ills. https://doi.org/10.1007/s12229-012-9096-1 Chase, M. W. [& al. 2009], Reveal, J. L. & Fay, M. F. (2009) A subfamilial classification for the expanded Asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Bot. J. Linn. Soc. 161(2): 132–136. https://doi.org/10.1111/j.1095-8339.2009. 00999.x Givnish, T. J. [& al. 2006], Pires, J. C., Graham, S. W., McPherson, M. A., Prince, L. M., Patterson, T. B., Rai, H. S., Roalson, E. H., Evans, T. M., Hahn, W. J., Millam, K. C., Meerow, A. W., Molvray, M., Kores, P. J., O'Brien, H. E., Hall, J. C., Kress, W. J. & Sytsma, K. J. (2006) Phylogenetic relationship of Monocots based on the highly informative plastid gene ndhF: Evidence for widespread concerted convergence. In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 28–51. Graham, S. W. [& al. 2006], Zgurski, J. M., McPherson, M. A., Cherniawsky, D. M., Saarela, J. M., Horne, E. F. C., Smith, S. Y., Wong, W. A., O'Brien, H. E., Biron, V. L., Pires, J. C., Olmstead, R. G., Chase, M. W. & Rai, H. S. (2006) Robust inference of Monocot deep phylogeny using an expanded multigene plastid data set.

U. Eggli In: Columbus, J. T. & al. (eds.): Monocots. Comparative biology and evolution excluding Poales. Aliso 22: 3–21. Jankalski, S. (2004) Sansevieria versus Dracaena. Sansevieria 8: 17–18, ill. Janssen, T. & Bremer, K. (2004) The age of major monocot groups inferred from 800+ rbcL sequences. Bot. J. Linn. Soc. 146(4): 385–398. https://doi.org/ 10.1111/j.1095-8339.2004.00345.x Judd, W. S. [& al. 2016], Campbell, C. S., Kellogg, E. A., Stevens, P. F. & Donoghue, M. J. (2007) Plant systematics: A phylogenetic approach. Ed. 4. Sunderland (US): Sinauer Associates. Kim, J.-H. [& al. 2010], Kim, D.-K., Forest, F., Fay, M. F. & Chase, M. W. (2010) Molecular phylogenetics of Ruscaceae sensu lato and related families (Asparagales) based on plastid and nuclear DNA sequences. Ann. Bot. (Oxford), n.s. 106: 775–790. https://doi.org/10.1093/aob/mcq167 Lu, P.-L. & Morden, C. W. (2014) Phylogenetic relationships among Dracaenoid genera (Asparagaceae: Nolinoideae) inferred from chloroplast DNA loci. Syst. Bot. 39(1): 90–104. https://doi.org/10.1600/ 036364414X678035 Nyffeler, R. & Eggli, U. (2010) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125-144. https://doi.org/10.5167/uzh35243 Perry, P. L. & Rudall, P. J. (1998) Eriospermaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 241–244, ills., key. Berlin (DE) etc.: Springer Verlag. https://doi.org/10.1007/978-3-662-03533-7_31 Rowley, G. D. (1990) Nolina en culture. Cact.-Avent. 5: 2–4. Rudall, P. J. [& al. 2000], Conran, J. G. & Chase, M. W. (2000) Systematics of Ruscaceae / Convallariaceae: A combined morphological and molecular investigation. Bot. J. Linn. Soc. 135(1–2): 73–92. https://doi.org/ 10.1111/j.1095-8339.2000.tb02346.x Seberg, O. [& al. 2012], Petersen, G., Davis, J. I., Pires, J. C., Stevenson, D. W., Chase, M. W., Fay, M. F., Devey, D. S., Jørgensen, T., Sytsma, K. J. & Pillon, Y. (2012) Phylogeny of the Asparagales based on three plastid and two mitochondrial genes. Amer. J. Bot. 99 (5): 875–889, ills. https://doi.org/10.3732/ajb.1100468 Steele, P. R. [& al. 2012], Hertweck, K. L., Mayfield, D., Mckain, M. R., Leebens-Mack, H. & Pires, J. C. (2012) Quality and quantity of data recovered from massively parallel sequencing: Examples in Asparagales and Poaceae. Amer. J. Bot. 99(2): 330–348. https://doi. org/10.3732/ajb.1100491 Stevenson, D. W. (1980) Radial growth in Beaucarnea recurvata. Amer. J. Bot. 67(4): 476–489. https://www. jstor.org/stable/2442287 Trelease, W. (1911) The desert group Nolineae. Proc. Amer. Philos. Soc. 50(200): 404–443, tt. 1–17. https://www. jstor.org/stable/983932 Walter, H. (1931) Die Hydratur der Pflanze und ihre physiologisch-ökologische Bedeutung. (Untersuchungen über den osmotischen Wert). Jena (DE): Verlag von Gustav Fischer.

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Beaucarnea Lemaire (Ill. Hort. 8(Misc.): 59, 1861). Type: Beaucarnea recurvata Lemaire. — Nolinoideae — Lit: Trelease (1911: synopsis); Hernández-Sandoval (1993: conservation); Lott & García-Mendoza (1994: Fl. Mesoamericana); Walker (2001: introduction); HernándezSandoval & al. (2012: management and conservation); Rivera-Lugo & Solano (2012: Fl. Tehuacán-Cuicatlán); Rojas-Piña & al. (2014: phylogeny, checklist, key); Martínez & al. (2014: leaf anatomy); Walker (2015: update). Distr: Mexico, C America (Guatemala, Belize, Honduras, Nicaragua?). Etym: For Monsieur Beaucarne (fl. 1861), Belgian succulent plant grower and notary from Eename near Audenarde, who first collected flowers of Beaucarnea recurvata. Incl. Pincenectitia hort. ex Lemaire (1861). Type: not typified. Incl. Calibanus Rose (1906). Type: Dasylirion caespitosum Scheidweiler. Hermaphrodite, dioecious or polygamodioecious shrubs to trees 4 at maturity, sessile or reduced, grouped at the stem apex; L erect, recurved and reflexed from above the base, linear, long, acuminate, 1.1–1.2 m  0.7–0.9 cm in the middle, glaucous-green, scarcely papillate, margins denticulate, teeth translucent, leaf base deltate; Inf paniculate, ovoid, 1.5–1.7 m; primary Br 28–30 cm, shorter Br 16–18 cm; Bra subtending the branches linear, 23–29 cm; Fl normally unisexual; male Fl Tep oblong, elliptic to orbicular, concave, 1.8–2.75  1.5–2 mm, OTep slightly smaller than the ITep; St 6, exserted, 2–3 mm, Anth dorsifixed, Ov reduced; female Fl Tep elliptic, orbicular to oblate, concave, 1.2–2 mm; Ov 3-carpellate, 2 ovules per carpel, unilocular, ellipsoid, 6-angled at the base; Sty  0.7 mm, Sti lobes tuberculate; Fr depressed globose, 4.5–5  4–4.5 mm, yellow with purple spots when mature, costae reddish; Se melonshaped, 2.5–4.5 mm ∅, testa rugose, red-brown. In the cladogram of Rojas-Piña & al. (2014) B. glassiana forms a “Calibanus clade” with B. compacta and B. hookeri, characterised by having subglobose caudices and greatly reduced branching. B. glassiana is closest to B. hookeri from which it differs in its general habit with fewer apical branches, significantly longer glaucous-green leaves, long exserted, wellbranched inflorescences, and smaller depressed globose fruits with reddish costae. Discovered in 1996 around Xichú by Charlie Glass, intrepid botanical explorer of Mexico and former editor of the Cactus & Succulent Journal (US). His collection later flowered in cultivation enabling confirmation of its generic placement. B. goldmanii Rose (Contr. US Nation. Herb. 12: 261, t. 20, 1909). Type: Mexico, Chiapas (Goldman 887 [US]). — Lit: Lott & García-Mendoza (1994); Rojas-Piña & al. (2014). Distr: SW Mexico (Chiapas), Guatemala; rocky slopes in deciduous forests, 800–1400 m. Stem to 10 m, slender, branched above, bark thin, rough; L linear, to 100  1.5 cm, strongly recurved, shiny, smooth, long acuminate towards the apices, margins minutely serrulate with

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appressed denticles; Inf compound-paniculate, nearly sessile, primary divisions 15–20 cm, Br few, to 10 cm; female Inf Bra linear-filiform, 19–21  1 cm; bracteoles 1–3 mm; Ped articulated; Tep  2 mm; Fr ellipsoid, glaucous, broadly 3-winged, very large, 18–20  12–15 mm; Se 5–5.5  5–5.5 mm, rugose, blackish, opaque. Resembles B. guatemalensis, but the leaves are large and the fruits are narrower and glaucous. It is especially poorly known and probably not in cultivation. B. gracilis Lemaire (Ill. Hort. 8(Misc.): 61, 1861). Type [neo]: Mexico, Puebla (Hernández S. 2354 [MEXU, TEX, UAT]). — Lit: Cardel & al. (1997: ecology); Rivera-Lugo & Solano (2012); Rojas-Piña & al. (2014). Distr: Mexico (Puebla, Oaxaca: Endemic to the Tehuacán-Cuicatlán valley); thorn scrub, 1300–2000 m. I: Cact. Succ. J. (US) 48: 64, 1976; Rowley (1987: 89, as Nolina); Walker (2001: 217, fig. 4). – Fig. 1.  Dasylirion gracile (Lemaire) J. F. Macbride (1918) (nom. illeg., Art. 53.1)  Nolina gracilis (Lemaire) Ciferri & Giacomini (1950); incl. Nolina histrix hort. (s.a.) (nom. inval., Art. 29.1); incl. Pincenectitia gracilis hort. ex Lemaire (1861) (nom. inval., Art. 34.1c?); incl. Beaucarnea oedipus Rose (1906). Dioecious tree, stem basally enormously swollen, circular in cross-section, variously and irregularly branched, to 12 m overall; L linear, erect, 30–60  0.4–0.7 cm, very glaucous, margins minutely but sharply serrulate-scabrous; Inf short-stalked, ovoid- or oblong-paniculate, 0.6–1 m, primary divisions to 30 cm, Br rather weak, to 14 cm; Ped 3.2–3.5 mm; Tep 1.2  1 mm; Fr round-ellipsoid, openly notched at tip and base, 10  7–9 mm; Se smooth, 3.6  2.8 mm. Morphologically similar to B. compacta, but differing in the elongated (instead of subglobose) stem, longer inflorescence branches, and larger flowers and fruits. It has a limited distribution in the Tehuacán-Cuicatlán Valley, Puebla/Oaxaca (Hernández-Sandoval & Zamudio 2003; RiveraLugo & Solano 2012). García-Franco & al. (1995) record a relatively high population density in the

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Fig. 1 Beaucarnea gracilis. (Copyright: J. Etter & M. Kristen)

valley of Zapotitlán de las Salinas (Puebla) with a sex ratio close to 1:1. A rare occurrence of parasitism by Psittacanthus calyculatus (Loranthaceae) is also reported. Cardel & al. (1997) recorded that plants with stem diameter < 0.7 m were absent from their study site (n = 404) and observed that low recruitment, removal of seedlings for commerce, grazing by goats and land clearance for corn cultivation, highway construction and urbanization, threaten the establishment and survival of seedlings. The leaves are used to make hats and arrangements for religious offerings (Rivera-Lugo & Solano 2012). B. guatemalensis Rose (Contr. US Nation. Herb. 10: 88, Fig. 1, 1906). Type: Guatemala (Kellermann 4320 [US, F, LL, MEXU, MICH, TEX, UC]). — Lit: Lott & García-Mendoza (1994); Rojas-Piña & al. (2014). Distr: Guatemala; rocky slopes, ecotone between deciduous forest to oak-pine forests, 700–1600 m.  Nolina guatemalensis (Rose) Ciferri & Giacomini (1950). Trees 6–12 m with thickened bulbous base, abruptly contracted into a slender stem often with slender multiple Br; L linear, to 100  2.5–3 cm, minutely scabrous, with papillose ridges, margins minutely papillate; Inf shortstalked, broadly ovoid-paniculate, primary divisions 30 cm, Br rather spreading, to 15 cm, branchlets to 6 cm; Bra 15 cm, lanceolate-linear; bracteoles 10–12 mm; male Fl Tep 3 mm; female

Fl Ped 8 mm, articulated above the middle; Fr ellipsoid-obovate,15–18  13–15 mm, openly notched at the tip for 4–5 mm; Se irregularly 3-lobed, 5 mm diam. Related to B. pliabilis, but the fruit has broader wings and the leaves are broader with a scabrous as opposed to smooth surface. B. hiriartiae L. Hernández (Acta Bot. Mex. 18: 25–29, ills., 1992). Type: Mexico, Guerrero (Hernández-Sandoval & Martínez 1629 [MEXU, TEX, UAT]). — Lit: Rojas-Piña & al. (2014). Distr: Mexico (Guerrero: Río Balsas drainage); deciduous tropical forest, in canyons on very steep slopes of shale or limestone, 250–700 m; reported to have two flowering and fruiting periods. Trees to 8 m; stem basally swollen, oval in cross-section, often with slender multiple Br; L recurved, linear, to 70–90  1–1.5 cm, pale green, concave, papillate, margin minutely serrulate; Inf short-stalked, ovoid-elliptically paniculate, primary divisions 20–25 cm, Br rather spreading, 3–10 cm; Ped 5.5–6.5 mm; Tep ovate, 2.5  2 mm; Fr obovate-oblong, 8–11  7–10 mm; Se ellipsoid-obovate, 3.5  3.1 mm. Considered to be closely related to B. stricta from neighbouring Oaxaca in the protologue, but appearing to differ only quantitatively in the length of organs such as leaves and pedicels. It differs though in that the stem base in cross-section is oval and the leaves are recurved. Hernández-

Beaucarnea RUSCACEAE

Sandoval (2001) compared B. hiriartiae with B. recurvata and B. sanctomariana. He recorded a unique stem architecture for this species, in that meristems of a single branch develop apical dominance, leaving the remaining branches with reduced growth rates. Consequently, the habit of B. hiriartiae resembles a tree with its branches growing only to one side. This feature alone enables this species to be distinguished easily in the field from B. recurvata and B. sanctomariana. B. hiriartiae has papillate leaves, in contrast to the smooth ones of the other two species (Hernández-Sandoval 2001). Vernacular names: “Izote Delgado”, “Ixtotl”. This species is probably not in cultivation. B. hookeri (Lemaire) Baker (J. Bot. 10: 327, 1872). Type: K (ex cult. Kew 1873). — Lit: Hernández-Sandoval & Zamudio (2003); Rojas-Piña & al. (2014). Distr: Mexico (Hidalgo, San Luis Potosí); apparently well-camouflaged on hilltops. I: Glass & Foster (1970: 269); Rowley (1987: 32, 90); Brand (2007); Spee (2007). – Fig. 2.  Dasylirion hookeri Lemaire (1859)  Calibanus hookeri (Lemaire) Trelease (1911)  Nolina hookeri (Lemaire) G. D. Rowley (1990); incl. Dasylirion hartwegianum Hooker (1859) (nom. illeg., Art. 53.1); incl. Dasylirion caespitosum Scheidweiler (1861)  Calibanus caespitosus (Scheidweiler) Rose (1906); incl. Dasylirion flexile K. Koch (1867). Fig. 2 Beaucarnea hookeri. (Copyright: T. Wardhaugh)

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Caudex globose, slightly flattened, to 50 cm ∅, bark thick, corky and fissured, later woody, with numerous crowns of leaves from very short monocarpic Br not elevated over the surface of the caudex; L thin, grass-like, wiry, narrowly linear, somewhat concave and keeled, to 30 cm  2–3 mm, margins serrulate-scabrous, tip entire; Inf simple panicles, normally unisexual or rarely with some hermaphrodite flowers, to 25 cm, shorter than the leaves, shortly pedunculate, lax with thin spreading Br 6–8 cm; Bra scarious, ovate or lanceolate, much shorter than the subtended branches; Tep elliptic-obovate, 2–3  1.5–2 mm, scale-like, dirty whitish-yellow, translucent; St subequal, 2.5–3 mm; Anth medifixed, 1.2–1.5 mm; Ov ovoid, 2.5–3  2 mm, 3-locular with 1 ovule in each locule; Sti sessile, 3-lobed; Fr globose-ovoid, 8–9  6–7 mm, pale straw-brown, indehiscent; Se melon-shaped, 3–4  3 mm. — Cytology: 2n = 38 (Johnson & Gale 1983). B. hookeri was first discovered around 1845. It was not described until 1859 when it was misidentified as Dasylirion hartwegianum by Sir William Hooker at Kew. Rose (1906) established the monotypic genus Calibanus for this species, named for Shakespeare’s monster Caliban from the play “The Tempest”, presumably for the massive caudex. The plant is currently well known in cultivation as Calibanus hookeri. Most recently, Rojas-Piña & al. (2014) provided convincing molecular evidence that Calibanus is nested within Beaucarnea and hence not distinct at the

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generic level. B. hookeri is closely related to B. compacta and B. glassiana, characterised by having subglobose caudices and greatly reduced branches. B. hookeri was rediscovered by Charlie Glass and Bob Foster in 1968 in San Luis Potosí, where it appears to be well camouflaged in habitat (Glass & Foster 1970: 269, ill.). Most of the material now widespread in cultivation originates from this introduction (ISI 688), since the plant has taken well to cultivation (Walker 2001). It has proved to be almost hardy, and easily propagated from seed. Johnson & Gale (1983) report successful artificial pollination with one-year old refrigerated pollen. Brand (2007) reports that seed-raised plants first flowered after 9 years. B. hookeri is a remarkable caudiciform, eventually developing a large woody caudex, curious rather than attractive. It is known locally in Mexico as “Sacamecate” and its leaves are used for thatching and for scouring dishes. B. olsonii V. Rojas & L. O. Alvarado (Phytotaxa 286(1): 15–18, ills., 2016). Type: Mexico, Puebla (Olson & al. 1044 [MEXU, MEXU, MO]). — Distr: Mexico (Puebla: Izúcar-Acatlán); dense tropical deciduous forest on steep slopes, 1300 m. Trees 3–5 m; stem basally massively swollen, 2.4–4 m ∅, abruptly tapering into few slender Br 1–3 m long, dark grey and squamose forming irregular grooves along the length of the stem; L straight and terminally erect, 60–85 cm  0.9–1.7 cm at the middle, green to bluish glaucous green, canaliculate, papillate, margin slightly irregular; Inf a thyrse with 3 orders of branching, last unit a reduced rhipidium, Bra narrowly triangular to triangular, apex acuminate to long caudate; female Fl with Ped 4–5 mm, articulated at of the length below the flower; Tep ovate, whitish 1.5–1.7  1.3–2 mm; male Fl unknown; Fr (immature) obovate, 3-winged, 7–9 mm; Se (immature) 1 per fruit, brown, testa rugose. Similar to B. hiriartiae but differs in its tessellated bark, straight leaves and short style. It is easily distinguished from other Beaucarnea species by its swollen base that abruptly tapers into a

C. C. Walker

slender stem and in its pedicel articulation that occurs at of the length of the pedicel below the flower. B. olsonii is a very rare, localised species, known only from its type locality in SW Puebla with an extent of occurrence estimated to be 1 m, nearly sessile, broadly ovoid-paniculate, primary Br to 30 cm, lower Br to 15 cm, branchlets 5 cm; male Fl Ped articulated above the middle; Tep 2–2.5 mm; female Fl Tep 2.5–4 mm; Fr ellipsoid to slightly obovate, 12–14  9–10 mm, pale yellow, tip and base emarginate, apical notch 1–2  1–2 mm; Se ellipsoid to ovoid, 3.5–4.5  3–4 mm, lobes not prominent, testa rough to smooth, bright red-brown. Similar to B. sanctomariana, but differing in the larger trunk bases, longer leaves, and larger ellipsoid fruits. Both species occur in the Isthmus of Tehuantepec, Oaxaca, where B. recurvata is restricted to the Pacific slopes, whereas B. sanctomariana occurs on the side facing the Gulf of Mexico (Hernández-Sandoval 2001). Most populations of B. recurvata are in a critical condition, resulting from the continued fragmentation and habitat destruction, caused by the expansion of agriculture, livestock grazing, removal of firewood, timber and urban expansion (Contreras & al. 2008). However, B. recurvata is readily grown from seed and therefore now extremely common in cultivation, and widely distributed through the commercial nursery trade. It retains its immature form of an almost globular caudex for many years before the stem tip starts to elongate, only branching after flowering. As a young plant, therefore, with the long narrow leaves emerging from the globose base it makes an unusual and attractive pot plant. First flowering of 15 year old pot-grown specimens has been reported at 1 m tall and  20–25 cm basal diameter. At maturity it has a rather different form, basally swollen with a tapering main stem and a few terminal branches. A variegated cultivar is also sometimes encountered in cultivation and was named ‘Stripy Ponytail’ (Walker 2015). Vernacular name, especially in the USA: “ponytail palm”. B. sanctomariana L. Hernández (Novon 11 (1): 50–53, ills., 2001). Type: Mexico, Oaxaca

C. C. Walker

(Wendt 5914 [MEXU (†?), CHAPA, LL (†?), MO (†?), UAT]). — Lit: Rojas-Piña & al. (2014). Distr: Mexico (Oaxaca); restricted to xerophytic vegetation on steep karstic rock outcrops that emerge from the surrounding tropical deciduous forest, 200–250 m. Tree 4–7 m, stem profusely branched, base widely conical, 1–1.5 m ∅, bark fissured with rectangular to polyhedral pattern, brown-grey; Br elongate, slender, with leaf Ros spherical to widely ovoid; L linear, recurvate, very long acuminate, 75–85  1.2–1.6 cm, smooth, bright green, margin pale green, denticulate; Inf ovoid to largely ellipsoid, 0.8–1 m  1 cm ∅ at the base; primary Br 10–15 cm, shorter Br 3–8 cm; male Fl fragrant; Ped 1.5–2.3 mm, articulated below the flower; Tep cream-yellow, ovate, 1.4–1.8 mm; female Fl Ped 1.5–2 mm, articulated at or above the middle; Tep cream-yellow, ovate, 1–1.5 mm; Fr ellipsoid to almost globose, 6–8  5–7 mm, pale yellow at maturity; Se (immature) with apices of lobes very irregular, testa rugose, red-brown. Morphologically similar to B. hiriartiae and B. recurvata. It shares with B. hiriartiae the recurved leaves, the undulate bracts, and the general colour of the inflorescence. It shares with B. recurvata the shape of the trunk base, and the recurved smooth leaves. Distinguishing features of B. sanctomariana are the short inflorescence branches and the small flowers and fruits. Its very limited distributional range characterises it as an endangered micro-endemic species. B. stricta Lemaire (Ill. Hort. 8(Misc.): 61, 1861). Type [neo]: Mexico, Oaxaca (Hernández S. 2391 [MEXU, TEX, UAT]). — Lit: Matuda (1960); Rivera-Lugo & Solano (2012); Rojas-Piña & al. (2014). Distr: Mexico (Oaxaca); tropical deciduous forest on acid soils of volcanic origin, 1200–1500 m. I: Lyons (1969: 53–54); Walker (2001: 218, fig. 5). – Fig. 5.  Beaucarnea recurvata var. stricta (Lemaire) Baker (1880)  Dasylirion strictum (Lemaire) J. F. Macbride (1918)  Nolina stricta (Lemaire) Ciferri & Giacomini (1950); incl. Pincenectitia glauca Lemaire (1861) (nom. inval., Art. 34.1c)  Beaucarnea glauca (Lemaire) Hereman (1868) (nom. inval., Art. 34.1c).

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This is another tall tree, with a stem greatly swollen and circular in cross section at the base. Its specific name refers to the erect, straight leaves. It has a limited distribution in Oaxaca.

References

Fig. 5 Beaucarnea stricta. (Copyright: U. Eggli)

Trees to 10 m; stem greatly swollen, circular in cross-section at the base, irregularly and modestly branched, bark corky, fissured; L linear, erect, to 55–80  0.8–1.5 cm, stiffly spreading, keeled, pale or glaucous grey, margin yellowish, usually minutely serrulate-scabrous; Inf shortstalked, ovoid-paniculate, primary divisions 20 cm, Br short, the lower with branchlets to 3 cm; Ped 2.8–3 mm; Tep 2.5  2 mm; Fr broadly ellipsoid, openly notched at tip and base, 12  8–10 mm; Se irregularly 3-lobed, smooth, 3.6  3.3 mm. In the cladogram of Rojas-Piña & al. (2014), B. stricta occupies a distinct clade, separate from B. purpusii, that was until recently included as a synonym of B. stricta. B. stricta is morphologically similar to B. purpusii, B. hiriartiae and B. gracilis, but is distinguished by the septal and distal nectaries; B. stricta also differs from B. gracilis in its larger leaves and bright yellow-green inflorescences (Rivera-Lugo & Solano 2012).

Brand, T. (2007) Calibanus hookeri in Blüte — leichter als gedacht? Kakt. and. Sukk. 58(10): 273–276, ills. Cardel, Y. [& al. 1997], Rico-Gray, V., García-Franco, J. G. & Thien, L. B. (1997) Ecological status of Beaucarnea gracilis, an endemic species of the semiarid Tehuacan Valley, Mexico. Conservation Biol. 11(2): 367–374. https://doi.org/10.1046/j.1523-1739.1997.95322.x Contreras, A. [& al. 2008], Osorio, M. L., Equihua, M. & Benítez, G. (2008) Conservación y approvechamiento de Beaucarnea recurvata, especie forestal no maderable. Cuad. Biodivers. 28: 3–9. García-Franco, J. [& al. 1995], Rico-Gray, V. & PalaciosRios, M. (1995) Parasitismo de Psittacanthus calyculatus (Loranthaceae) sobre Beaucarnea gracilis (Nolinaceae) en el Valle de Zapotitlán de las Salinas, Puebla, México. Cact. Suc. Mex. 40(3): 62–65, ill. Glass, C. & Foster, R. (1970) Mexican logbook. Cact. Succ. J. (US) 42(2): 73–77, (3): 107–112, 142, (4): 174–178, (5): 229–234, (6): 263–269. Hernández-Sandoval, L. G. (1993) Beaucarnea, un genero amenazado ? Cact. Suc. Mex. 38(1): 11–13, ill. Hernández-Sandoval, L. G. (2001) Beaucarnea sanctomariana (Nolinaceae), a new micro-endemic species of Ponytail Palm from the Isthmus de Tehuantepec, Oaxaca, Mexico. Novon 11(1): 50–53, ills. https://www.jstor.org/stable/3393207 Hernández-Sandoval, L. G. & Zamudio, S. (2003) Two new remarkable Nolinaceae from Central Mexico. Brittonia 55(3): 226–232, ills. https://www.jstor.org/sta ble/3218443 Hernández-Sandoval, L. G. [& al. 2012], Osorio Rosales, M. L., Pérez Farrera, A., Contreras Hernández, A., Malda Barrera, G., Espadas Manrique, C., Almanza Rodríguez, K. E., Castilloa Gómez, H. A. & Félix Álvarez, A. (2012) Manejo y conservación de las especies con valor comercial de Pata de Elefante (Beaucarnea). Querétaro (MX): Ediciones Universitarias, Universidad Autónoma de Querétaro. Johnson, M. A. T. & Gale, R. M. O. (1983) Observations on the leaf-anatomy, pollen, cytology and propagation of Calibanus hookeri (Lem.) Trelease. Bradleya 1: 25–32, ills. https://doi.org/10.25223/brad.n1.1983.a4 Lott, E. J. & García-Mendoza, A. (1994) Agavaceae. In: Davidse, G. & al. (eds.): Flora Mesoamericana, Volumen 6, Alismataceae a Cyperaceae; pp. 35–47, keys. México (MX): UNAM, Instituto de Biología / St. Louis (US): Missouri Botanical Garden / London (GB): Natural History Museum. https://www.tropicos. org/Project/FM

1328 Lundell, C. L. (1939) Studies of Mexican and Central American plants. VIII. Bull. Torrey Bot. Club 66(9): 583–604. https://doi.org/10.2307/2480801 Lyons, G. W. (1969) The Huntington Desert Garden today. Cact. Succ. J. (US) 41(2): 51–56, (3): 120–123, (4): 151–154, (5): 221–224, ills. Martínez, M. [& al. 2014], Hernández-Sandoval, L. & Carrillo, L. (2014) Foliar anatomy of Beaucarnea Lemaire (Nolinaceae s.s.). Pl. Syst. Evol. 300(10): 2249–2258, ills., key. https://doi.org/10.1007/s00606014-1048-2 Matuda, E. (1960) Beaucarnea stricta. Cact. Suc. Mex. 5 (4): 93–94, ill. Rivera-Lugo, M. & Solano, E. (2012) Flora del Valle de Tehuacán-Cuicatlan. Fascículo 99. Nolinaceae Nakai. México D.F. (MX): Instituto de Biología UNAM. Rojas-Piña, V. [& al. 2014], Olson, M. E., AlvaradoCárdenas, L. O. & Eguiarte, L. E. (2014) Molecular phylogenetics and morphology of Beaucarnea (Ruscaceae) as distinct from Nolina, and the submersion of Calibanus into Beaucarnea. Taxon 63(6): 1193–1211, ills., key. https://doi.org/10.12705/636.9 Rose, J. N. (1906) Studies of Mexican and Central American plants. No. 5. Liliaceae. Dasylirion and its allies. Contr. US Nation. Herb. 10(3): 87–92, tt. 23–25. https://www. biodiversitylibrary.org/page/41739857#page/193

C. C. Walker 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. Spee, B. (2007) Calibanus hookeri, geen plant om mee te nemen. Succulenta 86(3): 123–126, ills. Thiede, J. (2012) Nomenclatural status of unranked names published by Trelease (1911) in Beaucarnea, Dasylirion, and Nolina (Asparagaceae — Nolinoideae). Phytoneuron 2012(77): 1–4. https://www.phytoneuron. net/77PhytoN-Beaucarnea.pdf? Trelease, W. (1911) The desert group Nolineae. Proc. Amer. Philos. Soc. 50(200): 404–443, tt. 1–17. https://www.jstor.org/stable/983932 Vadillo-Pro, M. C. [& al. 2016], Hernández-Sandoval, L., Malda-Barrera, G., Osorio-Rosales, M. L. & MataRosas, M. (2016) In vitro regeneration from longitudinal sections of seedlings of Beaucarnea purpusii Rose, an endemic and endangered species. HortScience 51(3): 279–284, ills. Walker, C. C. (2001) Introducing the Nolinaceae. Brit. Cact. Succ. J. 19(4): 214–221, ills. Walker, C. C. (2015) Beaucarnea updated. CactusWorld 33(4): 267–272, ills.

Dasylirion RUSCACEAE C. C. Walker

Dasylirion Zuccarini (Allg. Gartenzeitung 6(33): 258, 1838). Type: Dasylirion graminifolium Zuccarini. — Nolinoideae — Lit: Trelease (1911: synopsis); Bogler (1994a: systematics); Bogler (1994b: phylogeny); Bogler (1995: sytematics, phylogeny); Hochstätter (2011: synopsis); Hochstätter (2012: synopsis). Distr: S USA to Mexico. Etym: Gr. ‘dasys’, dense, rough, shaggy; and Gr. ‘leirion’, lily; presumably for the long and untidy appearance of the leaves. Incl. Dasylirium Lemaire (1865) (nom. inval., Art. 61.1). Dioecious, short arborescent perennial shrubs with thick unbranched stems crowned with dense rosettes, sometimes rosettes  stemless; L serrulate or prickly, or unarmed, linear, hard and fibrous; Inf paniculate, elongate, conspicuously bracteate, to 6 m; Ped jointed at the base of the flowers; Per small, whitish, persistent; St 6, vestigial in female flowers; Ov 1-celled, ovules 2 or 3 but only 1 developing; Sty very short, erect; Sti 3; Fr 1-seeded (occasionally 2-seeded) small capsules, 3-winged, thin-walled. — Cytology: 2n = 38 (Satô 1935, 1942; Gómez-Pompa & al. 1971).

C. C. Walker (*) School of Environment, Earth and Ecosystem Sciences, The Open University, Milton Keynes, England e-mail: [email protected]

A genus of 20 species, many poorly understood. All are xerophytic shrubs and not truly succulent, but some are sometimes cultivated in succulent plant collections. Several species, commonly known by the Indian vernacular name “Sotol”, have various uses. Their stems were used for building and for fuel. The leaves are trimmed off and the remaining stump is roasted or boiled. The heads are often baked in pits dug in the ground. The roasted stems are also allowed to ferment to produce an alcoholic beverage. The leaves are much used for thatching, mats, baskets, rough hats etc., and their fibres for rough cordage. Dried and varnished expanded leaf bases, called “Desert Spoons”, are widely used in flower arrangements etc. Trelease (1911) divided the genus into 2 unranked groups: Eudasylirion and Quadrangulatae. Krause (1930) cited “Dasylirion Sek. II Quadrangulatae Trelease” with full reference, and this was accepted by Thiede (2012d) as a valid, albeit unintended, new combination. Hochstätter (2011) published an additional sect. Glaucophyllum for some species previously placed in sect. Dasylirion. This sectional classification is adopted here: [1] Sect. Dasylirion (= Eudasylirion Trelease 1911, nom. inval. ICN Art. 21.3): L 2-edged, usually somewhat concave and irregularly keeled, margins prickly and usually rough with minute intervening denticles. — 9 species.

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_26

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[2] Sect. Quadrangulatae (Trelease) Kraus 1930: L 4-sided, unarmed. — 4 species. [3] Sect. Glaucophyllum Hochstätter 2011: L waxy-glaucous. — 9 species. Only selected species of these xerophytes are dealt with here, namely those that have a long history, are of particular interest or are widely distributed. Many species not treated here are either very localised and rare or recently recognised. The following name is of unresolved application but is referred to this genus: Dasylirion robustum hort. (s.a.) (nom. inval., Art. 29.1). D. acrotrichum (Schiede) Zuccarini (Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 3: 226, 228, t. 1: Fig. 4, 1840). Type: Mexico, Puebla (Schiede s.n. [B †?]). — Distr: Mexico (Hidalgo, Puebla, Querétaro, San Luis Potosí); dry rocky slopes, in grassland or among shrubs or in oak-pine forest or open oak woodland, sometimes on limestone, 1800–2600 m. I: Curtis’s Bot. Mag. 84: t. 5030, 1858; McVaugh (1989: 175, Fig. 28); Hochstätter (2011: 115).  Yucca acrotricha Schiede (1829); incl. Roulinia gracilis Brongniart (1840)  Barbacenia gracilis (Brongniart) Brongniart (1840) (nom. inval., Art. 34.1c)  Bonapartea gracilis (Brongniart) Otto (1841) (nom. inval., Art. 34.1c)  Yucca gracilis (Brongniart) Otto (1841) (nom. illeg., Art. 53.1)  Dasylirion gracile (Brongniart) Zuccarini (1845)  Littaea gracilis (Brongniart) Verschaffelt (1864). [1] Stem to 1 m; outer L recurved, linear, green and glossy or somewhat glaucous and dull, 60–100  0.9–1.8 cm, tip split into 20–30 spreading fibres, margin distinctly and finely toothed between the prickles, prickles 5–10 ( 15) mm apart, to 2 mm long, rather straight, pale yellowish with slightly brown tips; Inf paniculate, 3–5 m; Bra ovate, entire; Tep 2–3 mm; Fr round-cordate, shallowly notched, wings broadening upwards, 8–9  6–7 mm; Se 3.5  3 mm. — Cytology: 2n = 38 (Satô 1942). McVaugh (1989) argued that the correct spelling of the epithet should be ‘acrotriche’, since this

C. C. Walker

is seemingly a bastard word, neither a noun in apposition to the generic name nor a proper Latin adjectival form. In contrast, most authors since Zuccarini transferred this species to Dasylirion from Yucca, have used the form ‘acrotrichum’, and this usage is here followed. This species is widely distributed in S-C Mexico. Plants are robust, short- to tall-stemmed. Leaves characteristically have a matt or dull surface with coarsely brushy tips (Hochstätter 2011). D. glaucophyllum Hooker (Curtis’s Bot. Mag. 84: t. 5041 + text, 1858). Type: K. — Distr: E-C Mexico (Hidalgo); dry rocky slopes, 1900–3000 m. I: Ríha & Subik (1981: Fig. 225); Hochstätter (2012: 162–163). Incl. Bonapartea glauca hort. (s.a.) (nom. inval., Art. 29.1); incl. Dasylirion glaucum Carrière (1872). [3] Stem short; L linear, acuminate, intensely glaucous, bluish-green, 60–120  1.2–1.8 cm, tip entire, marginal prickles 1–2 mm long, horny, deep yellow, margin finely toothed between the prickles; Inf densely paniculate, to 1.2 m; Tep greenish-white, red-tipped, 2 mm; Fr subellipsoid, 9–10  6 mm; Se 4  2.5 mm. This species was described by Hooker (1858) based on a single staminate specimen that flowered at Kew in the unusually warm summer of 1857. Hooker observed that, compared to D. acrotrichum, the leaves of his new species were held more erect and rigid, were very glaucous and the tips were not broken into tufts or brushes. This species is the type of the new sect. Glaucophyllum Hochstätter. It develops a robust short stem, with characteristic waxy-glaucous leaves. It is apparently very rare, and geographically isolated in E-C Mexico. D. graminifolium (Zuccarini) Zuccarini (Allg. Gartenzeitung 6(33): 259, 1838). Type: not typified. — Distr: Mexico (San Luis Potosí). I: Hochstätter (2011: 106).  Yucca graminifolia Zuccarini (1837). [1] Stem short, 6 m, 25 cm ∅, basally swollen, modestly branched; L linear, tapering to 120  4 cm, pale green, very flexible, with slightly curling tip 15 cm long, dead leaves forming a dense covering of the branches below the terminal rosette; Inf paniculate, >2.3 m tall, elongate, branched, primary Br 15–30 cm; Fl 20 mm ∅, white; Fr and Se not known. One of the largest and hardiest members of the genus. It is adapted to alpine conditions, where temperatures below 15 C are not infrequent. It grows sympatrically with N. nelsonii, but no hybrids have yet been observed (Hochstätter & Donati 2010). N. interrata Gentry (Madroño 8: 181, fig. 1, t. 1, 1946). Type: USA, California (Gentry 7330 [SD]). — Distr: USA (S California: San Diego County), Mexico (N Baja California); rocky hillsides, chaparral, 200–700 m; very rare. I: Hochstätter (2010: 44–45). [4] Plants with underground branching rhizome to 3 m  30 cm ∅, above-ground stems

C. C. Walker

not obvious, mature Ros with < 45 L; L linear, glaucous, 70–100  0.8–1.5 cm, base barely expanded, tip dry, slender, not filiferous, margin minutely and persistently serrate, armed with denticles of 2 sizes; Inf open compound panicles 1.5–2 m, base of peduncle 0.5–1.8 cm ∅; Bra persistent, 20–40 cm; female Fl with staminodes inserted on the Tep; Fr large, broader than long, 12–15 mm wide; Se reddish-brown to yellowish, wrinkled, 5  4 mm. Distinguished by the glaucous leaves with coarse armature. The horizontal underground stem or rhizome is especially noteworthy, but this structure may be present in other species assumed to be acaulescent, since this feature may be readily overlooked. Known locally in San Diego County as “Dehesa Nolina” and “Dehesa Beargrass”. N. interrata is very rare and hence one of the most endangered plant species in California, occurring in only two small stands. The populations are stable but are slowly being affected by land clearance for housing development. Three additional populations are known in Baja California. N. matapensis Wiggins (Contr. Dudley Herb. 3: 65, 1940). Type: Mexico, Sonora (Wiggins 7515 [DS]). — Distr: Mexico (Sonora, Chihuahua); mountain woodland, gravelly slopes, 900–1700 m. I: Walker (2001: 221, figs. 11–12); Hochstätter (2010: 40). [4] Small trees, stem to 3–10 m, basally swollen, simple or modestly branched, bark fissured; Ros small; L linear, 70–120  1–1.5 cm, recurved, hard, striate, greenish-yellow, somewhat glaucous, margin finely serrulate; Inf paniculate, 1.5–3  longer than the L, primary Br 15–35 cm, branchlets 10–15 cm; Tep oblonglinear, 2 mm; Fr broadly ellipsoid, depressed, deeply notched at both ends, tardily dehiscent, 8–9  5 mm; Se pale brown, nearly globose, 2.5–3 mm ∅. This species has a narrow distribution range. The leaves are used for basket-making, the stems for posts. A relatively unknown species, first introduced into cultivation in 1976 as ISI 982. Plants from this introduction are now multiheaded specimens at the Huntington Botanical

Nolina RUSCACEAE

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Garden. The species is recommended as an attractive landscape plant for subtropical areas such as S California. Local vernacular names: “Palmito”, “Tuya”. N. nelsonii Rose (Contr. US Nation. Herb. 10: 92, 1906). Type: Mexico, Tamaulipas (Nelson 4489 [US]). — Distr: Mexico (Tamaulipas), dry bush, 1200–2500 m. I: Hochstätter (2010: 39). [4] Tree, stem to 4.5 m, openly branched, with a large crown of stiff L forming small Ros; L rigid, erect to recurving, 50–70  3–4 cm, flexible, blue to blue-green, margin serrulate; Inf erect, paniculate, 2–3 m  20–60 cm; Bra scarious, lacerate; Fl 3 mm  3 mm ∅, cream; Fr papery, 8–10 mm ∅; Se globose to ovate, brownish, 2–3 mm ∅. This is a small tree similar to N. parryi, but the leaves differ in size and form. Superficially it looks like Yucca rostrata. Reported as frost hardy to 12  C. N. parryi S. Watson (Proc. Amer. Acad. Arts 14: 247, 1879). Type: USA, California (Parry s.n. [GH, MO]). — Lit: Mitich (1982); Hess & Dice (1995). Distr: USA (California); rocky slopes in desert and pinyon-juniper woodland, 900–2100 m. I: Hochstätter (2010: 30–31). – Fig. 3.  Nolina bigelovii var. parryi (S. Watson) L. D. Benson (1945)  Nolina bigelovii ssp. parryi (S. Watson) E. Murray (1983); incl. Nolina parryi ssp. parryi; incl. Nolina parryi ssp. wolfii Munz (1950)  Nolina bigelovii var. wolfii (Munz) L. D. Benson (1954)  Nolina wolfii (Munz) Munz (1974)  Nolina bigelovii ssp. wolfii (Munz) E. Murray (1983). [4] Arborescent, stem 1–2 (–6) m, basally branching, mature Ros with up to 200 L; L linear, 50–150  2–4 cm, almost pungent, rather thick, concave, keeled, margin serrulatescabrous, base strongly expanded; Inf to 4 m, with rather narrow Br 15–30 cm long and spreading, densely flowered branchlets < 4 cm; Bra large, papery, persistent; Fl large; Tep 4 mm, creamy-white; Fr very large, orbicular, deeply notched at both ends, 12–15 mm ∅; Se reddishbrown, 4  3 mm.

Fig. 3 Nolina parryi. (Copyright: U. Eggli)

N. parryi is differentiated from N. cismontana by the taller stems, larger number of leaves per rosette and broader leaf bases (Dice 1988, Hess & Dice 1995). It is endemic to California. Mitich (1982) records spectacular specimens in the Kingston Mountains, Mojave Desert. Here some individuals reach a height of over 6 m with stems 1 m ∅, and with leaves 1.5 m long; the plants are 4 m across. Inflorescences are enormous too, being 4 m long with the fertile portion 2 m in length. He reports that N. parryi grows readily from seed, requiring a mild climate for outdoor cultivation. In such conditions plants flower after 7–8 years. Common names: “Parry’s Nolina”, “Giant Nolina”. N. parviflora (Kunth) Hemsley (Biol. Centr.Amer., Bot. 3: 372, 1884). Type: Mexico, México (Humboldt & Bonpland 4031 [P]). — Lit: McVaugh (1989: 242–244, with ills.); RiveraLugo & Solano (2012); Ruiz-Sanchez & Specht (2013); Ruiz-Sanchez & Specht (2014). Distr:

1350

Mexico (Chihuahua?, Durango, Zacatecas, Nayarit, Jalisco, Michoacán, Querétero?, Hidalgo, Mexico, Puebla, Veracruz, Oaxaca); widely distributed in the pine forest, pine-oak-juniper forests, xerophytic scrub and tropical dry forest localities of the Trans-Mexican Volcanic Belt, Sierra Madre Occidental, Sierra Madre Oriental and Sierra Madre del Sur mountain ranges: steep slopes, mountainsides, barrancas, sandy or rocky soils, 1300–2400 m. I: Rowley (1990: figs. 1–2, 4, as N. longifolia); Folsom & al. (1995: 104–105, as N. longifolia); Hochstätter (2010: 36; 41–42, as N. longifolia).  Cordyline parviflora Kunth (1815)  Dracaena parviflora (Kunth) Willdenow (1829)  Beaucarnea parviflora (Kunth) Baker (1872); incl. Yucca longifolia Karwinsky ex Schultes fil. (1830)  Dasylirion longifolium (Karwinsky ex Schultes fil.) Zuccarini (1840)  Beaucarnea longifolia (Karwinsky ex Schultes fil.) Baker (1872)  Nolina longifolia (Karwinsky ex Schultes fil.) Hemsley (1884); incl. Roulinia humboldtiana Brongniart (1840); incl. Roulinia karwinskiana Brongniart (1840); incl. Dasylirion humboldtii Kunth (1844); incl. Nolina altamiranoana Rose (1905); incl. Nolina elegans Rose (1906). [4] Arborescent, stem 1–3 (rarely >5) m, swollen at base, with 1–6 Br, bark thick, corky and rough, with a large crown forming Ros of several hundred L; L initially rigid, later flexible, longattenuate, erect to recurving, ultimately drooping, to 1–1.5 m  1.5–2.5 cm, broadened to 5 cm at the base, pale green, smooth and ribbed on both surfaces, margin serrulate, tip entire; Inf erect, paniculate, widely-branched, 0.8–2 m; primary Br to 30 cm, spreading; Bra very long and narrow above a dilated base, foliaceous, to 60 cm; Fl very numerous; Tep 2.5–3 (–4) mm, whitecreamy or with green-brown midline; Fr globose, 8–12 mm  8–12 mm ∅; Se ovoid to nearly globose, light brown, 3.5–4  3–3.5 mm. McVaugh (1989) substantially broadened the concept of N. parviflora to include N. longifolia and N. elegans, giving the species a very widespread distribution from (?) Chihuahua S to Oaxaca. In contrast, Trelease (1911) and Hochstätter (2010) maintained their separation.

C. C. Walker

N. elegans has been of uncertain status since its original description by Rose as “probably acaulescent”. Characteristic features of N. parviflora are the short, broad, widely branched panicles with short very showy long bracts and dense, numerous flowers. It is a small tree similar to N. parryi, but the leaves differ in size and form, being generally longer and more flexible in N. parviflora, initially erect, then spreading and drooping. Easily mistaken for Beaucarnea recurvata, but in contrast N. parviflora has a more arborescent habit and the leaves are long, lax, and not significantly thicker at the base. Superficially it also looks like Yucca filifera. Plants flower annually, but there are mass flowering cycles that last up to three years. Bees, wasps and flies are pollinators (Ruiz-Sanchez & Specht 2013). It is surprisingly little-known and seldom cultivated. It is frost-hardy to 10  C. N. pollyjeanneae Hochstätter (Acta Succ. 1 (1): 7, ills. (pp. 4–8), 2013). Type: USA, Oklahoma (Hochstätter 523.50 [SRP, HEID]). — Distr: USA (Oklahoma); isolated gentle limestone slopes, on dry rocky hills in open woodland 1300 m. [2] Low-growing acaulescent spreading shrubs forming dense clumps 1–2 m ∅; L thin, grass-like, quite flexible, 40–80  0.2–0.4 cm, deep green, convex at the base then concave, tip fibrous and tapered, upper surface smooth, lower surface rough, margin finely denticulate; Inf paniculate, 40–80 cm, branched, generally exceeding the leaves; Fl 2–3 mm ∅, white to greenish; Fr capsules 3–4 mm ∅, unilocular, light brown to almost transparent; Se globose, 2–3 mm ∅. Closest geographically and ecologically to the grass-like species N. greenei (not treated here) and N. texana, but endemic to Oklahoma. N. texana S. Watson (Proc. Amer. Acad. Arts 14: 248, 1879). Type: USA, Texas (Lindheimer 712 [MO]). — Distr: USA (Texas, C New Mexico), adjacent Mexico (?); rocky hillsides (limestone, granite), grasslands and shrublands, 200–2800 m. I: Small (1916); Gentry (1972: 183, fig. 72); Hochstätter (2010: 15).

Nolina RUSCACEAE

 Beaucarnea texana (S. Watson) Baker (1880); incl. Nolina erumpens [?] compacta Trelease (1911)  Nolina erumpens var. compacta (Trelease) Trelease (1912)  Nolina texana var. compacta (Trelease) I. M. Johnston (1943). [2] Low-growing acaulescent spreading shrubs; L narrowly linear, 70–120  0.3–0.4 cm, deeply rounded below, margins somewhat serrulate or smooth; Inf paniculate, slender, 30–60 cm, primary Br 10–15 cm; Bra caudate-attenuate, surpassing the branches, dry, yellowish, persisting; Fl 2.5–3.5 mm, white to creamy; Fr 5–7  4–5 mm, round-ovate; Se globose, 2.5–4 mm ∅. — Cytology: 2n = 38. In contrast to other members of Sect. Erumpentes, the inflorescence of N. texana is longer. It has a wide geographical range in the USA, and Gentry (1972) anticipates localities being found in NW Mexico (Sonora). This is one of the hardiest species in the genus and can tolerate temperatures below 20  C and hence is winter-hardy in Europe. Common names: “Texas Sacahuiste”, “Bunchgrass”.

References Dice, J. C. (1988) Systematic studies in the Nolina bigelovii — N. parryi (Nolinaceae) complex. San Diego (US): San Diego State University, unpubl. M. Sc. thesis. Folsom, D. B. [& al. 1995], Trager, J. N., Folsom, J., Clements, J. & Scott, N. (1995) Dry climate gardening with succulents. New York (US): Pantheon Books, Knopf Publishing Group. Gentry, H. S. (1972) The Agave family in Sonora. Agricultural handbook No. 399. Washington D.C. (US): US Department of Agriculture. URL: http://naldc.nal. usda.gov/download/CAT87209091/PDF Hess, W. J. (2002) Nolina Michaux. In: Flora of North America Editorial committee (eds.): Flora of North America North of Mexico, Volume 26: Magnoliophyta:

1351 Liliidae: Liliales and Orchidales; pp. 415–421, ills., key. New York (US)/Oxford (GB): Oxford University Press. Hess, W. J. & Dice, J. C. (1995) Nolina cismontana (Nolinaceae), a new species name for an old taxon. Novon 5(2): 162–164, key. URL: http://www.jstor. org/stable/3392239 Hochstätter, F. (2010) Il genere / The genus Nolina (Nolinaceae). Piante Grasse 30 (1: Suppl.): 1–48, ills., map, key. Hochstätter, F. & Donati, D. (2010) Una nuova specie appartenente al genera Nolina. A new species belonging to genus Nolina: Nolina hibernica Hochstätter & Donati. Piante Grasse 30(2): 72–77, ills. Hodgson, W. C. (2001) Food plants of the Sonoran Desert. Tucson (US): University of Arizona Press. McVaugh, R. (1989) Flora Novo-Galiciana. A descriptive account of the vascular plants of Western Mexico. Vol. 15: Bromeliaceae to Dioscoreaceae. Ann Arbor (US): University of Michigan Herbarium. Mitich, L. W. (1982) Parry’s Nolina (Nolina parryi). Cact. Succ. J. Gr. Brit. 44: 67–68, ill. Rivera-Lugo, M. & Solano, E. (2012) Flora del Valle de Tehuacán-Cuicatlan. Fascículo 99. Nolinaceae Nakai. México D.F. (MX): Instituto de Biología UNAM. Rowley, G. D. (1990) Nolina en culture. Cact.-Avent. 5: 2–4. Ruiz-Sanchez, E. & Specht, C. D. (2013) Influence of the geological history of the Trans-Mexican Volcanic Belt on the diversification of Nolina parviflora (Asparagaceae: Nolinoideae). J. Biogeogr. 40: 1336–1347. https://doi. org/10.1111/jbi.12073 Ruiz-Sanchez, E. & Specht, C. D. (2014) Ecological speciation in Nolina parviflora (Asparagaceae): Lacking spatial connectivity along of the Trans-Mexican Volcanic Belt. PLoS One 9(6): e98754, 11 pages, maps, ills. https://doi.org/10.1371/journal.pone.0098754 Small, J. K. (1916) Nolina texana. Addisonia 2: 1–2, t. 41. Thiede, J. (2012) Nomenclatural status of unranked names published by Trelease (1911) in Beaucarnea, Dasylirion, and Nolina (Asparagaceae — Nolinoideae). Phytoneuron 2012(77): 1–4. http://www.phytoneuron. net/77PhytoN-Beaucarnea.pdf? Trelease, W. (1911) The desert group Nolineae. Proc. Amer. Philos. Soc. 50(200): 404–443, tt. 1–17. URL: http://www.jstor.org/stable/983932 Walker, C. C. (2001) Introducing the Nolinaceae. Brit. Cact. Succ. J. 19(4): 214–221, ills.

Sansevieria RUSCACEAE L. E. Newton

Sansevieria Thunberg (Prodr. Fl. Cap., 65, 1794). Type: Sansevieria thyrsiflora Thunberg [nom. illeg., = Aloe hyacinthoides Linné (ICBN 2006: 280)]. — Dracaenoideae — Lit: Brown (1915: synopsis); Binojkumar (2002: synopsis India); Chahinian (2005a: ill. synopsis); Mbugua (2007: Flora Tropical E Africa); Carlquist & Schneider (2007: anatomy); Croix (2010: Flora Zambesiaca); Newton (2012b: with key to arborescent spp.); Mansfeld (2013a: ill. synopsis); Mansfeld (2014c: synopsis fieldnumbers); Lebrun & Stork (2014: 228–252, synopsis tropical Africa). Distr: Africa, Arabian Peninsula (Yemen, Oman), Comores, Madagascar, India, Sri Lanka, Myanmar. Etym: For Pietro Antonio Sanseverino, Count of Chiaromonte (1724–1771), Italian patron of horticulture in Naples and founder of a private botanical garden. Incl. Cordyline Adanson (1763) (nomen rejiciendum, Art. 56.1). Type: Aloe hyacinthoides Linné. Incl. Acyntha Medikus (1786) (nomen rejiciendum, Art. 56.1). Type: Aloe hyacinthoides Linné. Incl. Sanseverinia Petagna (1787) (nomen rejiciendum, Art. 56.1). Type: Sanseverinia thyrsiflora Petagna.

L. E. Newton (*) Barking, Essex, UK e-mail: [email protected]

Acaulescent or caulescent perennials, sometimes branching near the base, with subterranean rhizomes or stolons above ground; L solitary, few or many, distichous or in Ros, succulent or leathery, lanceolate, linear or lorate and flat, or cylindrical or semi-cylindrical and usually with a groove on the upper surface, sessile, sometimes narrowed at the base to resemble a petiole, apex often a pungent spine, green, often with lighter blotches, scattered or in transverse bands; Inf terminal, paniculate or unbranched, the fertile portion a dense or lax thyrse, elongated or sometimes capitate, with extrafloral nectaries associated with the bracts; Fl solitary or (usually) in a dense helicoid cyme, bracteate, pedicellate, actinomorphic, usually nocturnal and sweetly scented; Ped articulated; Tep united at the base to form a tube with 6 free lobes that curl back at anthesis, mostly whitish; St 6, extended beyond the perianth tube, exposed at anthesis by curling back of tepals; Sty simple, as long as the stamens or slightly longer; Ov with 3 locules, but often only 1 or 2 seeds mature; Fr resembling berries, but seeds covered with a fibrous endocarp. — Cytology: The few published chromosome numbers suggest that there is some polyploidy, with base numbers of x = 20 or 21. Early counts were summarised by Darlington & Wylie (1955). The genus has been variously included in the families Agavaceae, Asparagaceae, Dracaenaceae, Haemodoraceae, Liliaceae, and Ruscaceae. In the latest general classification it is placed in the Asparagaceae (APG 2009, 2016), though Nyffeler & Eggli (2010) proposed recognition of Ruscaceae

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_29

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s.l. Menale & al. (2013) argue that the name should be attributed to Petagna, since Thunberg merely used an erroneous spelling (Sansevieria, now a conserved name) for Petagna’s Sanseverinia. Symoens (2014) resolved the etymology of the name. Following morphological studies, Bos (1984) suggested that Sansevieria should be included in Dracaena. In a later publication (Bos 1998) Dracaena does include Sansevieria, but no transfers of individual species were made. Molecular studies by Lu & Morden (2014) supported this sinking of Sansevieria. Consequently new combinations were published by Mabberley (2017: 1101) and Christenhusz & al. (2018), transferring the species to Dracaena. Whilst the genus could be included in Dracaena eventually, Christenhusz & al. (2018) missed out several species, and some of their new names (e.g. for S. pinguicula fa. disticha, see there) are unacceptable, and so they are not followed in this account. Seedlings of species with cylindrical leaves, as well as young plants raised from cuttings, have short, flat leaves, and they look different from mature plants. In the descriptions below, leaf dimensions are length and width, plus thickness from front to back in the case of cylindrical and laterally compressed leaves. Although often referred to as “spike-like racemes”, the fertile (flowering) portions are neither spikes nor racemes — they are compound structures with flower clusters arranged along an axis, and not opening from the base upwards (Newton 2007). Jankalski (2009a) regarded the inflorescence as a thyrse, interpreting the flower clusters as axillary fascicled cymes, stating that the central flower in each cluster is the first to open. It was shown by Newton (2011) that the central flower is not the first to open, at least in some species examined, and so the neutral term glomerule was proposed. Budweg (2016) agreed that the inflorescences are simple or branched thyrses, but with helicoid cymes, confirming that the central flower is not the first to open. Useful diagnostic characters, such as whether leaf surfaces are rough or smooth, and the colour of rhizomes, are often not mentioned in protologues. Earlier literature stated that the seeds

L. E. Newton

have a fleshy covering, the sarcotesta, and no pericarp. Budweg (2014) discussed past interpretations of the fruit, and has shown that the seeds are covered with a fibrous endocarp. Many of the species included in Brown’s account were described from cultivated plants of undocumented wild origin, and in some cases even the country of origin was unknown. Many species are poorly represented in herbaria, and more field work is required to determine distribution and variation. Classification: An infra-generic classification was suggested by Pfennig, but it was not presented formally with names for the groups (Pfennig 1977a; Newton 2005a). Mbugua (2007) gave subgeneric names to the groups proposed by Pfennig, but without valid publication. Jankalski (2009a) published names for the same groups proposed by Pfennig, at the rank of section. Mansfeld (2013a) took this further by proposing subsections. Jankalski (2015) proposed an alternative classification within Sect. Sansevieria, with 16 species groups, without formal names. — The following synopsis is based on Pfennig’s scheme, with slight modification: [1] Inf branching (Sect. Dracomima Jankalski 2009) [2] Inf unbranched, fertile portion capitate (Sect. Cephalantha Jankalski 2009) [3] Inf unbranched, fertile portion elongated (Sect. Sansevieria) [3a] Plants with stolons above ground. [3b] Plants with underground rhizomes; L cylindrical or semicylindrical, upper surface often grooved. [3c] Plants with underground rhizomes; L flattened or folded. Succulence: Although most flat-leaf species appear more leathery than succulent, it has been shown by Koller & Rost (1988a) and Koller & Rost (1988b) that leaves of all Sansevieria species have succulent tissue. This occupies a central mesophyll and is unusual in having a branched network of living cells and solitary or grouped dead water storage cells, called tracheids by Carlquist & Schneider (2007). In all other succulent plants examined the water-storage cells are living.

Sansevieria RUSCACEAE

Ecology: With nocturnal white flowers, emitting a strong sweet scent, sansevierias are typically phalaenophilous, specifically sphingophilous. This pollination syndrome was observed by Newton (1994) and photographs of hawkmoths visiting flowers were shown by Rulkens & Baptista (2009a) and Mansfeld (2013b). An early morning visit to Sansevieria flowers by a sunbird was not regarded as effecting pollination (Newton 2004). It has been reported that some wild animals chew leaves of sansevierias without swallowing them, perhaps for moisture content, including baboon (Newton & Mbugua, 1993), elephant (Newton 2009b) and rhinoceros (Newton 2003b), or possibly even for medicinal purposes (Newton 2003b). Ethnobotany: Some species have been used at a local level as a source of cordage fibre, giving rise to such vernacular names as “Bowstring hemp” and “African sisal”. Hybrids with improved fibre quality have been produced in the USA (Joyner & al. 1951; Wilson 1962). Arising from their popularity as ornamentals, numerous cultivars have been described, especially with varying degrees of variegation, and some cases of small-growing (“dwarf”) variants. Jankalski (2009b) compiled a list of known hybrids and cultivars. Several other uses were documented by Khalumba & al. (2008), Takawira-Nyenya & Stedje (2011), and Takawira-Nyenya & al. (2014). The following names are of unresolved application but are referred to this genus: Sansevieria cylindrica Baum (1903) (nom. illeg., Art. 53.1); Sansevieria ehrenbergii K. Schumann (1900) (nom. illeg., Art. 53.1); Sansevieria fulvocincta Haworth (1819); Sansevieria glauca Haworth (1812); Sansevieria guineensis var. γ Schultes (1829); Sansevieria laetevirens Haworth (1812); Sansevieria polyphylla Haworth (1812); Sansevieria pumila Spin (1818) (nom. illeg., Art. 53.1); Sansevieria stenophylla Link (1821); Sansevieria striata G. Don ex Steudel (1841) (nom. inval., Art. 32.1c); Sansevieria venosa G. Don ex Steudel (1841) (nom. inval., Art. 32.1c). S. aethiopica Thunberg (Prodr. Fl. Cap., 65, 1794). Type: RSA, Eastern Cape (Thunberg s.n. [UPS]). — Distr: Botswana, Namibia,

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Fig. 1 Sansevieria aethiopica. (Copyright: L. E. Newton)

Malawi, Moçambique, Zimbabwe, RSA (Northern Cape, North-West Prov., Eastern Cape, Gauteng); dry open places or bushland on well-drained soil. I: Obermeyer & al. (1992: 6). – Fig. 1.  Dracaena aethiopica (Thunberg) Byng & Christenhusz (2018); incl. Aletris hyacinthoides var. zeylanica Aiton (1789) (nom. illeg., Art. 53.1); incl. Sansevieria zeylanica Redouté (1809) (nom. illeg., Art. 53.1); incl. Sansevieria glauca Gérôme & Labroy (1903) (nom. illeg., Art. 53.1); incl. Sansevieria thunbergii Mattei (1918) (nom. illeg., Art. 52.1); incl. Sansevieria caespitosa Dinter (1926); incl. Sansevieria scabrifolia Dinter (1932); incl. Sansevieria transvaalensis hort. ex Morgenstern (1979) (nom. inval., Art. 36.1, 37.1). [3c] Acaulescent, rhizomatous; rhizome 1 cm ∅; L 13–30, rosulate, ascending-spreading, linear or linear-lanceolate, 13–43  1–2 cm, upper surface concave, base sometimes slightly narrowed, gradually narrowed above the middle

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to a subulate green tip 1.6–2.5 mm long, soon becoming white, otherwise dark green with bluish hue,  glaucous, sometimes with paler green transverse bands, margin red or whitish, surface slightly rough; Inf 35–75 cm, simple, elongate, fertile portion dense, 4–6 Fl per cluster; Bra ovate-lanceolate, acute, 0.5–1 mm; Ped to 5 mm; Fl white, sometimes purple or cream, tube 20–35 mm, lobes 15–20 mm. Croix (2010) includes Malawi and Moçambique in the distribution as new records. Mbugua (2007) described a plant in Kenya as S. aethiopica ssp. itumei (as ‘itumea’), but this is now regarded as a separate species (see S. itumei) (Newton 2016). S. arborescens Cornu ex Gérôme & Labroy (Bull. Mus. Hist. Nat. (Paris) 9: 170, 172–173, ills., 1903). Type [neo]: Kenya, Coast Prov. (Ngwiri s.n. in Newton 5884 [EA]). — Lit: Newton (2009a). Distr: Kenya, N Tanzania; lowland Acacia bushland, 40–375 m. I: Sansevieria J. 3(1): 36–37, 1994.  Dracaena arborescens (Cornu ex Gérôme & Labroy) Byng & Christenhusz (2018). [1] Caulescent, rhizomatous, rhizome 1 cm ∅, pale brown; stem erect, to 20–150  2–2.5 cm; L many, densely spiralled, spreading or recurved, lanceolate or linear-lanceolate, upper surface concave, 15–22  2–4.5 cm, base scarcely or slightly narrowed, green with whitish or reddish slightly wavy margin, surface smooth, with a 8.5–25.4 mm pale brown stout pungent subulate tip; Inf 50 cm, paniculate, lower branches to 8 cm, fertile portion subdense, 4–6 Fl per cluster; Bra triangular, 2 mm; Ped 3 mm; Fl white, tube 6 mm, lobes 9 mm. Originally described with flowers unknown. Cornu labelled living plants with the name, but did not publish it. Brown (1915) erroneously applied this name to the plants later described as S. dumetescens and S. ascendens. S. ascendens L. E. Newton (Bradleya 28: 19–21, ills., 2010). Type: Kenya, Coast Prov. (Newton 5842 [K, EA]). — Distr: S Kenya, N Tanzania; dry bushland, 160–350 m.  Dracaena ascendens (L. E. Newton) Byng & Christenhusz (2018).

L. E. Newton

[1] Caulescent, rhizomatous; rhizome 2 cm ∅, yellow-buff in colour; stem unbranched, to 1.5 m, erect or becoming decumbent; L spiral, spreading, to 60  3 cm, 10–15 mm thick, canaliculate above, narrowed to a pungent 1 cm red-brown spine-like tip, mid-green, lower surface with 10–15 dark green longitudinal lines for most of the length, margin 1 mm, red-brown with whitish fraying edge, both surfaces smooth; Inf to 1 m  35 cm, paniculate with 20 ascending branches, lower ones to 40 cm with secondary branching, fertile portion erect, to 21 cm, subdense, 4–6 Fl per cluster; Bra rounded, 1  2 mm, base brownish, white above; Ped 3 mm, pale green; Fl white, finely speckled brown, tube 5  3 mm, lobes 8  1.5 mm. Suggested to be the hybrid S. dumetescens  S. ehrenbergii by Jankalski (2015), but this is unlikely, as S. ascendens is known from 3 distinct populations, and S. ehrenbergii is not recorded from anywhere near this area (Newton 2016). S. aubrytiana Carrière (Rev. Hort. (Paris) 33: 448–449, 1861). Type: Gabon (Aubry le Comte s.n. [P]). — Distr: Angola, Gabon, Tanzania; dry areas.  Dracaena aubrytiana (Carrière) Byng & Christenhusz (2018); incl. Sansevieria bracteata Baker (1878). [3c] Acaulescent, rhizomatous; L erect, lanceolate, 38–60  5–7 cm, base narrowed from the middle or below into a channelled petiole, dark green with wide pale green irregular bands or blotches on both surfaces, glaucous, narrowed above to a 6.4 mm tip, margin hard, brownishred, sometimes with whitish edge; Inf 45–60 cm, simple, elongated, fertile portion 5–6 cm, dense; Bra lanceolate or oblong-lanceolate, acute, 1.3–1.9 mm; Ped 2–3 mm; Fl white, tube 88–114 mm, lobes 25–32 mm. The first edition of this handbook, as well as Mbugua (2007) treat this as S. bracteata, but the name S. aubrytiana has priority. Treated as a doubtful species by Lebrun & Stork (2014), though accepted by Chahinian (2005a) and Mansfeld (2015a). Lebrun & Stork (2014) also treat S. bracteata as distinct.

Sansevieria RUSCACEAE

S. bacularis Pfennig ex A. Butler & Jankalski (Sansevieria 22: 3–4, ills. (pp. 3–7), 2010). Type: Democratic Republic of the Congo [Zaïre], Kivu Prov. (Bequaert 5367 [BR]). — Distr: E Democratic Republic of the Congo [Zaïre] (Sud-Kivu). I: Mansfeld (2015b).  Dracaena bacularis (Pfennig ex A. Butler & Jankalski) Byng & Christenhusz (2018); incl. Sansevieria cylindrica De Wildemans (1921) (nom. illeg., Art. 53.1). [3b] Acaulescent, rhizomatous; L 1–2, usually upright, cylindrical or slightly compressed, 125–170  1.1–1.5 cm, apex soft, dark green with lighter cross banding, verrucose, somewhat rough, base sheathed in 2.5  2–3 cm cataphylls, deep purple, pale brown when dry; Inf to 115 cm, simple, elongated, fertile portion 30–45  7.5 cm, semidense, 2–8 Fl per cluster; Bra lanceolate, 5–10 mm; Ped 6–7 mm; Fl white with purple stripes, tube 5  2.5 mm, lobes 15–17  2–2.5 mm. The name was proposed by Pfennig and appeared in print by other authors without validation until 2010. S. bagamoyensis N. E. Brown (Bull. Misc. Inform. Kew 1913: 306, 1913). Type: Tanzania, Bagamoyo Distr. (Sacleux 672 [P]). — Lit: Newton (2012b: with ills.). Distr: N Tanzania; halfshade of coastal bushland, 100–200 m.  Dracaena bagamoyensis (N. E. Brown) Byng & Christenhusz (2018). [1] Caulescent, rhizomatous; rhizome 1–1.5 cm ∅, pale brown; stem erect to 2 m; L spiralled, recurved or recurved-spreading, linear-lanceolate or linear, upper surface concave, 34–40  1.6–2.5 cm, dark green, narrowed to a 4.2 mm brown hard spine-like tip, margin narrowly red-brown with white membranous edge, surface smooth; Inf 40–55 cm, paniculate, fertile portion lax,  6 Fl per cluster; Bra lanceolateacute, 2–3 mm; Ped to 6 mm; Fl cream-white, tube 6–7 mm, lobes 6.5 mm. S. ballyi L. E. Newton (Brit. Cact. Succ. J. 22(1): 11–13, ills., 2004). Type: Kenya, Coast Prov. (Newton 5594 [K, EA]). — Lit: Mansfeld (2016a: history of discovery, with ills.); Jankalski

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(2016: synopsis cultivars). Distr: Kenya, Tanzania.  Dracaena ballyi (L. E. Newton) Byng & Christenhusz (2018). S. ballyi var. ballyi — Lit: Newton (2010). Distr: S Kenya, N Tanzania; rocky areas, often in shade, 65–975 m. Incl. Sansevieria tsavoensis Pfennig ex Heitz & Zeller (1990) (nom. inval., Art. 36.1, 37.1). [3a] Acaulescent, stoloniferous; branching freely above ground; stolons to 18 cm; L 6–10, rosulate, spreading, almost cylindrical, 6–12  6–9 cm, with a groove on the upper surface from the base to of the leaf length, narrowed to a pungent 7 mm red-brown spine-like tip, with alternating bands of dark and light green, margins of the groove with red-brown line and narrow white edge, surface rough; Inf to 15.5 cm, simple, elongated, fertile portion to 8 cm, subdense, 2 Fl per cluster; Bra triangular, 3  3 mm; Ped 1–2 mm; Fl pale pink, tube 18–22  2–2.7 mm, lobes 10–13  1.5 mm, whitish inside. For many years cultivated as Sansevieria ‘Bally 12681’. No herbarium specimen was deposited by Bally, but the type was collected at the same locality. S. gracilis var. humbertiana in the sense of Mbugua (2007) belongs here. Mansfeld & Budweg (2015a) describe a cross with S. parva (male), and report that the reciprocal cross was not possible. S. ballyi var. robertsoniae L. E. Newton (Sansevieria 33: 11–12, ill., 2015). Type: Kenya, Coast Prov. (Newton & al. 5981 [EA]). — Distr: S Kenya (Coast Prov.); savanna woodland, in shade, 100 m.  Sansevieria ballyi ‘Ann Robertson’ L. E. Newton (2010). [3a] Differs from var. ballyi: L with paler colour and very smooth surface. First described in 2010 as the cultivar S. ballyi ‘Ann Robertson’. S. bella L. E. Newton (Cact. Succ. J. (US) 72 (4): 224–226, ills., 2000). Type: Kenya, Rift Valley Prov. (Newton 3945 [K, EA]). — Distr: S

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Kenya; under shrubs or amongst rocks in dry bushland, 1700–2030 m.  Dracaena bella (L. E. Newton) Byng & Christenhusz (2018). [3a] Caulescent, with runners, branching freely above ground; runners spreading or ascending, to 15 cm; L up to 8,  distichous, ascending, cylindrical with a groove on the upper surface from the base to  of their length, to 70 cm, to 3.5 cm thick, with distinct dark and light green transverse bands and darker green narrow longitudinal lines, tip a red-brown spine to 5 mm, pungent or relatively blunt, margins with a red-brown line and a narrow colourless flange, surface very rough; Inf to 60 cm, simple, elongated, peduncle light green with a slight bloom, fertile portion to 55 cm, dense, up to 7 Fl per cluster; Bra triangular, 3  2 mm; Ped to 5 mm; Fl white, tube 10–15 mm, lobes 13–18 mm. Close to S. suffruticosa and possibly better treated as a variety of that. S. bhitalae R. H. Webb & L. E. Newton (Sansevieria 37: 12–15, ills., 2018). Type: Tanzania, Iringa Region (Bhitala 1000 [MO]). — Distr: Tanzania (Iringa Region); in deep shade on hillslopes and on old anthills, 1360 m. [2] Rhizomatous, acaulescent; L 1–3, erect, laterally compressed (folded) with a dorsal broad U-shaped groove 1.2 cm deep, 40–90  5–7 cm, yellow-green to blue-grey-green, with yellowgreen mottling that ranges from prominent to very faint, and upper surface with faint longitudinal striations (up to 8 near the base) nearly the entire leaf length, surface smooth; Inf capitate; peduncle reddish, 15 cm, clusters with 2–3 flowers; Bra broadly ovate, 16  5 mm; Fl white to lilac with a purple-violet blush, tube 75 mm, lobes 28 mm. There is a juvenile form, possibly persistent for a few years, with 2–6 leaves, ovate-elliptical to oblanceolate, blue-green to grey-green, mottled with yellow-green markings and lines, rigid and nearly prostrate to spreading with a pointed tip and a red leaf margin with some shredding white fibres. S. braunii Engler & K. Krause (Bot. Jahrb. Syst. 45: 153–154, 1911). Type: Tanzania,

L. E. Newton

Kigoma (Braun s.n. [EA]). — Distr: SW Tanzania; woodland in deep shade, 500–1160 m.  Dracaena testudinea Byng & Christenhusz (2018). [2] caulescent, rhizomatous, rhizome 3 cm ∅, brownish-orange; L usually 2, lanceolate-oblong, 50–70  7–11 cm, base slightly narrowed, green with few whitish bands or blotches, tip acuminate, margin hardened, red-brown; Inf 45 cm, simple, cylindrical, peduncle reddish, fertile portion 15 cm ∅, dense; Bra lanceolate-acute, 20  6 mm; Ped 10 mm; Fl white, tube 82 mm, lobes 25–30 mm. The plants treated as S. braunii by Mbugua (1998) are S. frequens. S. burdettii Chahinian (Brit. Cact. Succ. J. 18 (3): 132–133, ills., 2000). Type: Malawi, Southern Region (Burdett s.n. in Chahinian 318 [K, MO]). — Lit: Rulkens & Baptista (2013); Thiede & Campbell-Barker (2015). Distr: C to S Malawi, NE Moçambique; rocky woodland, 100–500 m.  Dracaena burdettii (Chahinian) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; rhizome to 4 cm ∅, orange; L 3–6, distichous, slightly spreading, cylindrical with a groove on the upper surface from the base to to of the leaf length, to 90 cm, base 2.5 cm wide, dark green with several longitudinal lines, sometimes with faint transverse bands on young leaves, margins green with a withered non-fibrous flange, tip a withered spine, surface smooth, waxy; Inf 16–20 cm, simple, elongated, peduncle green with small white spots, fertile portion 12–14.75 cm, dense, 4–6 Fl per cluster; Bra 1–2 mm; Ped 4 mm; Fl white tinged pinkish-brown, tube 20–25 mm, lobes 18–20 mm. Type cited erroneously as Chahinian 316 in the protologue. S. burmanica N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 228–230, Fig. 12, 1915). Type: Myanmar, Mandalay Prov. (Clayton s.n. [K]). — Lit: Mansfeld & Ott (2015). Distr: N Myanmar, S India (Tamil Nadu); scrub forest,  100 m. I: Mansfeld (2016b).

Sansevieria RUSCACEAE

 Dracaena burmanica (N. E. Brown) Byng & Christenhusz (2018); incl. Sansevieria roxburghiana Hooker fil. (1896) (nom. illeg., Art. 53.1); incl. Sansevieria maduraiensis Binojkumar (2002). [3c] Acaulescent, rhizomatous; rhizome 1.25–1.8 cm ∅; L 8–18, rosulate, erect, linear or linear-lanceolate, flat or upper surface concave, 45–76  1–3.2 cm, green with paler transverse bands, upper surface with 1–3 longitudinal striations, lower surface with 6–9 striations, narrowed above to a 2.5–10 cm green soft subulate tip, margin green, becoming whitish with age, surface smooth; Inf 60–76 cm, simple, elongated, fertile portion lax, 2–5 Fl per cluster; Bra lanceolate, acute, 2–4 mm; Ped 7.5–8.5 mm; Fl greenish-white, tube 7.5–8.5 mm, lobes 8.5 mm. After being lost in cultivation for many years, this species was rediscovered in habitat in 1982 (Mansfeld & Ott 2015). S. canaliculata Carrière (Rev. Hort. (Paris) 33: 449, 1861). Type [neo]: Comores, Mayotte (Boivin 3079 [P, BM]). — Distr: Madagascar, Comores, N Moçambique, Tanzania; sandstone outcrops, 50–100 m. I: Brown (1915: 224).  Dracaena canaliculata (Carrière) Byng & Christenhusz (2018); incl. Sansevieria sulcata Bojer ex Baker (1875) (nom. inval., ICN Art. 36.1c); incl. Sansevieria sulcata Baker (1887); incl. Sansevieria schimperi Baker (1898); incl. Sansevieria livingstoniae Rendle (1932); incl. Sansevieria pfennigii Mbugua (2007). [3b] Acaulescent, rhizomatous; rhizome 1–1.25 cm ∅; L 1–2, to 5 cm apart along the rhizome, erect or slightly curved, cylindrical or slightly laterally compressed, with 5–6 shallow longitudinal grooves, 15–76 cm, 1–2 cm thick, dark green, narrowed shortly to a whitish hard acute tip, surface almost smooth; Inf 5–16 cm, simple, elongated, fertile portion lax, Fl usually 3 per cluster, solitary near the tip; Bra ovateacute,  4 mm; Ped 1–1.6 mm; Fl white tinged green, tube 25 mm, lobes 17 mm. — Cytology: 2n = 42 (Sharma & Chaudhuri 1964). Described from a cultivated plant of unknown origin. Brown (1915) cited a specimen from

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Somalia, but Thulin (1995) found no material agreeing with the description. The species was reported by Perrier (1938) to be naturalized in Madagascar. S. caulescens N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 200, Fig. 2A–B, 1915). Type: Kenya (Powell s.n. [K]). — Distr: Kenya.  Dracaena caulescens (N. E. Brown) Byng & Christenhusz (2018). [3a] Caulescent; stem erect, to 60  2.5–3.8 cm, branching at the base; L many, densely spiralled, spreading and slightly recurved, cylindrical with a groove along the upper face almost to the tip, 45–84  2–3.4 cm, 1.3–1.8 cm thick, dark green, lower surface with 9–12 dark longitudinal striations, with whitish edges, when young with indistinct transverse dark green bands, gradually narrowed to a firm green tip becoming pale brown or whitish, surface slightly rough; Inf 60–70 cm, simple, elongated, fertile portion 40–46 cm, dense, 6–12 Fl per cluster; Bra very small, inconspicuous; Ped 7.4–9.5 mm; Fl whitish, tube 15 mm, lobes 19 mm. I am not aware of any recent field gatherings matching this description. Lebrun & Stork (2014) followed Mbugua (2007) in treating this as a synonym of S. gracilis, but that species has stems that rarely exceed 8 cm in length. S. chahinianii R. H. Webb & Myklebust (Sansevieria 37: 6–7, ills., 2018). Type: Somalia (Specks 14026 [MO]). — Distr: E Somalia (W of Eyl); mixed savanna, 180 m. [3c] Acaulescent, rhizomatous, rhizomes to 20 cm; L (1–) 2 (–4), ascending to spreading, oblanceolate, 10–25  6–7 cm, grey-green with dark green irregular transverse bands, with a shallow central channel bearing longitudinal lines that are more prominent towards the base, tip 5 mm, white, chartaceous, margin entire, with outer thin white line adjacent to a thin red line, upper surface smooth, lower surface very rough; Inf 33 cm, simple, fertile portion to 15 cm, subdense, with 2 flowers per cluster; Bra triangular, scarious, 2  5 mm; Ped 1–2 mm; Fl yellow-white, tube 20–22 mm, lobes 15 mm.

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S. concinna N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 233–234, Fig. 14, 1915). Type: Moçambique, Sofala Prov. (Dawe 1 [K]). — Distr: Moçambique, Tanzania, Zimbabwe, NE RSA (KwaZulu-Natal); in the shade of coastal forest in sandy soil. I: Jaarsveld (1994).  Dracaena spathulata Byng & Christenhusz (2018)  Sansevieria subspicata var. concinna (N. E. Brown) Mbugua (2007). [3c] Acaulescent, rhizomatous; rhizome 1.25 cm ∅; L  5, rosulate, ascendingspreading, lanceolate, 15–25  1.25–3 cm, base shortly narrowed to a 3.8–8.9 cm channelled petiole, green with pale green transverse bands, with a 4.2–8.5 mm green subulate tip, margin scarcely hardened, green, surface smooth; Inf 15–30 cm, simple, elongate, peduncle green, tinged and dotted purple, fertile portion subdense, 1 or 2 Fl per cluster; Bra lanceolate-acuminate, 6–8.5 mm; Ped 3–4 mm; Fl white, tube 45 mm, lobes 21 mm. Treated as a variety of S. subspicata by Lebrun & Stork (2014), but accepted by Chahinian (2005a) and Mansfeld (2015a). S. conspicua N. E. Brown (Bull. Misc. Inform. Kew 1913: 306, 1913). Type: Kenya, Coast Prov. (Powell 12 [K]). — Distr: Kenya, Malawi, Tanzania; coastal thicket and grassland, to 375 m. I: Powys (1996). – Fig. 2.  Dracaena conspicua (N. E. Brown) Byng & Christenhusz (2018). [3c] Acaulescent, rhizomatous; rhizome 1.7–1.9 cm ∅; L 3–5, rosulate, ascendingspreading, lanceolate, 23–76  5–8.3 cm, base narrowed from below the middle, dull green, upper surface with darker longitudinal striations, with a 2.1–3.2 mm hard tip, margin hardened, reddish-brown and edged white, surface smooth; Inf to 60 cm, simple, elongated, peduncle greyishgreen tinged dull purple, fertile portion 25.5–30 cm, dense, 1–3 Fl per cluster; Bra linear-lanceolate, 0.6–1.3 mm; Ped 0.4–0.6 mm; Fl greenish-white below, white above, tube 38–42 mm, lobes 25–32 mm. S. cylindrica Bojer ex Hooker (Curtis’s Bot. Mag. 85: t. 5093 + text, 1859). Type: Angola

L. E. Newton

Fig. 2 Sansevieria conspicua. (Copyright: L. E. Newton).

(Welwitsch 3749 [K, P]). — Distr: Angola, Zambia, Zimbabwe. S. cylindrica var. cylindrica — Distr: Angola, Zambia, Zimbabwe. Incl. Sansevieria angolensis Welwitsch ex Hooker (1858) (nom. inval., Art. 32.1d)  Dracaena angolensis (Welwitsch ex Carrière) Byng & Christenhusz (2018); incl. Sansevieria guineensis Weiner (1887) (nom. illeg., Art. 53.1). [3b] Acaulescent, rhizomatous; rhizome 2.5–3.8 cm ∅; L 3–4, distichous, erect, cylindrical or slightly laterally compressed, 60–150 cm, 2–3 cm thick, green or whitish-green, with dark green transverse bands, gradually narrowed to a 4.2–6.4 mm hard whitish acute tip, surface slightly rough; Inf 60–90 cm, simple, elongated, fertile portion 38–75 cm, 5–6 Fl per cluster; Bra lanceolate or ovate-lanceolate, 4–10 mm; Ped 4–8.5 mm; Fl white or tinted, tube 17–25 mm, lobes 17–19 mm. — Cytology: 2n = 40 (Roy

Sansevieria RUSCACEAE

1956), 92 (Sharma & Chaudhuri 1964), 102–104 (Darlington & Wylie 1955). S. cylindrica var. patula N. E. Brown (Bull. Misc. Inform. Kew 1915 (5): 218–219, Fig. 5D, 1915). Type: Angola (Anonymus s.n. [K]). — Distr: Angola. [3b] Differs from var. cylindrica: L 3–6, diverging, recurved or spreading; Fl tube 11.5–19 mm, lobes 15–19 mm. Not recognised by Croix (2010), who suggests that it should probably be treated only as a cultivar. S. dawei Stapf (J. Linn. Soc., Bot. 37: 529, 1906). Type: Uganda, Central Distr. (Dawe 109 [K]). — Lit: Mbugua (2003). Distr: Kenya, Uganda, Burundi; open woodland, 600–1300 m.  Dracaena dawei (Stapf) Byng & Christenhusz (2018). [3c] Acaulescent, rhizomatous; rhizome to 2.5 cm ∅, pale brown; L 2–3, ascending or suberect, lanceolate, 60–150  5.7–11 cm, base narrowed from below the middle to a long or short channelled petiole, dull dark green and glaucous, narrowed above the middle to an acute tip, margin reddish-brown, upper surface smooth, lower surface slightly rough; Inf 45–75 cm, simple, elongated, dense, 3–4 Fl per cluster; Bra ovate or ovate-oblong, acute or subobtuse, 8.5–17 mm; Ped 4–6 mm; Fl white, tube 19–25 mm, lobes 17–22 mm. Mansfeld (2014b) suggests that S. dawei should be included in the synonymy of S. forskaoliana, though he did not examine material from the type locality, in Uganda. S. dhofarica T. A. McCoy & Lavranos (Avonia 35(1): 65, ills. (pp. 64–67), 2017). Type: Oman, Dhofar (McCoy 5014 [FT]). — Distr: Oman (Dhofar); limestone hills,  80 m. [1] Acaulescent, rhizomatous, forming large clumps; rhizome 2–2.3 cm ∅, greyish-tan; L 4–5, distichous, ascending, laterally compressed, deeply channelled low down, 50–60  5–6 cm, 5–7 cm thick at the base, with 4–7 deep grooves on the flattened sides, light to dark green, surface slightly rough, margin entire,

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reddish-brown, with thin membranous edge, tip acute, pungent; Inf 100 cm, with 6–12 Br, peduncle green or reddish-green; fertile portion 6–8 cm, 4–7 Fl per cluster; Bra attenuate-acute, 0.2  0.1 mm, green; Ped 0.3 mm; Fl tube 12 mm, colour and lobe size not given in the protologue. S. dooneri N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 231–233, Fig. 13A–B, 1915). Type: Kenya, Rift Valley Prov. (Dooner s.n. [K]). — Lit: Newton (2003a). Distr: Kenya, Tanzania; shade in dry forest,  2000–2100 m.  Dracaena dooneri (N. E. Brown) Byng & Christenhusz (2018). [3c] Acaulescent or shortly caulescent, rhizomatous or with runners; rhizome 0.6–0.85 cm ∅; stem to 5 cm, concealed by leaf bases; L to 20, in lax rosettes, lanceolate or lorate, base erect or ascending, recurved-spreading above, 10–43  1.5–3 cm, base gradually narrowed from near or above the middle, dark green, lower surface slightly paler, both surfaces with faint and irregular pale green transverse bands, shortly narrowed to a 6–50 mm green soft subulate tip, margin green, surface smooth; Inf 30–38 cm, simple, elongated, peduncle green, fertile portion 15–19 cm, lax, 2–3 Fl per cluster; Bra ovatelanceolate, acute, 2–4 mm; Ped 3–4 mm; Fl dull pink or pale purplish, whitish inside, tube 12 mm, lobes 11.6–12.7 mm. Similar to, and possibly conspecific with, S. parva, whose type locality is only 55 km away from that of this species. Included in S. parva by Mbugua (2007), Lebrun & Stork (2014) and Mansfeld (2015a), though regarded as distinct by Chahinian (2005a). S. downsii Chahinian (Brit. Cact. Succ. J. 18 (3): 133–135, ills., 2000). Type: Malawi, Northern Region (Downs 1/75 [K, MO]). — Lit: Thiede & al. (2009: with ills.). Distr: N Malawi, NE Zambia; dry woodland, 1000–1500 m.  Dracaena downsii (Chahinian) Byng & Christenhusz (2018). [3a] Acaulescent, with runners, runners to 2 cm ∅, covered with leaf sheaths; L 6–14, rosulate, spreading-recurved, cylindrical with a groove on

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the upper face from the base to of the leaf length, 14–45 cm, base 3.2 cm wide, medium bluish-green with sparse grey-green transverse bands and dark green longitudinal lines, margins with a chestnut-brown line and a narrow colourless flange, tip a spine, sometimes chestnut-brown, surface slightly rough, somewhat shiny, waxy; Inf to 1.6 m, simple, elongated, peduncle medium green, speckled, fertile portion < of the peduncle length, lax, 3–5 Fl per cluster; Bra 1–2 mm; Ped 3 mm; Fl white tinged green, tube 10 mm, lobes 12 mm. The plants reported from Malawi as possibly being S. gracilis by Thiede (1993) are S. downsii. S. dumetescens L. E. Newton (Bradleya 27: 156–158, ills., 2009a). Type: Kenya, Kilifi Distr. (Newton 3761 [K, EA]). — Distr: S Kenya; dry bushland, 100–200 m.  Dracaena dumetescens (L. E. Newton) Byng & Christenhusz (2018). [1] Caulescent, rhizomatous, forming dense clumps to 2 m ∅ or more; rhizome 1.8–2.3 cm ∅, pale brown; stem erect, to 1.6 m  2–2.5 cm; L spiral, spreading or recurved, lanceolate or linear-lanceolate, 40–55  3–4.5 cm, narrowed to a pungent 13–17 mm pale brown spine-like tip, green with whitish or reddish slightly wavy margin, both surfaces smooth; Inf to 55 cm, paniculate, with up to 18 Br, the lowest to 20 cm with secondary branching, fertile portion subdense, 4–6 Fl per cluster; Bra triangular, 1–2  1 mm, colourless; Ped 3 mm; Fl white, tube 6  2.5 mm, lobes 9  1.8 mm. Here belongs S. arborescens in the sense of Brown (1915). S. ebracteata (Cavanilles) C. R. Suresh (in Nicolson, Interpret. Van Rheede’s Hort. Malab., 271, 1988). Type: [icono] Rheede, Hort. Malab. 11: 83, t. 42, 1692. — Lit: Mansfeld & Ott (2015). Distr: S India (Kerala, Karnataka, Tamil Nadu, Hyderabad); sandy places.  Salmia ebracteata Cavanilles (1795)  Dracaena ebracteata (Cavanilles) Byng & Christenhusz (2018); incl. Aletris zeylanica var. β Lamarck (1783); incl. Sansevieria lanuginosa Willdenow (1797) (nom. illeg., Art. 52.1)  Acyntha

L. E. Newton

lanuginosa (Willdenow) Kuntze (1891) (nom. illeg., Art. 52.1). [3b] Acaulescent, rhizomatous; rhizome stout; L 3–5, erect, semiterete with concave channel down the upper surface and several grooves down the lateral and lower surfaces, 45–90  1.7–2 cm, narrowed to an acute tip, green with “woolly” grooves, surface rough; Inf  60 cm, simple, elongated, density unknown, 2–5 Fl per cluster; Ped  4.2 mm; Fl whitish-green, tube 8.5–10 mm, lobes 10–15 mm. S. ehrenbergii Schweinfurth ex Baker (J. Linn. Soc., Bot. 14: 549, 1875). Type: Sudan (Schweinfurth 31 [B?, K, P]). — Distr: Yemen, Djibouti, Eritrea, Ethiopia, Kenya, Somalia, Sudan, Tanzania, Uganda; rocky ground, usually in the shade of thickets or small trees, 400–1100 m. I: Teketay (1995). [1] Acaulescent or caulescent, rhizomatous; rhizome 3 cm ∅; stem erect, to 25 cm; L 5–9, distichous, erect or spreading, laterally compressed with a groove along the upper surface, 76–180 cm, 3–4.5 cm thick, narrowed to an abrupt 6.5–20 mm hard spine-like tip, dark green with 5–12 blackish-green shallow longitudinal grooves, margin reddish-brown with white membranous edge, surface slightly rough; Inf to 2 m, paniculate, fertile portion with 4–7 Fl per cluster; Ped 2–4 mm; Fl purple to white, tube 5–6.5 mm, lobes 7–18 mm. — Cytology: 2n = 40 (Sharma & Chaudhuri 1964). Often confused with S. robusta, and possibly not as widespread as shown above (from literature records). The species is quite variable. Some more robust variants reported in Yemen (Vrskovy 2009) are probably S. dhofarica. S. eilensis Chahinian (Sansevieria J. 4(1): 9–11, ills., 1995). Type: Somalia, Nugaal Distr. (Lavranos & Horwood 10178 [MO, UPS]). — Distr: E Somalia; shaded limestone,  120 m. I: Lavranos (1994: as S. sp.). – Fig. 3.  Dracaena eilensis (Chahinian) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; L 2–3, mostly distichous, cylindrical, sometimes with a groove on the upper surface, 7–18 cm, 1.9–2.5 cm thick,

Sansevieria RUSCACEAE

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elliptic or oblong-elliptic, 16–30  9–14 cm, 4–5 mm thick in the middle, distinctly canaliculate, tip broadly and shortly attenuate and acute, margins sometimes undulate, both surfaces dark green with irregular transverse bands of pale grey-green blotches, sometimes irregularly reticulate, margins purple-brownish with narrow white edge, upper surface slightly rough, lower surface very rough; Inf 20–35 cm, simple, with 1–2 basal bracts, elongated, fertile portion 15 cm, 3–5 Fl per cluster; Bra 15–20  7–8 mm, lanceolate, acuminate, membranous, whitish; Fl unknown. The protologue does not include details of flowers. Included in S. forskaoliana by Thulin (1995) and Mbugua (2007), but Jankalski (2007c) argues for its recognition as a distinct species, with spreading leaves smooth above and rough below, and foliaceous bracts on the peduncle (though see note after S. forskaoliana). Included in S. forskaoliana by Lebrun & Stork (2014) and Mansfeld (2015a). Fig. 3 Sansevieria eilensis. (Copyright: J. Trager).

narrowed from the middle to the base, narrowed abruptly to a 5 mm acute spine-like tip, medium grey-green, with light grey-green transverse bands and up to 12 medium green longitudinal lines, margin green becoming brown edged white, surface rough; Inf 34 cm, simple, spike-like, peduncle light green, fertile portion 23 cm, lax, 2–4 Fl per cluster; Bra 2–6  1–3 mm; Ped 4–5 mm; Fl greenish-white, tube 8 mm, lobes 14 mm. The type citation Lavranos 10179 in the protologue is an error. S. elliptica (Chiovenda) Chiovenda ex Guidotti (Agric. Colon. 26: 549, 1932). Type: not typified. — Distr: Somalia, Kenya.  Acyntha elliptica Chiovenda (1932); incl. Acyntha abyssinica var. sublaevigata Chiovenda (1932)  Sansevieria abyssinica var. sublaevigata (Chiovenda) Cufodontis (1971). [3c] Acaulescent, rhizomatous; rhizome 2 cm ∅; L 1–3, erect or slightly spreading-recurved,

S. erythraeae Mattei (Boll. Stud. Inform. Reale Giardino Colon. [Palermo] 4: 170, 1918). Type [lecto]: Eritrea (Schweinfurth & Riva 1468 [K, FT]). — Lit: Jankalski (2004). Distr: Eritrea, N Ethiopia, Sudan; semi-arid areas along rivers, 1200–2100 m.  Dracaena erythraeae (Mattei) Byng & Christenhusz (2018); incl. Sansevieria schweinfurthii Täckholm & Drar (1954) (nom. inval., Art. 36.1); incl. Sansevieria cylindrica Schweinfurth (1894) (nom. illeg., Art. 53.1). [3b] Acaulescent, rhizomatous; L 6–8, erect, cylindrical with a short groove on the upper surface and 5 furrows on the lower surface, 40–50 cm; Inf to 50 cm, simple, elongated, fertile portion dense, 3–5 Fl per cluster; Bra ovatelanceolate, acute; Ped 7–8 mm; Fl white, tube 5–7 mm, lobes >7 mm. S. fasciata Cornu ex Gérôme & Labroy (Bull. Mus. Hist. Nat. (Paris) 9: 170, 172–173, Fig. 3, 1903). Type: Congo Free State [Democratic Republic of the Congo [Zaïre]] (Dybowski / Lecomte s.n. [P, K]). — Distr: Democratic Republic of the Congo [Zaïre].

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 Dracaena fasciata (Cornu ex Gérôme & Labroy) Byng & Christenhusz (2018); incl. Sansevieria lasciata L. Gentil (1907) (nom. inval., Art. 61.1). [3c] Acaulescent, rhizomatous; L 2–5, recurvedspreading, lanceolate, 38–84  3.8–11.5 cm, narrowed from about the middle to the channelled petiole, with a 2.1–6.4 mm green acute tip, upper surface pale green broken into patches by irregular transverse zigzag dark green bands, lower surface whitish-green with irregular narrow transverse dark green bands, margin green, becoming reddish or whitish, surface smooth; Inf and Fl not known. Regarded as a plant of uncertain status by Lebrun & Stork (2014). S. fischeri (Baker) Marais (Kew Bull. 41(1): 58, 1986). Type: Tanzania, Kilimanjaro Distr. (Fischer 9 [B?, K [fragment]]). — Distr: S Ethiopia, Kenya, Somalia, Tanzania; sandy soils at edges of thickets and in dense bush, 260–900 m. I: Rauh (1963: 126, as S. singularis).  Buphane fischeri Baker (1898); incl. Sansevieria singularis N. E. Brown (1911)  Dracaena singularis (N. E. Brown) Byng & Christenhusz (2018). [2] Acaulescent, rhizomatous; rhizome to 4.5 cm ∅; L solitary, erect, cylindrical with 4–6 longitudinal furrows, 45–240 cm, narrowed slightly upwards to near the tip, then narrowing shortly to a whitish stout acute tip, dull greyishgreen or bluish-green, brighter green with pale green transverse bands when young, surface slightly rough; Inf to 10 cm, simple, capitate, peduncle subterranean, fertile portion dense; Bra acute, 3–5  3 mm; Ped 12.7 mm; Fl tube 20–50 mm, whitish sometimes tinged with violet, lobes 5–10 mm, white with violet venation outside. In growth habit and leaves this is very similar to S. stuckyi, and as both were described without inflorescences and flowers unknown, the two taxa have been confused in cultivation. Rauh (1963) first described the inflorescence and flowers of S. fischeri, and those of S. stuckyi were described by Jumelle (1923) and Pfennig (1981), confirming that the two species are distinct. Takawira-Nyenya (2006) cites specimens from Zimbabwe, but states that it is now extinct there.

L. E. Newton

S. formosa Chahinian (Sansevieria 26: 2–5, ills., 2012). Type: Malawi (Burdett s.n. in Chahinian 319 [K, MO]). — Distr: Malawi; ecology not described, 140 m.  Dracaena malawiana Byng & Christenhusz (2018). [3c] Acaulescent, rhizomatous; rhizome 1.8–2.6 cm ∅; L 1–2 or more, margins curved upwards to form a channel, to 50  5 cm, tip obtuse, dark greyish-green, sometimes with faint green banding, margins with narrow chestnutcoloured line, surface smooth; Inf simple,  capitate, to 21 cm, fertile portion 10–15 cm ∅, 2–4 Fl per cluster; Bra triangular, 16–20  4–7 mm; Fl grey-white, with purple tinge near the base, tube 80–82  2 mm, lobes 32–33  3 mm. Examination of specimens from N Malawi (Thiede et al., in press) suggests that the leaves may be up to 100 cm long, slightly spotted or distinctly spotted. S. forskaoliana (Schultes fil.) Hepper & Wood (Kew Bull. 38(1): 83, 1983). Type: Yemen (Forskål 9 [C]). — Lit: Hepper & Friis (1994); Friis (1995). Distr: Yemen, Sudan, Djibouti, Democratic Republic of the Congo [Zaïre], Eritrea, Ethiopia, Somalia, Kenya; dry rocky slopes and river beds, usually in shade, 550–2000 m. I: Teketay (1995: 53).  Smilacina forskaoliana Schultes fil. (1829)  Dracaena forskaoliana (Schultes & Schultes fil.) Byng & Christenhusz (2018); incl. Convallaria racemosa Forsskål (1775) (nom. illeg., Art. 53.1); incl. Sansevieria guineensis Schweinfurth (1894) (nom. illeg., Art. 53.1); incl. Sansevieria guineensis var. angustior Engler (1902)  Sansevieria abyssinica var. angustior (Engler) Cufodontis (1971); incl. Sansevieria abyssinica N. E. Brown (1913)  Acyntha abyssinica (N. E. Brown) Chiovenda (1916). [3c] Acaulescent, rhizomatous; rhizome 2 cm ∅; L erect, lanceolate, 60  6.3–7.5 cm, narrowed from the middle to a channelled petiole as long as the lamina, green, with a  2.1 mm brown hardened tip, margin wavy, hard, reddish-brown, 0.1 mm, upper surface smooth, lower surface rough, minutely papillose; Inf to 95 cm, simple, elongated, fertile portion

Sansevieria RUSCACEAE

with 4–5 Fl per cluster; Ped to 10 mm; Fl white, details not known. The spelling of the epithet is still frequently seen as forskaliana, but the form here used is mandatory according to ICN Art. 60.6. Originally described from a fruiting specimen without leaves, bracts and flowers. Status still uncertain: Thulin (1995) and Mbugua (2007) include S. elliptica here, but Jankalski (2007c) treats that as a distinct species, with leaves smooth above and rough below, in contrast to S. forskaoliana with both leaf surfaces rough. However, a plant collected recently from the S. forskaoliana type locality has leaves smooth above and rough below, and there is no mention of leaf surfaces in the protologue. Further studies of field-collected material are needed to resolve this. S. francisii Chahinian (Sansevieria J. 4(1): 12–14, ills., 1995). Type: Kenya, Coast Prov. (Horwood 432 [MO, UPS, ZSS]). — Distr: S Kenya (Coast Prov., Tana River Distr.); ecology not described. I: Speirs (1984: as S. sp); Mansfeld & Rosigkeit (2016).  Dracaena francisii (Chahinian) Byng & Christenhusz (2018). [3a] Caulescent, with runners; stem erect, to 30 cm; runners to 16  0.8–1.4 cm; L to 40, spiralled in 5 rows, cylindrical with a groove on the upper surface for of the length, to 8–15 cm, narrowed gradually to a 5 mm acute spine-like tip, dark green with grey-green transverse bands and 4–6 dark green longitudinal lines, margin brownish-red with a white edge on the basal , green above, surface slightly rough; Inf 12–25 cm, simple, elongated, fertile portion dense, 1–2 Fl per cluster; Bra triangular,  6  3 mm; Ped  2 mm; Fl greenish-white to brownish-green, tube 16–19 mm, lobes 8–10 mm. S. frequens Chahinian (Cact. Succ. J. (US) 72 (3): 130–132, ills., 2000). Type: Kenya, Laikipia Distr. (Chahinian 785 [MO, NY]). — Lit: Mbugua (1998: as S. braunii). Distr: Ethiopia, Kenya, Uganda; mostly rocky areas in grassland or open bush, 600–1750 m.  Dracaena frequens (Chahinian) Byng & Christenhusz (2018).

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[3c] Acaulescent, rhizomatous; rhizome to 5 cm ∅, yellow-buff coloured; L 4–8 (usually 6), rosulate, erect, leathery, oblanceolate, 90 or more 15 cm, dull grass-green, sometimes with a bloom, base slightly narrowed, tip obtuse, margins fibrous, chestnut-brown, surface smooth; Inf 60–90 cm, simple, elongated, peduncle green, fertile portion dense; Bra 3.35–8  2.5–3 mm; Ped shorter or longer than the bracts; Fl greenishwhite, tube 18–20 mm, lobes 26–28 mm. S. gracilis N. E. Brown (Bull. Misc. Inform. Kew 1911: 96, 1911). Type: Kenya, Coast Prov. (Powell 11 [K]). — Distr: Kenya, Tanzania; dry bushland and thicket, 30–600 m. I: Brown (1915: 204).  Dracaena serpenta Byng & Christenhusz (2018). [3a] Caulescent, with runners; stem erect, to 8 cm; runners slightly ascending, to 90  0.8 cm; L 8–12, densely spiralled, ascending or spreading, cylindrical with a groove on the upper surface for up to 12.5 cm from the base, 23–80 cm, to 0.6–1 cm thick, deep grass-green, sometimes with inconspicuous narrow transverse darker bands and slightly darker longitudinal lines, narrowed gradually to a 2.1–6.4 mm brown or whitish spine-like tip, margin membranous, white, surface smooth; Inf to 30 cm, simple, elongated, peduncle light green, fertile portion lax, 2 Fl per cluster; Bra lanceolate or linearlanceolate, acute, 2.1–3.2 mm; Ped 1–1.6 mm; Fl white, tube 19–25 mm, lobes 10.5–12.7 mm. Widespread in the Coast Province, but apparently extinct at the type locality, Mazeras, which has grown into a large town. Collections referred to as var. humbertiana by Mbugua (2007) are S. ballyi. S. gracillima Chahinian (Sansevieria 12: 3–4, ills., 2005). Type: Somalia, Bay Prov. (Lavranos & Bauer 27839 in Chahinian 336 [MO, NY]). — Distr: S Somalia; dense semi-deciduous forest on granite inselberg, 600 m.  Dracaena gracillima (Chahinian) Byng & Christenhusz (2018); incl. Sansevieria gracilis var. somaliensis Chahinian (2005) (nom. inval., Art. 36.1, 37.1).

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[3a] Acaulescent, with runners; runners 7–10 cm; L 8–10, densely spiralled, slightly spreading, cylindrical with shallow groove on the upper surface to the leaf length, 6–22  0.9 cm, base 9 mm thick, with spine-like tip, dark green with conspicuous bands of lighter green, edge of the groove reddish, surface smooth; Inf to 17 cm, simple, elongated, peduncle dark green, with 3 sterile bracts, fertile portion lax, 1–3 Fl per cluster; Bra triangular, 2  2 mm; Ped 2 mm; Fl brownish, tube 6–7 mm, lobes 14–15  2 mm. Described as similar to S. gracilis, but with shorter leaves and shorter flowers, which are a different colour. S. grandicuspis Haworth (Synops. Pl. Succ., 67, 1812). Type: not typified. — Distr: Unknown. Incl. Sansevieria ensifolia Haworth (1812); incl. Sansevieria pumila Haworth (1812). [3c] Acaulescent, rhizomatous; L 5–15, erect or ascending-spreading, linear-lanceolate, 18–50  1.25–3.8 cm, sometimes narrowed to a channelled petiole to 15 cm, with alternating transverse bands of dull dark and lighter green, with 5–7 longitudinal dark green impressed lines, shortly narrowed above to a 17–50 mm green flexible subulate tip, margin green, surface smooth; Inf and Fl not known. Originally described with flowers not seen, with no illustration or herbarium specimen, and still little-known. Possibly better treated as a name of unresolved application. Included in S. zeylanica by Chahinian (2005a) and Govaerts (2014+), and in S. aethiopica by Mansfeld (2015a). A more likely candidate is S. roxburghiana, which has long subulate leaf tips. The specimen referred to this species by Mbugua (2007) is S. dooneri (Newton & Webb 2011). S. hallii Chahinian (Sansevieria J. 5: 7–10, ills., 1996). Type: Zimbabwe, East Region (Hall 67/799 [MO, UPS]). — Lit: Rulkens & Baptista (2009b); Jaarsveld & Condy (2015). Distr: Moçambique, Zimbabwe, RSA (N Limpopo); rocky outcrops, 300–360 m. I: Jaarsveld (2016).  Dracaena hallii (Chahinian) Byng & Christenhusz (2018).

L. E. Newton

[2] Acaulescent, rhizomatous; rhizome 1.8–3 cm ∅, grey-orange; L 1–3, erect, cylindrical with a deep groove on the upper surface, to 60  5 cm, dark greyish-green with numerous longitudinal lines, with inconspicuous transverse bands when young, narrowed to a 6 mm obtuse or rounded tip, margin chestnut-brown edged white, surface rough; Inf to 18 cm, capitate, fertile portion to 16 cm ∅, dense; Bra triangular, 18  6 mm; Fl white tinged purple, tube 45–78 mm, lobes 25–29 mm. S. hargeisana Chahinian (Sansevieria J. 3(3): 53–55, ills., 1994). Type: Somalia, North-Central Region (Lavranos 7382 [MO, UPS]). — Distr: Somalia; limestone plateau with sparse vegetation, 1200 m.  Dracaena hargeisana (Chahinian) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; L 4–8, rosulate, spreading, cylindrical with a basal adaxial groove for of the length, with a white membranous margin, 8–20 cm, base 10–14 mm thick, tip cuspidate with brown band at the base, dark green with lighter green transverse bands and 5–7 dark green longitudinal lines, surface rough; Inf 14–18 cm, simple, peduncle with 3 sterile bracts, fertile portion 4.5–6 cm, semidense, 2–4 Fl per cluster; Ped 5 cm ∅, light straw-coloured, darker grey when older; L 4–6 (–15), linear, flat, 60–140  6–13 cm, uniformly green, margin red with white outer edge, tip narrowing for 5 cm to a sharp point, with the margin inrolled; Inf 70–130 cm, simple, peduncle 30 cm, with 4–5 broadly lanceolate acute bracts 4–6  1.4 cm, green, fertile portion 28–55 cm, 4–6 Fl per cluster, clusters 1 cm apart; Bra acute, 12–19  4–7 mm, papery; Ped 9–18 mm; Fl whitish, tube 64–95 mm, lobes 17–20 mm. Described from a single incomplete herbarium specimen, but more information, including comments on variation in the field, is provided by Rulkens & Baptista (2009a). Close to S. conspicua. S. perrotii Warburg (Tropenpflanzer 5: 190, ill. (p. 191), 1901). Type: Tanzania, Lindi Prov. (Perrot s.n. [not located]). — Lit: Newton (2014). Distr: Tanzania; among bushes on coral.  Dracaena perrotii (Warburg) Byng & Christenhusz (2018). [1] Caulescent; stem erect, to 20  2.5 cm, covered by leaf bases; L 8–12, distichous, ascending or spreading, cylindrical, laterally slightly compressed, with a deep and wide groove on the upper surface, 91–152 cm, 1.1–1.5 cm thick, narrowed gradually to a whitish hard acute tip, margin reddish-brown, edged white; Inf 1.2 m, paniculate, fertile portion lax, 2–4 Fl per cluster;

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Fl tube 1.27 mm, pale greenish, lobes 9.5 mm, whitish inside, purplish outside. Leaves described by Jankalski (2007a) as pale bluish-glaucous. Included in S. robusta by Mbugua (2007), though S. perrotii is an earlier name and would have priority if the two were considered to be conspecific (Newton 2014). A plant collected at the Kenya coast by Mr. Tony Dyer is possibly this species. S. pfisteri D. J. Richards (Sansevieria 19: 4–7, ills., 2009). Type: Angola (Downs & Pfister s.n. [SRGH]). — Distr: SW Angola; hot stony desert, below limestone ridges.  Dracaena pfisteri (D. J. Richards) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; rhizome 1.5–3 cm ∅, light brown; L 8–10 (–15), distichous, erect when young, spreading when displaced by new growth, to 64 cm, cylindrical with adaxial groove 2 cm wide  1.5 cm deep from the base to nearly of the leaf length, dull dark green with faint transverse paler green bands, adaxial groove with brownish margin, with 7 or more very shallow longitudinal grooves with broken dark lines, surface smooth; Inf 64 cm, simple, elongated, peduncle pale green with minute whitish linear marks, fertile portion semidense, 6–10 Fl per cluster; Fl dull yellow, purplish brown at the tip, tube 20–25  3 mm, lobes 10 mm. S. phillipsiae N. E. Brown (Hooker’s Icon. Pl. 30: t. 3000 + text, 1913). Type: Somalia (Lort Phillips s.n. [K]). — Distr: S & E Ethiopia, Somalia; in the shade of trees, 1250–1450 m.  Dracaena phillipsiae (N. E. Brown) Byng & Christenhusz (2018). [3a] Caulescent, in clumps to 38 cm tall, with runners; stem erect, branching at or above the base; runners to 20  1.3 cm; L 5–10, rosulate, ascending, becoming spreading, cylindrical with a deep groove on the upper surface for 5–8.9 cm at the base, 10.2–46 cm, 1.3–1.9 cm thick, narrowed gradually to a 2.1–3.2 mm brown hard acute or obtuse tip, dark and slightly bluish-green, with faint transverse paler green bands when young, margin white, surface smooth; Inf 35–46 cm, simple, elongated, fertile portion 23–30 cm, 3–6 Fl per

L. E. Newton

cluster; Bra ovate-lanceolate, acute, 3.2–6.4 mm; Ped 2.65–3.18 mm; Fl white, tube 10 mm, lobes 11.6–12.7 mm. S. pinguicula P. R. O. Bally (Candollea 19: 145–147, ills., 1964). Type: Kenya, Coast Prov. (Bally 4275 [K]). — Lit: Newton & Thiede (2015). Distr: Kenya.  Dracaena pinguicula (P. R. O. Bally) Byng & Christenhusz (2018). S. pinguicula fa. disticha (Pfennig ex Butler) L. E. Newton & Thiede (Cact. Succ. J. (US) 87(1): 31, ills. (pp. 30–31), 2015). Type: Kenya, Coast Prov. (Bally 14976 [EA]). — Distr: Kenya (Coast Prov.); sandy plains with open bushland, 120–230 m.  Sansevieria pinguicula ssp. disticha Pfennig ex Butler (2012)  Dracaena pinguicula ssp. disticha (Pfennig ex A. Butler) Byng & Christenhusz (2018). [1] Differs from ssp. pinguicula: L 10–15 cm, distichous, curving upwards. The type locality is not far from that of ssp. pinguicula, and the two variants have been found growing together, so the rank of subspecies, chosen upon transfer of the species to Dracaena by Christenhusz & al. (2018), is inappropriate as the rank of subspecies is usually used for geographically separated populations. A specimen in ZSS, cited in the protologue as an isotype, has a different date from that of the holotype and cannot be regarded as a duplicate (ICN Art. 8.2, 9.4). Hence it is regarded as a paratype. As the holotype does not show the distichous leaf arrangement, an epitype was designated (Newton & Thiede 2015). S. pinguicula ssp. nana (Chahinian) L. E. Newton & Thiede (Cact. Succ. J. (US) 87(1): 32, ills. (p. 31), 2015). Type: Kenya, Coast Prov. (Schwartz & LaFon s.n. in Chahinian 397 [MO, NY]). — Distr: Kenya (Coast Prov.); open bushland, 150 m.  Sansevieria pinguicula var. nana Chahinian (2013)  Dracaena pinguicula ssp. nana (Chahinian) Byng & Christenhusz (2018). [1] Differs from ssp. pinguicula: L shorter, darker green; Fl white.

Sansevieria RUSCACEAE

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Fig. 5 Sansevieria pinguicula ssp. pinguicula. (Copyright: U. Eggli)

A compact, dwarf-growing variant of S. pinguicula, growing some distance away from the type locality of the species. S. pinguicula ssp. pinguicula — Distr: Kenya; sandy plains with open bushland, 120–230 m. I: Pfennig (1977a: 510). – Fig. 5. [1] Shortly caulescent, with runners; stem erect; runners to 8 cm; L 5–7, rosulate, cylindrical with a deep groove on the upper surface and 2–7 narrow grooves on the lower surface, 12–30 cm, 2.8–3.5 cm thick, narrowing to a horny acute tip, green, margin brown, surface slightly rough; Inf 15–32 cm, paniculate, fertile portion dense, 4–6 Fl per cluster; Ped 1.5–2 mm; Per tube 4–5 mm, lobes 3–4 mm, white with brown mid-stripe. Chahinian (2005a) states that plants have rosulate leaves in juvenile stages, and distichous leaves when mature. Plants growing and spreading in my garden for over 20 years still have rosulate leaves, but see fa. disticha above. S. powellii N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 198, Fig. 1, 1915). Type: Kenya, Coast Prov. (Powell 5 [K]). — Distr: Kenya, Somalia; shade in thickets. I: Sansevieria J. 3(1): 12–13, 1994. – Fig. 6.  Acyntha powellii (N. E. Brown) Chiovenda (1932)  Dracaena powellii (N. E. Brown) Byng & Christenhusz (2018). [1] Caulescent, rhizomatous; stem erect, to 1.2 m  2.5 cm; L distichous, the ranks becoming twisted around the stem, spreading,

Fig. 6 Sansevieria powellii. (Copyright: L. E. Newton)

semicylindrical, 30–69  2.2–2.9 cm,  1.3 cm thick, narrowed gradually to a pale brown hard spine-like acute tip, faintly glaucous grass-green becoming dark bluish-green, margin red-brown edged white, surface slightly rough; Inf  46 cm, paniculate, fertile portion subdense, 4–6 Fl per cluster; Bra convex, fleshy, 1.1–2.1 mm;

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Ped  3.2 mm; Fl dull greenish-white, with dull brownish-purple slender lines outside, tube 6.4 mm, lobes 9.5 mm. Pfennig (1977a) has suggested that this might be a natural hybrid S. arborescens  S. robusta, but this has yet to be confirmed (Newton 2012a; Butler 2012a). S. powysii L. E. Newton (Bradleya 28: 23–25, ills., 2010). Type: Kenya, Coast Prov. (Powys 1250 [K, EA]). — Distr: S Kenya (Coast Prov.); in dense costal bushland.  Dracaena powysii (L. E. Newton) Byng & Christenhusz (2018). [1] Caulescent, rhizomatous; rhizome 2.6–3 cm ∅, pale yellow; stem erect, to 2 m; L spiral, spreading, becoming deflexed, triangular, to 43  2 cm, 1.7–2 cm thick, canaliculate above, narrowed to pungent 5 mm brownish spine-like tip, upper surface dull green, smooth, lower surface mid-green with 15 darker longitudinal lines in the lower , very slightly rough, margin 1 mm, red-brown with whitish fraying edge; Inf to 85  45 cm, paniculate with up to 20 spreading-ascending branches, lower ones to 35 cm with secondary branching, fertile portion subdense, 4–6 Fl per cluster; Bra rounded with acute apex, 1  1.5 mm, with brownish base and white apex; Ped 2 mm, pale green; Fl yellowish with brown mid-stripe on the lobes, tube 5–6  2.5 mm, lobes to 7  1.5–2 mm. Jankalski (2009c) suggested that this might be the plant in Somalia described as S. powellii by Thulin (1995), but Thulin described the leaves as “in 2 ranks”, which fits S. powellii but not S. powysii, and he did not mention the very thick leaves as in S. powysii. S. raffillii N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 252, Fig. 22, 1915). Type: Kenya, Coast Prov. (Powell 7 [K]). — Distr: Kenya, Tanzania, Uganda.  Dracaena raffillii (N. E. Brown) Byng & Christenhusz (2018). S. raffillii var. glauca N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 252–254, Fig. 22, 1915). Type: Kenya, Coast Prov. (Powell 8 [K]). — Distr: Kenya.

L. E. Newton

[3c] Differs from var. raffillii: L very dark bluish-green with distinct, but not very conspicuous, irregular spots or wavy transverse bands of lighter green 2.5–5 cm apart, distinctly bluishglaucous; Inf peduncle bluish-glaucous below, lighter green speckled with pale green above; Bra linear-lanceolate, acute, 8.5–25.4 mm; Ped slightly glaucous, 5.3–6.4 mm. Apparently collected only once, and possibly only a minor variant of the species. S. raffillii var. raffillii — Distr: Kenya, Tanzania, Uganda; Acacia bushland, 900–1700 m. I: Sansevieria J. 2(2): 36–37, 1993. [3c] Acaulescent, rhizomatous; rhizome 2–5 cm ∅, whitish; L 1–2, erect, lanceolate or lorate, 68–152  5.5–12.5 cm, narrowed below the middle to a sessile base or short petiole, with a short reddish-brown hard tip, with yellowishgreen closely placed blotches or irregular transverse bands on darker background, sometimes paler on the lower surface, slightly glaucous, with age with less conspicuous markings, margin hard, reddish-brown, surface smooth; Inf 90–115 cm, simple, elongated, fertile portion 61–76 cm, dense, 2–5 Fl per cluster; Bra ovate-lanceolate, acuminate, 5.3–17 mm; Ped 4.2–6.4 mm; Fl tube 25–28.6 mm, greenish-white, lobes 28.6–29.6 mm, white. S. robusta N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 207, 1915). Type [lecto]: Kenya, Taita Distr. (Grenfell 13 [K]). — Lit: Chahinian (2005b); Newton (2014). Distr: Kenya; dry bushland, 600–1500 m. I: Mbugua (1994); Newton (1994). [1] Caulescent, rhizomatous; rhizome to 2 cm ∅, yellowish-white; stem erect, to 60  2.5 cm; L 6–14, distichous, erect or slightly spreading, laterally compressed with a wide groove along the upper face, to 2.13 m  3.4 cm, to 4.5 cm thick, narrowed gradually to a 6.4–12.7 mm brown hard spine-like tip, dark green, faintly glaucous, with 14–30 longitudinal darker green lines, margin narrowly bordered red-brown, edged white, surface smooth; Inf 80–140 cm, paniculate, peduncle greenish-grey, fertile portion 12–20 cm, lax, 4–6 Fl per cluster; Ped

Sansevieria RUSCACEAE

11–19 mm; Fl white or greenish, tube 10–25 mm, lobes 5–10 mm. Originally described with flowers unknown, and based on 4 syntypes, all from Taita District. A lectotype was chosen by Mbugua (2007). Mbugua (2007) and Mansfeld (2015a) included S. perrotii here and this was followed by The Plant List, after correcting Mbugua’s failure to recognize its priority over S. robusta. The synonymization was disputed by Newton (2014). When growing in thick bush it can grow taller, but in open situations it does not exceed 60 cm. Chahinian (2005b) recorded variation in the species, and further field work is required to investigate this.

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irregular whitish-green elongated blotches, margin narrow, white and sometimes detached as a fibre, with thin inner red line, surface slightly rough; Inf 74–92 cm, simple, peduncle mid-green; fertile portion 55–69 cm, dense, with (1–) 4–8 (–9) flowers per cluster; Bra linearlanceolate, attenuate, 1.4–2.8  0.4–0.8 mm, green with white margin, with 3–5 darker nerves; Ped 3–5 mm; Fl white, tinged light green at base and apex, tube 30–42 mm, lobes 25–28 mm.

S. rorida (Lanza) N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 205, 1915). Type: Somalia (Macaluso 177 [PAL?]). — Lit: Mangani (1996: with ills.). Distr: Somalia; sandy places on the coast near sea-level.  Sanseverinia rorida Lanza (1910)  Acyntha rorida (Lanza) Chiovenda (1916). [1] Caulescent; stem to 23 cm; L 11–15, distichous, ascending-spreading, cylindrical with a groove along the upper surface, 30–53  2.5–3.4 cm, 1.9–2.5 cm thick, narrowed gradually to a spine-like tip, green, somewhat glaucous, with numerous darker longitudinal lines on the sides and lower surface, margin reddish-brown, edged white; Inf  1 m, paniculate, fertile portion 7.6–18 cm, lax, 3–6 Fl per cluster; Bra deltoid, acute; Ped  2 mm; Fl tube 6.4 mm, lobes 10.6–12.7 mm, whitish-yellow with reddish mid-stripe. Included in S. ehrenbergii by Thulin (1995) and Mansfeld (2015a), and mentioned under that species by Lebrun & Stork (2014).

S. roxburghiana Schultes fil. (Syst. Veg. 7: 357, 1829). Type: [icono]: Roxburgh, Pl. Corom. 2: 43, t. 184, 1805, as S. zeylanica. — Lit: Budweg & Mansfeld (2014: history); Mansfeld & Ott (2015). Distr: India (Coromandel Coast, Goa, Karnataka, Maharashtra, Tamil Nadu).  Acyntha roxburghiana (Schultes fil.) Kuntze (1891)  Cordyline roxburghiana (Schultes fil.) Merrill (1923)  Dracaena roxburghiana (Schultes fil.) Byng & Christenhusz (2018); incl. Sansevieria zeylanica Roxburgh (1805) (nom. illeg., Art. 53.1). [3c] Acaulescent, rhizomatous; L 6–24, rosulate, ascending, slightly recurved, linear with a deep groove on the upper surface, 20–60  1.3–2.5 cm, narrowed gradually to a 6.4–50 mm green soft tip, green with darker green irregular transverse bands and dark green longitudinal lines, margin green, becoming whitish with age, upper surface smooth, lower surface slightly rough; Inf 30–78 cm, simple, elongated, fertile portion 25–38 cm,  4 Fl per cluster; Bra lanceolate, attenuate, 3.2–4.2 mm; Ped 5.3–8.5 mm; Fl whitish-green, tube 6–8 mm, lobes 8.5–9.5 mm. Cytology: 2n = 40 (Darlington & Wylie 1955).

S. rosulata T. G. Forrest (Cact. Succ. J. (US) 89(6): 289–292, ills., 2017). Type: Uganda, Bukedea Distr. (Forrest 1506 [MHU]). — Distr: E Uganda; rock outcrops, sometimes on large ant hills, 1160 m. [3c] Acaulescent, rhizomatous, rhizome 4 cm ∅, light yellow/brown; L 4–12 (–16), rosulate, spreading, lanceolate, 60–100 (–104)  7–8 (–10) cm, base narrowing slightly, rarely petiolate, tip attenuate-subulate, 3–6 cm, green with

S. sambiranensis H. Perrier (Notul. Syst. (Paris) 5(2): 154–156, 1935). Type [syn]: Madagascar, Sambirano Distr. (Perrier 8379/8380 [P]). — Distr: Madagascar; shaded rocks in humid forest. I: Perrier (1938: 5); Vrskovy (2006); Rzepecky (2013).  Dracaena sambiranensis (H. Perrier) Byng & Christenhusz (2018). [2] Acaulescent, rhizomatous; rhizome to 30  1–2 cm; L 15–20, laxly rosulate,

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oblanceolate, 120  2.5–9 cm, narrowed below the middle to a 10–20 cm channelled petiole, narrowed to an acute tip, green; Inf 2–8 cm, capitate, fertile portion 4–5 cm ∅, dense; Bra lanceolate, acute, 6–7 mm; Ped 7–20 mm; Fl carminered, tube 10 mm, lobes 20–22 mm. The species was long lost, but 2 collections from 1987 and 1998 have recently been reported (Rzepecky 2013). A preliminary DNA study suggests that this taxon does not belong in Sansevieria or Dracaena (Mansfeld 2014a). S. scimitariformis D. J. Richards (Sansevieria 5: 8–9, ills., 2003). Type: Zimbabwe (Richards 2259 [SRGH]). — Distr: NE Zimbabwe; amongst rocks on granite hill, c. 890 m.  Dracaena scimitariformis (D. J. Richards) Byng & Christenhusz (2018). [2] Acaulescent, rhizomatous; rhizome 3–4 cm ∅; L 4–6, rosulate, spreading, flat-canaliculate, 50–70  6–10 cm, base 3 cm thick, tip blunt, overall dark green, sometimes with lighter transverse bands, margin slightly wavy, brown; Inf 15 cm, simple, peduncle with 4 sterile Bra, fertile portion 10 cm, dense, almost capitate, 4–6 Fl per cluster; Bra 20 mm; Fl purplish at the base, becoming lighter and yellowish-green above, 100–120 mm. The protologue describes the “floral tube” as 10–12 cm long, but as corolla lobes are not mentioned this probably refers to the whole flower. Croix (2010) suggests a possible occurrence in Zambia. This species and S. humiflora are close to S. hallii. Further field work to determine distribution and variation could lead to a reassessment of their distinction as species. S. senegambica Baker (J. Linn. Soc., Bot. 14: 548, 1875). Type [syn]: Senegal (Perrottet 782 [K]). — Distr: Gambia, Guinea, GuineaBissau, Ivory Coast, Senegal, Sierra Leone; shady places. I: Brown (1915: 236).  Dracaena senegambica (Baker) Byng & Christenhusz (2018); incl. Sansevieria cornui Gérôme & Labroy (1903); incl. Sansevieria liberiensis Cornu ex A. Chevalier (1920) (nom. inval., Art. 32.1d?).

L. E. Newton

[3c] Acaulescent, rhizomatous; rhizome 1.3–1.9 cm ∅, bright red, turning pale brownish in light; L 2–4, rosulate, suberect, recurved to spreading nearer the tip, linear-lanceolate to lanceolate, 30–69  3–6.4 cm, narrowed at the base to a 2.5–7.6 cm channelled petiole, narrowed gradually from the middle to a 4.2–12.7 mm green soft subulate tip, upper surface dark green sometimes with indistinct paler green transverse bands, lower surface slightly paler with more distinct transverse bands, margin green, surface smooth; Inf 30–50 cm, simple, elongated, peduncle light or dark green or mottled purplish, fertile portion lax, 3–6 Fl per cluster; Bra ovate-lanceolate or oblong-lanceolate, acute, 6.4–8.5  2.1–3.2 mm; Ped 5.3–8.5 mm; Fl white, tinged purple in sun, tube 6.4–12.7 mm, lobes 10.6–19 mm. — Cytology: 2n = 40 (Sharma & Chaudhuri 1964). Based on 2 syntpes, the other being Richard 72 (K). S. sinus-simiorum Chahinian (Sansevieria 3: 24–26, ills.; 4: 9 [erratum], 2002). Type: Malawi (Burdett s.n. in Chahinian 316 p.p. [MO, NY]). — Lit: Thiede & al. (2017). Distr: S Malawi; in shade on rocky slopes.  Dracaena sinus-simiorum (Chahinian) Byng & Christenhusz (2018); incl. Sansevieria malawiana hort. (s.a.) (nom. inval., Art. 29.1). [2] Acaulescent, rhizomatous, rhizome 5 cm ∅, grey; L 8–10, crowded, straight or slightly curved backwards or sideways, occasionally twisting, with a round channel  the width of the leaf at the bottom and widening to sometimes almost the full width of the leaf at the top, to 100–150 cm long, to 6 cm thick, 6 cm wide at the base, slowly tapering to the tip, overall with inconspicuous cross-banding when young, more obscure later, with numerous continuous and interrupted lines, most reaching the apex, margin (edge of channel) acute, chestnut-brown bordered by withered fibres, tip obtuse, surface smooth, waxy; Inf simple, capitate, peduncle 29 cm, 16 mm wide at the base, 13 mm at mid-height, with 6 sterile triangular Bra, withered, 32–35  11.13 mm; fertile portion 19 cm ∅, dense; Bra membranous, 4–8  2–4 mm; Ped 6–16 mm; Fl white tinged with green, when

Sansevieria RUSCACEAE

closed 30–32 mm, 2 mm wide at the tip, 1 mm wide at the bottom, when open tube cylindrical, 82–97 mm, 2 mm wide at the top, 3 mm wide at mid-length, inflated to 4 mm at the bottom, lobes 19–21  3 mm. The hyphen in the epithet was omitted by Christenhusz & al. (2018), contrary to ICN Art. 60.9. S. sordida N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 214–215, Fig. 8, 1915). Type: K [ex cult.]. — Distr: Unknown.  Dracaena sordida (N. E. Brown) Byng & Christenhusz (2018). [3b] Acaulescent or shortly caulescent, rhizomatous; L 4–12, distichous, slightly spreading, cylindrical, slightly compressed laterally with a channel down the upper surface, 69–107  0.9–1.3 cm, 1.3–1.9 cm thick, becoming flattened towards the base, narrowed to a 0.7–1.1 mm whitish or grey brown-based acute spine-like tip, dull bluish-green with numerous darker longitudinal lines, margin hardened, dark brown edged white, surface very rough; Inf 30–60 cm, simple, elongated, peduncle dull light green, with minute white dots, fertile portion 20–45 cm, lax, 7–14 Fl per cluster; Bra subulate, 3.2–6.4 mm; Ped 8.5–12.7 mm; Fl tube 7.4–10.6 mm, white or greenish, lobes 14.8–16.9 mm, white inside, green with minute dull purplish dots outside. I am not aware of any recent field gatherings matching this description, but Mbugua (2007)

Fig. 7 Sansevieria stuckyi. (Copyright: U. Eggli)

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claims a disjunct distribution in Kenya, Zambia and RSA. S. specksii R. H. Webb & Myklebust (Sansevieria 37: 7–10, ills., 2018). Type: Benin, Atakora (Specks 21430 [MO]). — Distr: W Benin; savanna, 720 m. [3c] Acaulescent, rhizomatous, forming dense clumps, rhizome 5 cm ∅; L (2–) 3 (–5), ascending to spreading, oblanceolate, canaliculate, (70–) 90 (–110)  6–7 cm, medium-green with light grey-green irregular transverse bands, with pronounced U-shaped central channel bearing longitudinal lines that are more prominent towards the base, with thin red-brown entire margin, upper surface smooth, lower surface moderately rough, tip 5 mm, white, chartaceous; Inf 90 cm, simple, fertile portion to 45 cm, dense, with (4–) 6 (–8) Fl per cluster; Bra triangular, scarious, 5  2 mm; Ped 1–2 mm; Fl cream, 80–100 mm. S. stuckyi Godefroy-Lebeuf (Sansev. Gigant. Afr. Orient., 13, 17, 33, ills., 1903). Type: Moçambique (Godefroy-Lebeuf s.n. [K]). — Distr: Moçambique, Zimbabwe. I: Brown (1915: 220). – Fig. 7.  Dracaena stuckyi (Godefroy-Lebeuf) Byng & Christenhusz (2018); incl. Sansevieria andradae Godefroy-Lebeuf (1903). [2] Acaulescent, rhizomatous; rhizome to 5 cm ∅; L 1–2 (rarely 3), erect, cylindrical with a groove along the upper surface, 122–275 

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3.8–6.4 cm, narrowing gradually to a pale brown hard acute subulate tip, dull green, slightly glaucous, with transverse paler green bands and 6–20 longitudinal lines, margin green, surface slightly rough; Inf 21–32 cm, simple, capitate, peduncle purple, speckled green, fertile portion 7  34 cm at anthesis, dense; Bra ovate-lanceolate, 8–20  4–10 mm; Fl tube 90–100 mm, lobes 40 mm. — Cytology: 2n = 116 (Sharma & Chaudhuri 1964). Originally described with flowers unknown. See comment for S. fischeri. Reported by TakawiraNyenya (2006) to be extinct in Zimbabwe. S. subspicata Baker (Gard. Chron., ser. 3, 6: 436, 1889). Type: Moçambique, Gaza Prov. (Monteiro s.n. [K]). — Distr: Moçambique; coastal thickets.  Dracaena subspicata (Baker) Byng & Christenhusz (2018); incl. Sansevieria splendens A. B. Graf (1978) (nom. inval., Art. 36.1, 37.1); incl. Sansevieria patersonii hort. ex Morgenstern (1979) (nom. inval., Art. 36.1, 37.1). [3c] Acaulescent, rhizomatous; rhizome 1.3–2.5 cm ∅, pale yellowish-brown; L 4–10, erect or recurved-spreading, lanceolate, 23–60  2.5–5.7 cm, narrowed from the middle or above to a 3.8–23  0.4–0.6 cm channelled petiole, with a 4.2–6.4 mm green soft subulate tip, deep green, sometimes faintly glaucous, margin green becoming whitish, surface smooth; Inf 30–40 cm, simple, elongated, fertile portion lax, 1–2 Fl per cluster; Bra lanceolate, acute, 2.1–6.4 mm; Ped 1.1–2.1 mm; Fl tube 23–30 mm, greenish-white, lobes 17–19 mm, white. S. subtilis N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 237–238, Fig. 17, 1915). Type: Uganda (Dawe s.n. [K]). — Distr: Uganda.  Dracaena subtilis (N. E. Brown) Byng & Christenhusz (2018). [3c] Acaulescent, rhizomatous; rhizome 0.85–1.1 cm ∅; L 2–4, erect or slightly recurved, linear-lanceolate, 53–69  2.5–4.5 cm, narrowed to a 5–30 cm channelled petiole, narrowed gradually from the middle or above to a 12.7–25.4 mm green soft subulate tip, green, lower surface sometimes with faint transverse bands, margin green,

L. E. Newton

surface smooth; Inf 38–53 cm, simple, elongated, fertile portion lax, 2–3 Fl per cluster; Bra lanceolate, acuminate, 3.2–4.2 mm; Ped 4.2–7.4 mm; Fl tube 6.4–8.5 mm, lobes 10.6–12.7 mm, white. S. suffruticosa N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 202, Fig. 3, 1915). Type: Kenya (Evans s.n. [K]). — Lit: Jankalski (2007b); Webb & Newton (2016). Distr: Kenya.  Dracaena suffruticosa (N. E. Brown) Byng & Christenhusz (2018). S. suffruticosa var. longituba Pfennig (Bot. Jahrb. Syst. 102 (1–4): 178, 1981). Type: Kenya, Nairobi Distr. (Pfennig 1336 [EA]). — Distr: Kenya; edges of thickets and on sandy cliffs, 1450 m. I: Chahinian (1992).  Dracaena suffruticosa var. longituba (Pfennig) Byng & Christenhusz (2018). [3a] Differs from var. suffruticosa: L surface very smooth; Fl tube 20 mm. S. suffruticosa var. suffruticosa — Distr: Kenya; edges of thickets, 1700–1900 m. [3a] Caulescent, with runners; stem erect, to 30 cm, branching freely 0.8–7.6 cm above ground; runners spreading or ascending, to 25  1.9 cm; L 7–18,  distichous or irregular, ascending or spreading, cylindrical with a groove on the upper surface from the base to – of the leaf length, 15–60 cm, 1.3–1.9 cm thick, narrowed gradually to a 3.2–4.2 mm brown hard acute spine-like tip, dark green with faint paler green transverse bands when young, with darker green longitudinal lines, surface rough, sometimes smooth near the base; Inf 30–38 cm, simple, elongated, peduncle green with numerous whitish minute linear dots, fertile portion 4.5–6.4 cm ∅, dense, 2–5 Fl per cluster; Bra ovate, acute or acuminate, 1.1–4.2 mm; Ped 2.8–4.2 mm; Fl whitish or greenish-white, sometimes slightly red-tinged outside, tube 10 mm, lobes 12.7–14.8 mm. Pfennig (1981) argued for a type locality near Nakuru. S. trifasciata Prain (Bengal Pl. 2: 1054, 1903). Type: Nigeria, Oban Distr. (Talbot s.n. [K]). — Lit:

Sansevieria RUSCACEAE

Chahinian (1987: synopsis cultivars); Chahinian (2012: updated synopsis cultivars); Chahinian (2013: updated synopsis cultivars); Chahinian (2014: updated synopsis cultivars); Mangani (2014: history). Distr: Nigeria, Democratic Republic of the Congo [Zaïre]; naturalized elsewhere (e.g. India).  Dracaena trifasciata (Prain) Mabberley (2017). Widely cultivated in numerous cultivars, see Chahinian (1987) and later updates. A dwarf cultivar (leaves to 50  7.5 cm) is ‘Hahnii’ and several derivatives with different leaf sizes, different degrees of variegation, and different shades of green exist. S. trifasciata var. laurentii (De Wildeman) N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 240, 1915). Type: Democratic Republic of the Congo [Zaïre] (Anonymus s.n. [K]). — Distr: Democratic Republic of the Congo [Zaïre]. I: Chahinian (1987: 21).  Sansevieria laurentii De Wildeman (1904). [3c] Differs from var. trifasciata: L with yellow margin to 1 cm wide. — Cytology: 2n = 40 (Sharma & Chaudhuri 1964: as S. laurentii). A periclinal chimaera, treated as a cultivar (S. trifasciata ‘Laurentii’) by Jankalski (2007a). S. trifasciata var. trifasciata — Distr: Nigeria. I: Chahinian (1987: 20). Incl. Aletris hyacinthoides Miller (1768) (nom. illeg., Art. 53.1); incl. Acyntha guineensis Medikus (1786); incl. Salmia guineensis Cavanilles (1795); incl. Sansevieria guineensis Gérôme & Labroy (1903) (nom. illeg., Art. 53.1); incl. Sansevieria zebrina L. Gentil (1907) (nom. inval., Art. 32.1c); incl. Sansevieria jacquinii N. E. Brown (1911). [3c] Acaulescent, rhizomatous; rhizome 1.3–2.5 cm ∅; L 1–2 (–6), erect, linearlanceolate, 30–122  2.5–7 cm, narrowed gradually from  or above the middle to a channelled petiole, with a 3.2–3.8 mm green subulate tip, with alternating transverse bands of light dull green or clear whitish-green and deep grassgreen to almost blackish-green, with slight glaucous bloom, margin green, surface smooth; Inf 30–76 cm, simple, elongated, peduncle green with

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pale green dots, fertile portion lax, 3–8 Fl per cluster; Bra ovate or ovate-lanceolate, acuminate, 3.2–12.7 mm; Ped 5.3–8.5 mm; Fl white, tube 6.4–12.7 mm, lobes 14.8–19 mm. — Cytology: 2n = 36 (Sharma & Chaudhuri 1964). S. varians N. E. Brown (Bull. Misc. Inform. Kew 1915(5): 209, 1915). Type: K [ex cult.]. — Distr: Known from cultivation only, possibly from RSA (see note below).  Dracaena varians (N. E. Brown) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; L 4–8, distichous, erect or ascending, cylindrical with a groove on the upper surface or semicylindrical with a shallow groove on the upper surface, 38–114 cm, 1.3–2.2 cm thick, narrowed gradually to a 4.2–8.5 mm whitish hard acute tip, dull dark grass-green with numerous dark green longitudinal lines, when young with indistinct lighter green transverse bands, margin green or red-brown often edged white, surface slightly rough; Inf 60–76 cm, simple, elongated, peduncle light glaucous-green or greyish-green, fertile portion with 6–10 Fl per cluster; Bra linear or filiform, acute, 4.2–6.4 mm; Ped 4.2–5.3 mm; Fl white, speckled with purple at the tips, tube 10.6–12.7 mm, lobes 14.8–16.9 mm. South African plants described and illustrated by Jaarsveld (1987) and Jaarsveld (1994) as a form of S. pearsonii are regarded as this species by Jankalski (2007a). Not included in Lebrun & Stork (2014). S. volkensii Gürke (in Engler, Pfl.-welt Ost-Afr., Teil C, 144, 1895). Type [neo]: Tanzania, Pare Distr. (Wingfield 1525 [K]). — Distr: Kenya, Tanzania; edge of thickets on sandy soil, 500–900 m. I: Brown (1915: 212, as S. intermedia).  Dracaena volkensii (Gürke) Byng & Christenhusz (2018); incl. Sansevieria intermedia N. E. Brown (1914); incl. Acyntha polyrhitis Chiovenda (1932)  Sansevieria polyrhitis (Chiovenda) Chiovenda ex Guidotti (1932); incl. Sansevieria quarrei De Wildeman (1932); incl. Sansevieria humbertiana Guillaumin (1940)  Sansevieria gracilis var. humbertiana (Guillaumin) Mbugua (2007)  Dracaena serpenta var.

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humbertiana (Guillaumin) Byng & Christenhusz (2018). [3b] Acaulescent, rhizomatous; L 2–7, erect or ascending, semicylindrical, 45–122 cm, 1.3–1.9 cm thick, narrowed gradually to a whitish acute spinelike tip, dull deep green, becoming slightly bluishgreen, sometimes with faint whitish transverse bands, margin green or whitish, surface slightly rough; Inf 20–46 cm, simple, elongated, peduncle light greyish-green, fertile portion 30–40 cm, dense, 3–6 Fl per cluster; Bra ovate or ovatelanceolate, acute, 2.1–4.2 mm; Ped 2.1 mm; Fl white, tube 14.8–19 mm, pale greenish, lobes 12.7–19 mm, white or greenish-white with minute purplish spots on the outside. S. humbertiana, included here as a synonym, was recombined as Dracaena serpenta var. humbertiana by Christenhusz & al. (2018). S. zanzibarica Gérôme & Labroy (Bull. Mus. Hist. Nat. (Paris) 9: 170, 172–173, fig. 19, 1903). Type: Tanzania / Zanzibar? (Sacleux s.n. [P]). — Distr: Tanzania or Zanzibar. Incl. Sansevieria ehrenbergii Gérôme & Labroy (1903) (nom. illeg., Art. 53.1); incl. Sansevieria ehrenbergii De Wildeman (1905) (nom. illeg., Art. 53.1). [–] Very shortly caulescent; L distichous, recurved-spreading, linear-lanceolate, 15–30  1.9–2.5 cm, dull dark green with bluish-grey bloom, margin reddish-brown; Inf and Fl not known. Originally described with flowers unknown, and still a little-known species. Jankalski (2007a) suggests that it is possibly S. perrotii, and he was followed by Lebrun & Stork (2014). Included in S. arborescens by Mansfeld (2015a). S. zeylanica (Linné) Willdenow (Spec. Pl. 2: 159, 1799). Type: [lecto — icono]: Commelin, Hort. Med. Amstelod. Pl. Rar. 2: 41, t. 21, 1701. — Lit: Mansfeld & Ott (2015). Distr: Sri Lanka, India (Tamil Nadu); rocky or sandy places in dry regions. I: Wijnands (1973: 110).  Aloe hyacinthoides var. zeylanica Linné (1753)  Aletris hyacinthoides var. zeylanica (Linné) Linné (1762)  Aloe zeylanica (Linné) Jacquin (1762)  Aletris zeylanica (Linné) Miller

L. E. Newton

(1768)  Dracaena zeylanica (Linné) Mabberley (2017); incl. Aletris zeylanica Lamarck (1789) (nom. illeg., Art. 53.1); incl. Sansevieria indica Herter (1956) (nom. illeg., Art. 52.1). [3b] Acaulescent, rhizomatous; rhizome 1.3–1.5 cm ∅; L 5–11, erect below, slightly recurved above, linear to semicylindrical with a groove on the upper surface, 45–76  0.9–2.1 cm, 5–8.5 mm thick, narrowed gradually to a 12.7–38 mm long green soft subulate acute tip, dark green with lighter green transverse bands, with 4–7 darker green longitudinal lines, margin green, surface almost smooth; Inf 55–65 cm, simple, elongated, lax, up to 6 Fl per cluster; Fl white, grey-tipped, tube 20–25 mm, lobes to 35 mm. — Cytology: 2n = 40, 42 (Darlington & Wylie 1955); 2n = 42 (Janaki-Ammal 1945). Described with flowers not seen, and still a little-known species. Binojkumar (2002) claims that this species also occurs in India. Mansfeld & Ott (2015) give the leaf size as 30–40  1.5–2 cm.

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 or ders and families of flowering plants: APG IV. Bot. J. Linn. Soc. 181(1): 1–20. https://doi.org/10.1111/boj.12385 Binojkumar, M. S. (2002) A study on the genus Sansevieria Thunb. (Dracaenaceae) in India. J. Econ. Taxon. Bot. 26(2): 455–463, ills., key. Bos, J. J. (1984) Dracaena in West Africa. Agric. Univ. Wageningen Pap. 84–1– 126 pp., ills., maps, key. http:// edepot.wur.nl/283694 Bos, J. J. (1998) Dracaenaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 238–241, ills., key. Berlin (DE) etc.: Springer Verlag. https://doi.org/ 10.1007/978-3-662-03533-7_30 Brown, N. E. (1915) Sansevieria. A monograph of all known species. Bull. Misc. Inform. Kew 1915(5): 185–261, ills., key, 2 tt. http://biodiversitylibrary.org/ page/11492707 Brown, S. H. (2016) Winter-Frühlingsblüher in Süd-Florida. Sansevieria Online 4(1): 28–32, ills. Budweg, H.-G. (2014) Die rätselhaften Früchte der Sansevierien. Sansevieria Online 2(1): 4–31, ills.

Sansevieria RUSCACEAE Budweg, H.-G. (2016) Die Blütenstände der Sansevierien. Sansevieria Online 4(2): 4–24, ills. Budweg, H.-G. & Mansfeld, P. A. (2014) Sansevieria roxburghiana von der Koromandelküste. Sansevieria Online 2(1): 45–51, ills. Butler, A. (2009) Is this the real Sanseveria braunii Engler & Krause? Sansevieria 19: 11–12, ills. Butler, A. (2012a) Sansevieria powellii, species or hybrid? Dr Horst Pfennig’s experiments. Sansevieria 26: 7–18, ills. Butler, A. (2012b) Sansevieria itumea (Mbugua) Jankal. history and further thoughts. Sansevieria 28: 19–20, ills. Carlquist, S. & Schneider, E. L. (2007) Origins and nature of vessels in monocotyledons. 9. Sansevieria. South Afr. J. Bot. 73(2): 196–203, ills. https://doi.org/ 10.1016/j.sajb.2006.11.002 Chahinian, B. J. (1987) The Sansevieria trifasciata varieties. A presentation of all cultivated varieties. Reseda (US): Trans Terra Publishing. Chahinian, B. J. (1992) Editors’ notes. Sansevieria J. 1(3): 33, 45, ill. Chahinian, B. J. (2005a) The splendid Sansevieria. Buenos Aires (AR): Published by the author. Chahinian, B. J. (2005b) The many faces of Sansevieria robusta N. E. Brown. Sansevieria 11: 3–6, ills. Chahinian, B. J. (2012) The Sansevieria trifasciata varieties — a new look — [part I] / part II. Sansevieria 27: 2–6, 28: 2–12, ills. Chahinian, B. J. (2013) The Sansevieria trifasciata varieties — a new look — part III / part IV. Sansevieria 29: 2–13, 30: 2–9, ills. Chahinian, B. J. (2014) The Sansevieria trifasciata varieties — a new look — part V / VI. Sansevieria 31: 2–11, 32: 2–10, ills. Christenhusz, M. J. M. [& al. 2018], Fay, M. F. & Byng, J. W. (eds.) (2018) The Global Flora. A practical flora to vascular plant species of the world. Special Edition, GLOVAP Nomenclature Part 1, Vol. 4. Bradford (GB): Plant Gateway. Croix, I. la (2010) Dracaenaceae. In: Timberlake, J. R. & Martins, E. S. (eds.): Flora Zambesiaca, vol. 13, part 2, pp. 13–35, ills., key. Richmond (GB): Royal Botanic Gardens, Kew, for the Flora Zambesiaca Managing Committee. Darlington, C. D. & Wylie, A. P. (1955) Chromosome atlas of flowering plants. London (GB): Allen & Unwin. Friis, I. (1995) Sansevieria forskaoliana, a new name for Sansevieria abyssinica. Sansevieria J. 4(1): 3–8, ills. Govaerts, R. H. A. (comp.) (2014+) World checklist of selected plant families [continuously updated]. Richmond (GB): Trustees of the Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp Hepper, F. N. & Friis, I. (1994) The plants of Pehr Forsskål’s Flora Aegyptiaco-Arabica. Richmond (GB) etc.: Royal Botanic Gardens Kew in association with the Botanical Museum Copenhagen. Jaarsveld, E. J. van (1987) The succulent riches of South Africa and Namibia and their adaptive strategies. Aloe 24(3–4): 45–92, ills. Jaarsveld, E. J. van (1994) The Sansevieria species of South Africa and Namibia. Aloe 31(1): 11–15, ills., key.

1383 Jaarsveld, E. J. van (2016) Sansevieria hallii in the Gonarezhou National Park (Eastern Zimbabwe). Sansevieria 35: 8–14, ills. Jaarsveld, E. J. van & Condy, G. (2015) Sansevieria hallii. Flow. Pl. Afr. 64: 42–46, ills., map. Janaki-Ammal, E. K. (1945) In: Darlington, C. D. & Janaki-Ammal, E. K.: Chromosome atlas of cultivated plants. London (GB): Allen & Unwin. Jankalski, S. (2004) Sansevieria erythraeae. Sansevieria 9: 18–21, ill. Jankalski, S. (2007a) Brown’s Sansevieria monograph — an update. Sansevieria 15: 21–29. Jankalski, S. (2007b) Notes on the Sansevieria suffruticosa group. Sansevieria 17: 7–13, ills., key. Jankalski, S. (2007c) Sansevieria species described since Brown’s monograph. Sansevieria 17: 19–28. Jankalski, S. (2009a) The Sanseveria inflorescence and new sections proposed. Sansevieria 19: 8–10, ills. Jankalski, S. (2009b) Sansevieria hybrids. Sansevieria 20: 15–21, ills. Jankalski, S. (2009c) The typification of Sansevieria arborescens. Sansevieria 20: 22–26, ills. Jankalski, S. (2015) Infrageneric species groups in Sansevieria. Sansevieria 33: 15–19, ills., key. Jankalski, S. (2016) Sansevieria ballyi and its cultivars. Sansevieria 35: 18–21, ills. Joyner, J. F. [& al. 1951], Pate, J. B., Gangstad, E. O. & Seale, C. C. (1951) The potentialities of Sansevieria for fiber production in South Florida. Proc. Soil Sci. Soc. Florida 11: 138–139. Jumelle, H. (1923) Le Sansevieria canaliculata et le Sansevieria stuckyi. Bull. Mus. Nation. Hist. Nat. 29(8): 607–612. Khalumba, M. L. [& al. 2008], Mbugua, P. K. & Kungu, J. B. (2008) Ethnobotanical studies on four species of the genus Sansevieria Thnb. in Kenya. East Afr. J. Bot. 1(2): 161–164. Koller, A. L. & Rost, T. L. (1988a) Leaf anatomy in Sansevieria (Agavaceae). Amer. J. Bot. 75(5): 615–633, ills. http://www.jstor.org/stable/2444196 Koller, A. L. & Rost, T. L. (1988b) Structural analysis of water-storage tissue in leaves of Sansevieria (Agavaceae). Bot. Gaz. 149(3): 260–274, ills. http:// www.jstor.org/stable/2995258 Lavranos, J. J. (1994) The discovery of Sansevieria sp. “Eyl”. Sansevieria J. 3(3): 51–52. Lebrun, J.-P. & Stork, A. L. (2014) Tropical African flowering plants. Ecology and distribution. Volume 8: Monocotyledons 2 (Anthericaceae – Palmae). Genève (CH): Ville de Genève, Éditions des Conservatoire et Jardin Botaniques. Lu, P.-L. & Morden, C. W. (2014) Phylogenetic relationships among Dracaenoid genera (Asparagaceae: Nolinoideae) inferred from chloroplast DNA loci. Syst. Bot. 39(1): 90–104. https://doi.org/10.1600/ 036364414X678035 Mabberley, D. J. (2017) Mabberley’s Plant-Book. A portable dictionary of plants, their classification and uses. Ed. 4. Cambridge (GB): Cambridge University Press.

1384 Mai, E. (2016) Die Dominikanische Republik auch für Sansevierienfreunde. Sansevieria Online 4(1): 4–11, ills. Mangani, R. (1996) Sansevieria rorida (Lanza) N. E. Brown. Sansevieria J. 5(1): 11–17, ills. Mangani, R. (2014) The history of Sansevieria trifasciata Prain. Sansevieria 32: 11–19, ills. Mansfeld, P. A. (2013a) Die Gattung Sansevieria. Alle Arten und ihre Pflege. Hamburg (DE): Published by the author / Books on Demand GmbH. Mansfeld, P. A. (2013b) Neugliederung der Gattung Sansevieria (Asparagaceae). Kakt. and. Sukk. 64(2): 35–38, ills., key. Mansfeld, P. A. (2014a) Madagascar und seine Sansevierien. Sansevieria Online 2(1): 32–37, ills. Mansfeld, P. A. (2014b) Sansevieria forskaoliana — ein neuer Name für Sansevieria dawei. Sansevieria Online 2(2): 4–13, ills., maps. Mansfeld, P. A. (2014c) Die Feldnummern der Sansevierien. Sansevieria Online 2014(Sonderheft): 1–23. Mansfeld, P. A. (2015a) Die Systematik der Gattung Sansevieria (Asparagaceae) — ein aktueller Stand. Sansevieria Online 3(1): 20–29. Mansfeld, P. A. (2015b) Die unverwechselbare Sansevieria bacularis. Sansevieria Online 3(2): 35–39, ills., map. Mansfeld, P. A. (2016a) Sansevieria ballyi L. E. Newton (Asparagaceae) and newly found field notes. Bradleya 34: 225–229, map, ills. Mansfeld, P. A. (2016b) Blütenbildung bei Sansevieria burmanica. Sansevieria Online 4(2): 45–51, ills. Mansfeld, P. A. & Budweg, H.-G. (2015) Erstbeschreibung von: Sansevieria ballyi  Sansevieria parva hybr. nov. Sansevieria Online 3(1): 12–19, ills. Mansfeld, P. A. & Ott, G. H. F. (2015) The history of Sansevieria burmanica N. E. Br. and its rediscovery. Bradleya 33: 105–109, map, key, ills. Mansfeld, P. A. & Rosigkeit, J. (2016) Pagodenhaft und blühfreudig: Sansevieria francisii. Sansevieria Online 4(1): 33–39, ills., map. Mbugua, P. K. (1994) Three Sansevieria species of Kenya. 1. Sansevieria robusta N. E. Br. Cact. Succ. J. (US) 66 (2): 87–88, ills., map. Mbugua, P. K. (1998) Three Sansevieria species of Kenya. II. Sansevieria braunii. Cact. Succ. J. (US) 70(6): 311–312, ills. Mbugua, P. K. (2003) Sansevieria dawei Stapf: Its description and distribution in Africa. Sansevieria 7: 5–6, ills. Mbugua, P. K. (2007) Sansevieria. In: Beentje, H. J. & Ghazanfar, S. A. (eds.): Flora of Tropical East Africa: Dracaenaceae; pp. 10–41. Richmond (GB): Royal Botanical Gardens, Kew. Menale, B. [& al. 2013], Luca, P. de & Guacchio, E. del (2013) A plea to restore Petagna’s authorship for the genus Sansevieria, nom. cons. (Liliaceae). Taxon 62 (2): 387–390, ill. https://doi.org/10.12705/622.7 Newton, L. E. (1994) Observations on flowering of Sansevieria robusta in Kenya. EANHS Bull. 24(1): 8–11. Newton, L. E. (2003a) Sansevieria dooneri and S. parva. Sansevieria 7: 10–11, ill.

L. E. Newton Newton, L. E. (2003b) Additional cases of Sansevieria being chewed by animals. Sansevieria 8: 3. Newton, L. E. (2004) Sunbirds feeding on Sansevieria flowers. Nat. East Africa 34(2): 8, ill. Newton, L. E. (2005a) Horst Pfennig (1933–1994). Sansevieria 11: 7–8, ill. Newton, L. E. (2005b) What is Sansevieria patens? Sansevieria 13: 3. Newton, L. E. (2007) The inflorescence in the genus Sansevieria. Sansevieria 15: 15–16, ill. Newton, L. E. (2009a) The identity of Sansevieria arborescens (Ruscaceae), with an amplified description, and description of a new species. Bradleya 27: 153–158, ills. Newton, L. E. (2009b) Elephants like Sansevierias. Sansevieria 20: 8–10, ills. Newton, L. E. (2010) Notes on Sansevieria ballyi, with description of a new cultivar. Sansevieria 22: 10–12, ills., map. Newton, L. E. (2011) Thoughts on the Sansevieria inflorescence. Sansevieria 24: 7–10, ills. Newton, L. E. (2012a) Comments on Sansevieria powellii. Sansevieria 26: 6. Newton, L. E. (2012b) The identity of Sansevieria bagamoyensis. Bradleya 30: 103–106, ills. Newton, L. E. (2014) Sansevieria robusta and Sansevieria perrotii. Sansevieria 32: 24–25, ills. Newton, L. E. (2016) Species versus hybrid Sansevierias. Sansevieria 34: 2–4, ills. Newton, L. E. & Mbugua, P. K. (1993) Sansevieria leaves: A thirst quencher for baboons. Sansevieria J. 2(1): 5–6. Newton, L. E. & Thiede, J. (2015) Rank adjustments for the infraspecific taxa of Sansevieria pinguicula P. R. O. Bally (Asparagaceae / Dracaenaceae). Cact. Succ. J. (US) 87(1): 28–32, ills. https://doi.org/10.2985/ 015.087.0105 Newton, L. E. & Webb, R. H. (2011) The Tanzanian “Sansevieria grandicuspis” unmasked. J. East Afr. Nat. Hist. 100(1–2): 141–144, ills. https://doi.org/ 10.2982/028.100.0110 Nyffeler, R. & Eggli, U. (2010) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. https://doi.org/10.5167/uzh-35243 Obermeyer, A. A. [& al. 1992], Immelmann, K. L. & Bos, J. J. (1992) Dracaenaceae, Asparagaceae, Luzuriagaceae and Smilacaceae. In: Leistner, O. A. (ed.): Flora of Southern Africa, Vol. 5, part 3. Pretoria (ZA): National Botanic Institute. Perrier, H. (1938) 40e famille. Liliacées (Liliaceae). In: Humbert, H. (ed.): Flore de Madagascar. Tananarive (MG): Imprimerie Officielle. http://biodiversitylibrary. org/page/12247520 Pfennig, H. (1977a) Rasenbildend bis baumartig: Die Sansevierien. Gartenpraxis 1977: 506–511, ills. Pfennig, H. (1977b) Die Kultur der Sansevierien. Gartenpraxis 1977: 550–553, ills. Pfennig, H. (1981) Zur Systematik und Kultur einiger ostafrikanischer Sansevieria-Arten (Agavaceae). Bot. Jahrb. Syst. 102(1–4): 169–179, map.

Sansevieria RUSCACEAE Powys, J. G. L. (1996) Island fairies of the night. Sansevieria J. 5(1): 3–6. Rauh, W. (1963) Über einige interessante Sukkulenten aus Kenia. Sukkulentenkunde 7/8: 108–127, ills. Roy, M. (1956) A cytological investigation of the different species of Sansevieria with the aid of the improved technique. Caryologia 8: 221–230. https://doi.org/ 10.1080/00087114.1956.10797561 Rulkens, A. J. H. & Baptista, O. J. (2009a) Field observations and local uses of the poorly known Sansevieria pedicellata from Manica Province in Mozambique. Sansevieria 20: 2–7, ills. Rulkens, A. J. H. & Baptista, O. J. (2009b) Field impressions and ethno-veterinary plant use of Sansevieria hallii and related species in central Mozambique. Sansevieria 21: 8–14, ills. Rulkens, A. J. H. & Baptista, O. J. (2013) Notes on the distribution of Sansevieria burdettii Chahinian. Sansevieria 29: 14–16, ills. Rzepecky, A. (2013) Sansevieria sambiranensis H. Perr. Sansevieria 29: 18–23, ills. Sharma, A. K. & Chaudhuri, M. (1964) Cytological studies as an aid in assessing the status of Sansevieria, Ophiopogon and Curculigo. Nucleus (Calcutta) 7(1): 43–58. Speirs, D. C. (1984) Sansevieria species FKH 432. Brit. Cact. Succ. J. 2(3): 81–82, ills. Symoens, B. (2014) The origin of the name Sansevieria revealed? Sansevieria 31: 12–17, ills. Takawira-Nyenya, R. (2006) A taxonomic study of the genus Sansevieria (Dracaenaceae) in Zimbabwe. In: Ghazanfar, S. A. & Beentje, H. J. (eds.): Taxonomy and ecology of African plants, their conservation and sustainable use. Proc. 17th. AETFAT Congress, Addis Ababa, Ethiopia; pp. 61–71. Richmond (GB): Royal Botanical Gardens Kew. Takawira-Nyenya, R. & Stedje, B. (2011) Ethnobotanical studies in the genus Sansevieria Thunb. (Asparagaceae) in Zimbabwe. Ethnobot. Res. Applic. 9: 421–443, ills., map. https://doi.org/10.17348/era.9.0.421-443 Takawira-Nyenya, R. [& al. 2014], Newton, L. E., Wabuyele, E. & Stedje, B. (2014) Ethnobotanical studies in the genus Sansevieria Thunb. (Asparagaceae) in Coast Province of Kenya. Ethnobot. Res. Applic. 12:

1385 51–69, ills. http://www.ethnobotanyjournal.org/vol12/ i1547-3465-12-051.pdf Teketay, D. (1995) The genus Sansevieria Thunb. in Ethiopia. A contribution to the flora of Ethiopia. Sansevieria J. 4(2): 43–58, ills., map, key. Thiede, J. (1993) Notes on the Sansevieria species of Malawi. Sansevieria J. 2(2): 27–34, (3): 51–52, ills., map, key. Thiede, J. & Campbell-Barker, T. P. (2015) Sansevieria burdettii (Asparagaceae / Dracaenaceae): Its history, introduction and distribution, with new localities. Bradleya 33: 27–33, map, ills. Thiede, J. [& al. 2009], Hargreaves, B. J. & Downs, P. E. (2009) A gorgeous gem. Sansevieria downsii at Njakwa Gorge, Malawi. Cact. Succ. J. (US) 81(5): 224–229, 255, ills., map. https://doi.org/10.2985/ 015.081.0501 Thiede, J. [& al. 2017], Campbell-Barker, T. P. & Hargreaves, B. J. (2017) New records for the flora of Mount Mulanje, Malawi: Sansevieria sinus-simiorum Chahin. (Asparagaceae / Dracaenaceae), Crassula swaziensis Schönland and Crassula setulosa Harvey (Crassulaceae). Bradleya 35: 129–138, ills., map. https://doi.org/10.25223/brad.n35.2017.a12 Thulin, M. (ed.) (1995) Flora of Somalia. Volume 4. Angiospermae (Hydrocharitaceae – Pandanaceae). Richmond (GB): Royal Botanic Gardens Kew. Vrskovy, B. P. (2006) Sansevierie z Cerveného ostrova a hyacintovky. Cactaceae etc. 16(4): 132–140, ills. Vrskovy, B. P. (2009) A brief account of the Sansevierias of Yemen. Sansevieria 20: 11–14, ills. Waidhofer, A. (1996) Sansevieria deserti (Agavaceae). Cact. Succ. J. (US) 68(1): 9–11, ills. Webb, R. H. & Newton, L. E. (2016) Sansevieria laevifolia R. H. Webb & L. E. Newton: A new species from the Rift Valley of Kenya, with a discussion of the Sansevieria suffruticosa complex. Sansevieria 34: 9–13, ills., map. Wijnands, D. O. (1973) Typification and nomenclature of two species of Sansevieria (Agavaceae). Taxon 22(1): 109–114. https://doi.org/10.2307/1218041 Wilson, F. D. (1962) Florida H-13, a promising Sansevieria hybrid as a source of cordage fiber. Gainesville (US): Everglades Experimental Station.

Part XVIII The Family Xanthorrhoeaceae

Xanthorrhoeaceae U. Eggli

Perennial woody pachycaul small trees to stemless herbs with massive underground Rstock, branched or unbranched, stems with secondary thickening growth; L numerous in dense Ros at the stem tips, xeromorphic, narrowly oblong to filiform-linear, often wiry, tough, tips often spiny, basally often broadened and resiniferous, persistent when old; Inf spike-like, massively cylindrical on a woody leafless scape, dense-flowered; Fl bisexual, sessile, actinomorphic, 3-merous, in tight clusters surrounded by Bra; Per with 6 dry and chaffy persistent Tep in 2 series (often called sepals and petals), Tep free, OTep chartaceous, ITep white or yellow; St 3 + 3; Anth dorsifixed, introrse; Ov superior, of 3 united Ca, with septal Nec, 3-locular with 2 rows of ovules; Sty with undivided Sti; Fr loculicidal capsules with 1–2 Se per locule; Se with black testa and copious endosperm. Order: Asparagales Important Literature: Bedford & al. (1986), Clifford (1998).

In traditional classifications, Xanthorrhoeaceae embraced 10 genera with  100 species, but the family has for long been known to be heterogeneous (Clifford 1998). In its strict sense it is now restricted to the genus Xanthorrhoea, and the genera Baxteria, Calectasia, Dasypogon and Kingia are placed in the separate family Dasypogonaceae (including Baxteriaceae, Calectasiaceae and Kingiaceae). Dasypogonaceae were regarded as unplaced at the base of the Commelinids (APG 2009), and were moved to Arecales by APG (2016); they are restricted to W Australia. On the other hand, APG (2009), following suggestions already made earlier (APG 2003), opted for enlarging Xanthorrhoeaceae by including Asphodelaceae and Hemerocallidaceae (no succulents) as synonyms. Such an amount of lumping of well-established families does not result in a phylogenetically more correct classification (Nyffeler and Eggli 2010), and we and Smith & Figueiredo (this volume for Asphodelaceae) prefer to maintain the Asphodelaceae as separate family, and accept the Xanthorrhoeaceae in their narrow circumscription.

Distribution: Tropical and subtropical Australia.

U. Eggli (*) Sukkulenten-Sammlung Zürich, Grün Stadt Zürich, Zürich, Switzerland e-mail: [email protected] © Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8_30

1389

1390

Species of Xanthorrhoea (28–30 species in Australia) are sometimes included in succulent plant collections but are more properly regarded as mere xerophytes with woody stems. There are no indications that any form of water storage exists.

References APG (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436. https://doi.org/10.1046/j.1095-8339.2003. t01-1-00158.x 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):

U. Eggli 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 Bedford, D. J. [& al. 1986], Lee, A. T., Macfarlane, T. D., Henderson, R. J. F. & George, A. S. (1986) Xanthorrhoeaceae. In: George, A. S. (ed.): Flora of Australia, 46: 88-171, ills., keys. Canberra (AU): Australian Government Publishing Service. Clifford, H. T. (1998) Xanthorrhoeaceae. In: Kubitzki, K. (ed.): The families and genera of vascular plants; 3: 467-470, ills. Berlin (DE) etc.: Springer Verlag. https://doi.org/10.1007/978-3-662-03533-7_29 Nyffeler, R. & Eggli, U. (2010) An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125-144. https://doi.org/10.5167/uzh35243

Index

A Abromeitiella, 879 Abromeitiella abstrusa, 880 Abromeitiella brevifolia, 880 Abromeitiella brevifolia ssp. chlorantha, 881 Abromeitiella chlorantha, 881 Abromeitiella lorentziana, 881 Abromeitiella lotteae, 884 Abromeitiella pulvinata, 881 Abromeitiella scapigera, 887 Acanthospora, 1107 Acanthostachys, 849 Acanthostachys exilis, 850 Acanthostachys pitcairnioides, 850 Acanthostachys strobilacea, 850 Acyntha, 1353 Acyntha abyssinica, 1364 Acyntha abyssinica var. sublaevigata, 1363 Acyntha elliptica, 1363 Acyntha guineensis, 1381 Acyntha lanuginosa, 1362 Acyntha massae, 1371 Acyntha polyrhitis, 1381 Acyntha powellii, 1375 Acyntha rorida, 1377 Acyntha roxburghiana, 1377 Adernia, 1231 Aechmaea, 851 Aechmea, 851, 1053, 1102 Aechmea ampullacea, 856 Aechmea ampullaceal var. longifolia, 856 Aechmea benrathii, 855 Aechmea cylindrata, 853 Aechmea cylindrata var. micrantha, 853 Aechmea distichantha, 853 Aechmea distichantha fa. albiflora, 853 Aechmea distichantha var. distichantha, 853 Aechmea distichantha var. glaziovii, 853–854 Aechmea distichantha var. schlumbergeri, 854 Aechmea distichantha var. vernicosa, 854 Aechmea excavata, 853 Aechmea fasciata var. purpurea, 1049 Aechmea glaziovii, 854 Aechmea gracilis, 854

Aechmea grandiceps, 854 Aechmea hookeri, 853 Aechmea hyacinthus, 853 Aechmea involucrata, 854 Aechmea involucrifera, 854 Aechmea legrelliana, 855 Aechmea microphylla, 853 Aechmea minor, 854 Aechmea myriophylla, 853 Aechmea nudicaulis, 854–855 Aechmea ortgiesii, 855 Aechmea pitcairnioides, 850 Aechmea platyphylla, 853 Aechmea polystachia, 853 Aechmea polystachia var. excavata, 853 Aechmea polystachia var. myriophylla, 853 Aechmea polystachya, 854 Aechmea polystachya var. longifolia, 853 Aechmea pulchella, 854 Aechmea recurvata, 855 Aechmea recurvata var. benrathii, 855, 1049 Aechmea recurvata var. ortgiesii, 855 Aechmea recurvata var. recurvata, 855–856 Aechmea regelii, 854 Aechmea rubra, 854 Aechmea rupestris, 855 Aechmea weberbaueri, 851–856 Agallostachys antiacantha, 998 Agallostachys variegata, 1036 Agapanthaceae, 425 Agavaceae, 9, 12 Agave, 11, 13, 15, 21 Agave abisaii, 40 Agave aboriginum, 51 Agave abortiva, 39 Agave abrupta, 47 Agave acicularis, 41 Agave acklinicola, 42 Agave acuispina, 78 Agave affinis, 242 Agave ajoensis, 42 Agave aktites, 42 Agave albescens, 43 Agave albicans, 172

© Springer-Verlag GmbH Germany, part of Springer Nature 2020 U. Eggli, R. Nyffeler (eds.), Illustrated Handbook of Succulent Plants: Monocotyledons, https://doi.org/10.1007/978-3-662-56486-8

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1392 Agave albicans var. ctenophora, 173 Agave albicans var. mediopicta, 173 Agave albicans var. variegata, 173 Agave alboaustralis, 43 Agave albomarginata, 44 Agave albopilosa, 44 Agave alibertii, 271 Agave aloides, 39 Agave aloina, 205 Agave altissima, 45 Agave amaniensis, 240 Agave americana, 39, 45, 54, 86, 146, 188, 241, 260 Agave americana [?] nairobensis, 46 Agave americana cv. subtilis, 46 Agave americana fa. virginica, 45 Agave americana ssp. americana, 45 Agave americana ssp. expansa, 47 Agave americana ssp. oaxacensis, 47 Agave americana ssp. protoamericana, 47 Agave americana var. americana, 45 Agave americana var. coccinea, 61 Agave americana var. expansa, 47 Agave americana var. franzosinii, 112 Agave americana var. latifolia, 197 Agave americana var. marginata, 45 Agave americana var. marginata alba, 45 Agave americana var. marginata aurea, 45 Agave americana var. marginata pallida, 45 Agave americana var. medio-picta, 45 Agave americana var. oaxacensis, 47 Agave americana var. picta, 45 Agave americana var. striata, 45 Agave americana var. subtilis, 46 Agave americana var. theometel, 45 Agave americana var. variegata, 45 Agave amica, 48 Agave amoena, 139 Agave amurensis, 279 Agave ananassoides, 100 Agave andreae, 48 Agave angustiarum, 49 Agave angustifolia, 50, 251, 343 Agave angustifolia fa. woodrowii, 53 Agave angustifolia ssp. deweyana, 51 Agave angustifolia ssp. letonae, 52 Agave angustifolia ssp. rubescens, 50 Agave angustifolia ssp. sargentii, 53 Agave angustifolia ssp. tequilana, 251 Agave angustifolia var. angustifolia, 50 Agave angustifolia var. deweyana, 51 Agave angustifolia var. letonae, 52 Agave angustifolia var. marginata, 52 Agave angustifolia var. nivea, 53 Agave angustifolia var. pes-mulae, 251 Agave angustifolia var. rubescens, 50 Agave angustifolia var. sargentii, 53 Agave angustifolia var. variegata, 53 Agave angustissima, 116 Agave angustissima var. ortgiesiana, 88

Index Agave anomala, 53 Agave antillarum, 54 Agave antillarum ssp. grammontensis, 54 Agave antillarum var. antillarum, 54 Agave antillarum var. grammontensis, 54 Agave apedicellata, 54 Agave applanata, 55 Agave applanata var. huachucensis, 199 Agave applanata var. major, 55 Agave applanata var. parryi, 196 Agave applanata var. spectabilis, 55 Agave applanata var. subnivea, 55 Agave aquariensis, 168 Agave arcedianoensis, 57 Agave argyrophylla, 337 Agave arizonica, 58 Agave armata, 137, 491 Agave artichaut, 134 Agave arubensis, 58 Agave aspera, 39, 332 Agave asperrima, 58 Agave asperrima ssp. asperrima, 59 Agave asperrima ssp. maderensis, 59 Agave asperrima ssp. potosiensis, 60 Agave asperrima ssp. zarcensis, 60 Agave atrovirens, 60, 224 Agave atrovirens ssp. mirabilis, 62 Agave atrovirens var. atrovirens, 60 Agave atrovirens var. cochlearis, 61 Agave atrovirens var. marginata, 61 Agave atrovirens var. mirabilis, 62 Agave atrovirens var. salmiana, 223 Agave atrovirens var. sigmatophylla, 61 Agave attenuata, 62 Agave attenuata [?] dentata, 63 Agave attenuata cv. Boutin Blue, 63 Agave attenuata cv. Nova, 63 Agave attenuata ssp. attenuata, 62 Agave attenuata ssp. dentata, 63 Agave attenuata var. brevifolia, 62 Agave attenuata var. coarctata, 62 Agave attenuata var. compacta, 62 Agave attenuata var. elliptica, 62 Agave attenuata var. glauca, 62 Agave attenuata var. latifolia, 62 Agave attenuata var. paucibracteata, 62 Agave attenuata var. serrulata, 63 Agave attenuata var. subdentata, 63, 106 Agave attenuata var. subdenudata, 39 Agave attenuata var. subundulata, 62 Agave aurea, 64 Agave aurea ssp. aurea, 64 Agave aurea ssp. promontorii, 65 Agave aurea var. capensis, 65 Agave auricantha, 212, 213 Agave australis, 1213 Agave avellanidens, 65 Agave azurea, 66 Agave bahamana, 67

Index Agave bakeri, 120, 147 Agave banlan, 39 Agave barbadensis, 146 Agave baxteri, 39 Agave beaucarnei, 148 Agave beaulueriana, 112 Agave bedinghausii, 67 Agave beguinii, 68 Agave bennetii, 39 Agave bergeri, 51 Agave bernhardii, 39 Agave bessereriana, 160 Agave bessereriana [?] amoena, 160 Agave bessereriana [?] candida, 160 Agave bessereriana [?] glauca, 160 Agave bessereriana [?] hystrix, 160 Agave bessereriana [?] longifolia glauca, 160 Agave bessereriana [?] longifolia viridis, 160 Agave bicolor, 68 Agave blissii, 68 Agave bogotensis, 93 Agave boldinghiana, 68 Agave bollii, 173 Agave bollii var. celsii, 172 Agave bonnetiana, 225 Agave bonnetii, 39 Agave boscii, 116 Agave botteri, 172 Agave bouchei, 172 Agave bourgaei, 139 Agave bovicornuta, 69 Agave braceana, 70 Agave brachystachys, 228 Agave brachystachys var. pubescens, 217 Agave brachystachys var. strictior, 228 Agave bracteosa, 71 Agave brauniana, 185 Agave breedlovei, 51 Agave brevipetala, 72 Agave breviscapa, 158 Agave brevispina, 72 Agave brittoniana, 72 Agave brittoniana ssp. brachypus, 72 Agave brittoniana ssp. brittoniana, 73 Agave brittoniana ssp. sancti-spirituensis, 73 Agave brittoniana var. brachypus, 72 Agave bromeliifolia, 50 Agave brunnea, 73 Agave bulbifera, 78 Agave bulbosa, 329 Agave bulbulifera, 74 Agave bulliana, 74 Agave bundrantii, 76 Agave cacozela, 76 Agave caeciliana, 59 Agave caerulescens, 157 Agave caespitosa, 205 Agave cajalbanensis, 77 Agave calderonii, 39

1393 Agave californica, 245 Agave calodonta, 77 Agave campaniflora, 64 Agave campanulata, 343 Agave canartiana, 61 Agave canartiana var. laevior, 61 Agave candelabrum, 78 Agave cantala, 78 Agave cantala ssp. acuispina, 78 Agave cantala var. acuispina, 78 Agave cantala var. cantala, 78 Agave cantula, 78 Agave capensis, 65 Agave caratas, 224 Agave carchariodonta, 279 Agave caribaea, 39, 146 Agave caribaeicola, 146 Agave carminis, 242 Agave caroli-schmidtii, 234 Agave cavanillesii, 79 Agave caymanensis, 79 Agave celsiana, 45, 172 Agave celsiana [?] albida, 172 Agave celsiana [?] nana, 173 Agave celsii, 172 Agave celsii var. albicans, 172 Agave celsii var. celsii, 172 Agave cernua var. serrulata, 63 Agave cerulata, 79 Agave cerulata ssp. cerulata, 80 Agave cerulata ssp. dentiens, 80 Agave cerulata ssp. nelsonii, 81 Agave cerulata ssp. subcerulata, 81 Agave cerulata var. nelsonii, 81 Agave chamelensis, 82 Agave chazaroi, 83 Agave chiapensis, 83 Agave chiapensis var. major, 83 Agave chihuahuana, 197 Agave chinensis, 224 Agave chisosensis, 121 Agave chloracantha, 39, 185 Agave chloracantha var. brauniana, 185 Agave chrysantha, 84 Agave chrysoglossa, 85 Agave cinerascens, 55 Agave coarctata, 225 Agave coccinea, 61, 188 Agave cochlearis, 224 Agave cocui, 85 Agave cocui var. cucutensis, 86 Agave cocui var. laguayrensis, 86 Agave coelum, 225, 226 Agave coetocapnia, 86 Agave coetocapnia ssp. clivicola, 87 Agave coetocapnia ssp. coetocapnia, 87 Agave coetocapnia ssp. pueblensis, 88 Agave cohniana, 243 Agave colimana, 88

1394 Agave colimillensis, 57 Agave collina, 89 Agave colorata, 89 Agave commelynii, 343 Agave communis, 45 Agave compacta, 62, 201 Agave complicata, 46 Agave compluviata, 224 Agave concinna, 39, 172, 173 Agave conduplicata, 271 Agave confertiflora, 90 Agave congesta, 91 Agave conjuncta, 167 Agave connochaetodon, 164 Agave considerantii, 268 Agave consociata, 99 Agave convallis, 92 Agave cookei, 39 Agave corderoyi, 147 Agave cordillerensis, 46 Agave cornuta, 279 Agave costaricana, 51 Agave costaricensis, 277 Agave couesii, 198 Agave crassispina, 224 Agave crassispina var. culta, 224 Agave crenata, 139, 167 Agave cubensis, 325, 332, 343 Agave cubensis var. striata, 341 Agave cucullata, 39 Agave cundinamarcensis, 92 Agave cupreata, 93 Agave cuspidata, 51 Agave cyanophylla, 158 Agave dasylirioides, 94 Agave dasylirioides var. dealbata, 94 Agave datylio, 95 Agave datylio ssp. vexans, 95 Agave datylio var. vexans, 95 Agave davillonii, 39 Agave dealbata, 94 Agave dealbata [?] angustifolia, 94 Agave dealbata [?] compacta, 94 Agave dealbata [?] compacta angustifolia, 94 Agave dealbata var. nana, 94 Agave deamiana, 39, 253 Agave debaryana, 63 Agave debilis, 96 Agave decaisneana, 39 Agave decipiens, 96 Agave deflexispina, 61 Agave delamateri, 97, 98 Agave demeesteriana, 100 Agave densiflora, 172, 185 Agave densiflora var. angustifolia, 185 Agave densiflora var. foliis striatis aureis, 185 Agave dentata, 63 Agave dentiens, 80 Agave deserti, 98, 99

Index Agave deserti ssp. deserti, 98, 100 Agave deserti ssp. pringlei, 216 Agave deserti ssp. simplex, 100 Agave deserti var. pringlei, 216 Agave deserti var. simplex, 100 Agave desmetiana, 100, 101, 134 Agave deweyana, 51 Agave diacantha, 39 Agave difformis, 101, 102 Agave disceptata, 154 Agave disjuncta, 233 Agave dissimulans, 92 Agave doctorensis, 102 Agave dolichantha, 103 Agave dominicensis, 54 Agave donnell-smithii, 51 Agave drimiifolia, 260 Agave duplicata, 86 Agave duplicata ssp. duplicata, 87 Agave duplicata ssp. graminifolia, 124 Agave duplicate ssp. pueblensis, 88 Agave duplicate var. clivicola, 87 Agave durangensis, 103, 104 Agave dussiana, 146 Agave dyckii, 224 Agave eborispina, 262 Agave eduardii, 270 Agave edwardii, 270 Agave eggersiana, 104, 105 Agave ehrenbergii, 172 Agave eichlamii, 191 Agave eichlamii var. interjecta, 191 Agave ekmanii, 257 Agave ekmanii var. microdonta, 257 Agave elegans, 212 Agave elizae, 39 Agave ellemeetiana, 105 Agave ellemeetiana ssp. ellemeetiana, 105, 106 Agave ellemeetiana ssp. subdentata, 106 Agave ellemeetiana var. subdentata, 106 Agave elliptica, 63 Agave elongata, 51, 111 Agave endlichiana, 51 Agave engelmannii, 106 Agave ensifera, 107 Agave ensiformis, 246 Agave entea, 39 Agave erosa, 39 Agave erubescens, 51 Agave evadens, 107, 108 Agave excelsa, 51 Agave expansa, 47 Agave expatriata, 148 Agave falcata, 245 Agave falcata var. espadina, 245 Agave falcata var. microcarpa, 245 Agave felgeri, 108, 109 Agave felina, 46 Agave fenzliana, 139

Index Agave ferdinand-cortez, 224 Agave ferdinandi-regis var. nigra, 226 Agave ferox, 225 Agave filamentosa, 109 Agave filifera, 109, 110 Agave filifera fa. ortgiesiana, 40 Agave filifera ssp. filifera, 109 Agave filifera ssp. microceps, 170 Agave filifera ssp. multifilifera, 177 Agave filifera ssp. schidigera, 230 Agave filifera var. adornata, 230 Agave filifera var. angustifolia, 230 Agave filifera var. candida superba, 109 Agave filifera var. compacta, 170 Agave filifera var. elatior, 109 Agave filifera var. filamentosa, 109 Agave filifera var. ignescens, 230 Agave filifera var. immaculata, 109 Agave filifera var. longifolia, 109 Agave filifera var. mediopicta, 109 Agave filifera var. pannosa, 230 Agave filifera var. schidigera, 230 Agave filifera var. splendens, 109 Agave filifera var. superba, 109 Agave filifera var. viridis, 109 Agave flaccida, 39, 50 Agave flaccifolia, 185 Agave flavescens, 160 Agave flavescens var. macracantha, 160 Agave flavescens var. macroacantha, 160 Agave flavovirens, 51 Agave flexispina, 110 Agave foetida, 329 Agave fortiflora, 110, 111 Agave fourcroydes, 39, 111, 112 Agave fourcroydes var. espiculata, 111 Agave fragrantissima, 39 Agave franceschiana, 240 Agave franzosinii, 39, 112, 113 Agave friderici, 39 Agave fuerstenbergii, 45 Agave funifera, 352 Agave funkiana, 113 Agave fusca, 113, 114 Agave galeottei, 39 Agave galvaniae, 114, 115 Agave garciae-mendozae, 115 Agave garciaruizii, 115, 116 Agave geminiflora, 116, 117 Agave geminiflora var. filamentosa, 116 Agave geminiflora var. filifera, 116 Agave geminiflora var. sonorae, 232 Agave geminiflora var. stricta-viridis, 116 Agave gentryi, 117, 118 Agave ghiesbreghtii, 118, 119 Agave ghiesbreghtii [?] dentata, 63 Agave ghiesbreghtii [?] mollis, 62 Agave ghiesbreghtii var. peacockii, 203 Agave gigantea, 329, 343

1395 Agave gigantensis, 119, 120 Agave gilbertii, 120 Agave gilbeyi, 118 Agave glabra, 39 Agave glaucescens, 39, 62 Agave glomeruliflora, 121 Agave goeppertiana, 172 Agave goldmaniana, 235 Agave gomezpompae, 121 Agave gonzaloi, 121, 122 Agave gracielae, 122 Agave gracilipes, 122, 123 Agave gracilis, 61, 123, 206, 207 Agave gracilispina, 45 Agave gracillima, 123, 124 Agave graminifolia, 124, 125 Agave grandibracteata, 39 Agave grandidentata, 134 Agave granulosa, 39 Agave grenadina, 146 Agave grijalvensis, 149 Agave grisea, 125 Agave grisea var. cienfuegosana, 125 Agave grisea var. grisea, 125 Agave grisea var. obesispina, 125 Agave guadalajarana, 125, 126 Agave guatemalensis, 234 Agave guccifolia, 243 Agave guedeneyrii, 39 Agave guemensis, 126 Agave guerrerensis, 126, 127 Agave guiengola, 127, 128 Agave guignardii, 128 Agave gustaviana, 167 Agave gutierreziana, 39 Agave guttata, 128, 129 Agave gypsicola, 129 Agave gypsophila, 129, 130 Agave hanburyi, 256 Agave harrisii, 130, 131 Agave hartmanii, 202, 203 Agave haseloffii, 172 Agave hauniensis, 131 Agave havardiana, 131, 132 Agave haworthiana, 158 Agave haynaldii, 101 Agave haynaldii var. utahensis, 261 Agave henriquesii, 39 Agave heteracantha, 39, 107, 157 Agave heteracantha var. glomeruliflora, 121 Agave heteracantha var. splendens, 279 Agave heteracantha var. univittata, 157 Agave heteracantha var. vittata, 157 Agave heterodon, 139 Agave hexapetala, 332 Agave hidalgensis, 115 Agave hiemiflora, 133 Agave hookeri, 133, 134, 139, 185, 243 Agave horizontalis, 39

1396 Agave horizontinalis, 39 Agave horrida, 134, 148 Agave horrida ssp. horrida, 134, 135 Agave horrida ssp. perotensis, 135 Agave horrida var. gilbeyi, 118 Agave horrida var. laevior, 134 Agave horrida var. latifrons, 134 Agave horrida var. macrodonta, 134 Agave horrida var. maigretiana, 134 Agave horrida var. micracantha, 134 Agave horrida var. monstruosa, 134 Agave horrida var. recurvispina, 134 Agave horrida var. triangularis, 256 Agave horrida var. viridis, 134 Agave houghii, 270 Agave houllettii, 51 Agave howardii, 136 Agave huachucensis, 199 Agave huehueteca, 118 Agave humboldtiana, 39 Agave humilis, 228 Agave hurteri, 136, 137 Agave hybrida, 137 Agave hystrix, 246, 247 Agave impressa, 137, 139 Agave inaequidens, 139 Agave inaequidens ssp. barrancensis, 139 Agave inaequidens ssp. inaequidens, 139, 140 Agave inaguensis, 140 Agave indagatorum, 140, 141 Agave ingens, 46 Agave ingens var. picta, 45 Agave inghamii, 39 Agave inghamii [?] longissima, 39 Agave inopinabilis, 148 Agave integrifolia, 160 Agave intermixta, 141 Agave intrepida, 94 Agave involuta, 141, 142 Agave isthmensis, 142, 143 Agave ixtli, 39, 50, 111 Agave ixtlioides, 51 Agave ixtli var. elongata, 111 Agave ixtli var. excelsa, 51 Agave jacobiana, 224 Agave jacquiniana, 50 Agave jaiboli, 143 Agave jaliscana, 143, 144 Agave jarucoensis, 144 Agave jimenoi, 144, 145 Agave justosierrana, 145 Agave kaibabensis, 261 Agave karatto, 145, 146 Agave karwinskiana, 147 Agave karwinskii, 146, 147 Agave katherineae, 167 Agave kavandivi, 147, 148 Agave kellermaniana, 39, 253 Agave kellockii, 63

Index Agave keratto, 146 Agave kerchovei, 148, 149 Agave kerchovei var. beaucarnei, 148 Agave kerchovei var. brevifolia, 148 Agave kerchovei var. diplacantha, 148 Agave kerchovei var. distans, 148 Agave kerchovei var. glauca, 148 Agave kerchovei var. inermis, 256 Agave kerchovei var. macrodonta, 148 Agave kerchovei var. miniata, 148 Agave kerchovei var. pectinata, 148 Agave kerchovei var. variegata, 148 Agave kerrato, 146 Agave kerratto, 146 Agave kewensis, 149, 150 Agave killischea, 39 Agave killischkii, 134 Agave kirchneriana, 51 Agave knightiana, 116 Agave kochii, 279 Agave kristenii, 151 Agave laetevirens, 241 Agave lagunae, 151, 152 Agave lamprochlora, 39 Agave langlassei, 228 Agave lata, 271 Agave laticincta, 39 Agave latifolia, 212 Agave latissima, 224 Agave laurentiana, 188 Agave laxa, 78, 147 Agave laxifolia, 96 Agave lechuguilla, 152, 154 Agave lechuguilla fa. glomeruliflora, 121 Agave lechuguilla var. caerulescens, 157 Agave legrelliana, 188 Agave legrelliana var. breviflora, 188 Agave leguayana, 39, 118 Agave lehmannii, 224 Agave lemairei, 39 Agave lempana, 39 Agave leopoldii, 154, 196 Agave lepida, 158 Agave lespinassei, 51 Agave letonae, 52 Agave letonae var. marginata, 52 Agave lindleyi, 39 Agave linearis, 160 Agave littaeoides, 39 Agave littoralis, 154, 155 Agave longibracteata, 155 Agave longiflora, 155, 156 Agave longifolia, 111 Agave longifolia var. picta, 45 Agave longipes, 156, 157 Agave longisepala, 39 Agave lophantha, 157, 158 Agave lophantha fa. caerulescens, 157 Agave lophantha var. angustifolia, 157

Index Agave lophantha var. brevifolia, 157 Agave lophantha var. caerulescens, 157 Agave lophantha var. gracilior, 157 Agave lophantha var. latifolia, 107 Agave lophantha var. pallida, 157 Agave lophantha var. poselgeri, 152 Agave lophantha var. subcanescens, 157 Agave lophantha var. tamaulipasana, 152 Agave lophantha var. univittata, 157 Agave lurida, 50, 158, 159, 206 Agave lurida var. jacquiniana, 50 Agave lyobaa, 159, 160 Agave macracantha, 160 Agave macrantha, 160, 173 Agave macrantha var. concinna, 173 Agave macroacantha, 160, 161 Agave macroacantha var. concinna, 172 Agave macroacantha var. integrifolia, 160 Agave macroacantha var. latifolia, 160 Agave macroacantha var. macrantha, 173 Agave macroacantha var. planifolia, 160 Agave macroculmis, 61, 117 Agave maculata, 39, 161 Agave maculata var. brevituba, 161 Agave maculosa, 161, 162 Agave maculosa var. brevituba, 161 Agave maculosa var. minor, 161 Agave madagascariensis, 329 Agave madrensis, 162 Agave magni, 158 Agave maigretiana, 134 Agave malinczii, 39 Agave manantlanicola, 163 Agave mangui, 158 Agave mapisaga, 163, 164 Agave mapisaga ssp. lisa, 163 Agave mapisaga var. lisa, 163 Agave marcusea, 198 Agave marcusii, 197 Agave marensii, 198 Agave margaritae, 164, 165 Agave maria-patriciae, 165 Agave maritima, 88 Agave marmorata, 165, 166 Agave martiana, 146 Agave massiliensis, 166 Agave maximiliana, 166, 280 Agave maximiliana var. bovicornuta, 69 Agave maximiliana var. katherineae, 167 Agave maximiliana var. maximiliana, 167, 168 Agave maximilianea, 224 Agave maximowicziana, 39 Agave mayoensis, 270 Agave mckelveyana, 168, 169 Agave medioxima, 146 Agave megalacantha, 139 Agave megalodonta, 169 Agave melanacantha, 188 Agave melliflua, 46

1397 Agave mescal, 139 Agave mexicana, 70, 158 Agave mexicana var. picta, 45 Agave mezortillo, 157 Agave michoacana, 103 Agave micracantha, 172, 185 Agave micracantha [?] variegata, 173 Agave micracantha var. albicans, 172 Agave micracantha var. albidior, 172 Agave micracantha var. mitis, 172 Agave micracantha var. picta, 172 Agave microceps, 169, 170 Agave milleri, 39, 45 Agave milleri var. picta, 45 Agave millspaughii, 170 Agave minarum, 39, 253 Agave minor, 170, 171 Agave mirabilis, 62 Agave miradorensis, 100 Agave miradorensis var. regeliana, 100 Agave missionum, 171, 172 Agave mitis, 172, 174 Agave mitis ssp. albidior, 172 Agave mitis var. albidior, 172 Agave mitis var. mitis, 172 Agave mitriformis, 225 Agave monostachya, 39 Agave monstruosa, 39 Agave montana, 174 Agave montezumae, 224 Agave montium-sancticaroli, 174, 175 Agave montserratensis, 146 Agave mooreana, 39 Agave moranii, 175 Agave morrisii, 171, 241, 260 Agave mortolensis, 175 Agave muelleriana, 40 Agave muilmannii, 185 Agave mulfordiana, 232, 233 Agave multicolor, 176 Agave multifilifera, 176, 177 Agave multiflora, 40 Agave multilineata, 152 Agave murpheyi, 177, 178 Agave myriacantha, 185 Agave nanchititlensis, 178 Agave nashii, 178, 179 Agave nayaritensis, 179 Agave neglecta, 275, 276 Agave nelsonii, 81 Agave neocernua, 179 Agave neokewensis, 180 Agave neomexicana, 197 Agave neonelsonii, 180 Agave neopringlei, 55 Agave nevadensis, 262 Agave nevidis, 146 Agave newberryi, 262 Agave nickelsiae, 181

1398 Agave nigra, 40, 226 Agave nigromarginata, 40 Agave nirvana, 40 Agave nissonii, 40 Agave nivea, 53 Agave nizandensis, 182, 183 Agave noackii, 205 Agave noacksii, 205 Agave noah, 190 Agave noli-tangere, 148 Agave nussaviorum, 183 Agave nuusaviorum, 183 Agave nuusaviorum ssp. deltoidea, 183 Agave nuusaviorum ssp. nuusaviorum, 183, 184 Agave oaxacana, 184 Agave obducta, 146 Agave oblongata, 172 Agave obscura, 184, 185 Agave ocahui, 186 Agave ocahui ssp. longifolia, 186 Agave ocahui var. longifolia, 186 Agave ocahui var. ocahui, 186, 187 Agave occidentalis, 187 Agave odorata, 332 Agave offoyana, 187 Agave oligophylla, 160 Agave oliveriana, 228 Agave opacidens, 191 Agave orcuttiana, 236 Agave ornata, 45 Agave ornithobroma, 188, 189 Agave oroensis, 189 Agave ortgiesiana, 40, 88 Agave ortgiesiana var. brevifolia, 88 Agave ottonis, 61 Agave ousselghemiana, 172 Agave ousselghemiana [?] albo-picta, 173 Agave ousselghemiana [?] picta, 173 Agave ovatifolia, 190 Agave owenii, 51 Agave pablocarrilloi, 190, 191 Agave pachyacantha, 236 Agave pachycentra, 191 Agave pacifica, 51 Agave pallida, 40, 271 Agave palmaris, 251 Agave palmeri, 192, 193 Agave palmeri ssp. chrysantha, 84 Agave palmeri var. chrysantha, 84 Agave palustris, 193 Agave pampaniniana, 40 Agave panamana, 51 Agave paniculata, 194 Agave papyrocarpa, 194 Agave papyrocarpa ssp. macrocarpa, 195 Agave papyrocarpa ssp. papyrocarpa, 195 Agave parayi, 198 Agave parrasana, 195 Agave parreyi, 198

Index Agave parryi, 196 Agave parryi fa. integrifolia, 198, 199 Agave parryi ssp. couesii, 198 Agave parryi ssp. huachucensis, 199 Agave parryi ssp. neomexicana, 197 Agave parryi ssp. parryi, 197 Agave parryi ssp. truncata, 200 Agave parryi var. couesii, 198 Agave parryi var. huachucensis, 199 Agave parryi var. neomexicana, 197 Agave parryi var. parryi, 199 Agave parryi var. truncata, 200 Agave parva, 200 Agave parvidentata, 201 Agave parviflora, 201, 202 Agave parviflora ssp. densiflora, 202 Agave parviflora ssp. flexiflora, 202 Agave parviflora ssp. parviflora, 202 Agave paryi, 198 Agave patonii, 197 Agave paucifolia, 160, 245 Agave paupera, 40 Agave pavoliniana, 40 Agave pax, 203 Agave payrii, 198 Agave peacockii, 203 Agave pedrosana, 251 Agave pedunculifera, 63 Agave pelona, 204, 205 Agave pendula, 205 Agave perbella, 137 Agave perlucida, 40 Agave perplexans, 230 Agave perringea, 40 Agave pes-mulae, 251 Agave petiolata, 206 Agave petrophila, 206 Agave petskinil, 207 Agave pfersdorffii, 208 Agave phillipsiana, 208 Agave picta, 45 Agave pintilla, 208, 209 Agave planera, 40 Agave planifolia, 209 Agave platyphylla, 210 Agave polianthes, 48 Agave polianthiflora, 211, 212 Agave polianthoides, 228 Agave polyacantha, 40, 107 Agave polyacantha var. densiflora, 185 Agave polyacantha var. xalapensis, 185 Agave polyantha, 107 Agave polyanthoides, 228 Agave polyphylla, 158 Agave portoricensis, 171 Agave poselgeri, 152 Agave poselgeri var. kerchovei, 148 Agave potatorum, 40, 212, 224 Agave potatorum var. minor, 212

Index Agave potatorum var. verschaffeltii, 212 Agave potosina, 213 Agave potrerana, 214, 215 Agave prainiana, 51 Agave pratensis, 215 Agave pringlei, 40, 216 Agave producta, 216 Agave prolifera, 51 Agave promontorii, 65 Agave prostrata, 40 Agave protuberans, 128 Agave pruinosa, 63 Agave pseudofilifera, 109 Agave pseudotequilana, 251, 252 Agave pubescens, 217 Agave pugioniformis, 160 Agave pulcherrima, 40, 205 Agave pulchra, 212 Agave pulverulenta, 40 Agave pumila, 40, 217, 218 Agave punctata, 50 Agave purpurea, 40 Agave purpusorum, 118 Agave pygmae, 234, 235 Agave pygmaea, 234 Agave quadrata, 212 Agave quilae, 218 Agave quiotepecensis, 219 Agave quiotifera, 224 Agave ragusae, 40, 224 Agave ramosa, 45 Agave rasconensis, 46 Agave recurva, 246 Agave regeliana, 100, 134 Agave regelii, 135 Agave regia, 40 Agave reginae, 139 Agave repanda, 97 Agave revoluta, 220 Agave rhodacantha, 220, 221 Agave richardsii, 246 Agave rigida, 50, 51, 67 Agave rigida var. elongata, 51, 111 Agave rigida var. longifolia, 111 Agave rigida var. sisalana, 240 Agave robusta, 40 Agave roezliana, 118 Agave roezliana var. gilbeyi, 118 Agave roezliana var. inghamii, 118 Agave roezliana var. peacockii, 203, 204 Agave roezlii, 118 Agave rohanii, 40, 118 Agave romanii, 221 Agave roseana, 242 Agave rosei, 221 Agave rosselloana, 222 Agave rovelliana, 222 Agave rubescens, 50 Agave rubrocincta, 205

1399 Agave rudis, 40 Agave rufocincta, 205 Agave rumphii, 78 Agave rupicola, 172 Agave rupicola var. bouchei, 172 Agave rupicola var. brevifolia, 172 Agave rupicola var. longifolia, 172 Agave rupicola var. rubridentata, 172 Agave rutteniae, 222, 223 Agave rzedowskiana, 223 Agave salm-dyckii, 146 Agave salmiana, 223 Agave salmiana ssp. crassispina, 224 Agave salmiana ssp. ferox, 225 Agave salmiana ssp. salmiana, 224 Agave salmiana ssp. tehuacanensis, 225 Agave salmiana var. angustifolia, 163 Agave salmiana var. cochlearis, 224 Agave salmiana var. contorta, 224, 225 Agave salmiana var. ferox, 225 Agave salmiana var. glauca, 224 Agave salmiana var. gracilispina, 45 Agave salmiana var. mitriformis, 225 Agave salmiana var. recurvata, 163 Agave saltilloensis, 226 Agave samalana, 136 Agave sanpedroensis, 226, 227 Agave santanamichelii, 227 Agave saponaria, 228 Agave saponifera, 40 Agave sartorii, 205 Agave sartorii var. caespitosa, 205, 206 Agave sartorii var. oblongata, 172 Agave saundersii, 212 Agave scaberrima, 216 Agave scabra, 59, 228 Agave scabra ssp. maderensis, 60 Agave scabra ssp. potosiensis, 60 Agave scabra ssp. zarcensis, 60 Agave scaphoidea, 262 Agave scaposa, 229 Agave scheuermaniana, 146 Agave schidigera, 229, 230 Agave schidigera ssp. huicholensis, 230 Agave schidigera var. angustifolia, 230 Agave schidigera var. ignescens, 230 Agave schidigera var. ortgiesiana, 88 Agave schidigera var. plumosa, 230 Agave schidigera var. taylorii, 230 Agave schlechtendahlii, 61 Agave schneideriana, 231 Agave schnittspahnii, 212 Agave schottii, 231 Agave schottii var. atricha, 231 Agave schottii var. schottii, 232 Agave schottii var. serrulata, 232 Agave schottii var. treleasei, 231 Agave scolymus, 40, 77, 212 Agave scolymus var. polymorpha, 212

1400 Agave scolymus var. seemanniana, 234 Agave sebastiana, 233 Agave seemanniana, 234 Agave seemanniana ssp. pygmaea, 234 Agave seemanniana var. perscabra, 234 Agave seemanniana var. pygmaea, 234 Agave seemannii, 234 Agave seemannii var. acuta, 234 Agave seemannii var. nana, 234 Agave seemannii var. papillosa, 234 Agave seemannii var. parvispina, 234 Agave seemannii var. rotundifolia, 234 Agave segurae, 240, 241 Agave serrata, 40 Agave serrulata, 40, 50 Agave shaferi, 235 Agave shawii, 81, 235 Agave shawii ssp. goldmaniana, 235 Agave shawii ssp. shawii, 236 Agave shawii var. goldmaniana, 235 Agave shawii var. sebastiana, 233 Agave shrevei, 237 Agave shrevei ssp. magna, 237 Agave shrevei ssp. matapensis, 237 Agave shrevei ssp. shrevei, 237 Agave sicifolia, 51 Agave sileri, 238 Agave silvestris, 40 Agave simoni, 239 Agave simonii, 137 Agave simonis, 218 Agave singuliflora, 239 Agave sisalana, 240 Agave sisalana fa. armata, 240 Agave sisalana fa. marginata, 240 Agave sisalana fa. medio-picta, 240 Agave sisalana var. armata, 240 Agave slevinii, 242, 243 Agave smithiana, 40 Agave sobolifera, 67, 241 Agave sobolifera [?] marginata, 241 Agave sobolifera fa. spinidentata, 241 Agave sobolifera var. serrulata, 51 Agave sobria, 241 Agave sobria ssp. frailensis, 241, 242 Agave sobria ssp. roseana, 242 Agave sobria ssp. sobria, 242 Agave sobria var. roseana, 242 Agave sordida, 40 Agave spatularia, 55 Agave spectabilis, 45, 51, 62, 63 Agave spicata, 228, 243 Agave spinosa, 343 Agave spiralis, 96 Agave splendens, 279 Agave stenophylla, 325 Agave stictata, 244 Agave striata, 244, 246 Agave striata fa. recurva, 246

Index Agave striata fa. stricta, 247 Agave striata ssp. falcata, 244 Agave striata ssp. striata, 245 Agave striata ssp. stricta, 247 Agave striata var. californica, 245 Agave striata var. echinoides, 247 Agave striata var. mesae, 246 Agave striata var. nana, 246 Agave striata var. recurva, 246 Agave striata var. stricta, 247 Agave stricta, 246, 247 Agave stringens, 248 Agave subfalcata, 160 Agave subinermis, 40 Agave subsimplex, 248, 249 Agave subtilis, 46 Agave subzonata, 46 Agave sudburyensis, 160 Agave sullivanii, 111 Agave taeniata, 40 Agave taylorea, 249 Agave taylorii, 230, 249 Agave tecta, 249, 250 Agave tehuacanensis, 225 Agave telegraphica, 250 Agave temacapulinensis, 250 Agave tenuifolia, 251 Agave tenuispina, 191 Agave teopiscana, 83 Agave teoxomuliana, 40 Agave tequilana, 251 Agave terraccianoi, 222 Agave theometel, 45 Agave theoxmuliana, 50 Agave thomasiae, 252, 253 Agave thomsoniana, 40 Agave tigrina, 271 Agave tingens, 46 Agave titanota, 253 Agave titanota [?] minor, 253, 254 Agave todaroi, 166 Agave toeniata, 40 Agave toneliana, 40 Agave toumeyana, 254 Agave toumeyana ssp. bella, 254 Agave toumeyana ssp. toumeyana, 255, 256 Agave toumeyana var. bella, 255 Agave toumeyana var. toumeyana, 255, 256 Agave trankeera, 146 Agave treleasei, 231 Agave triangularis, 256 Agave triangularis var. rigidissima, 256 Agave triangularis var. subintegra, 256, 257 Agave troubetskoyana, 40 Agave tuberosa, 48, 342 Agave tuberosa var. β spinis duplicibus, 330 Agave tuberosa var. β spinis solitariis, 343 Agave tubulata, 257 Agave tubulata ssp. brevituba, 257

Index Agave turneri, 258, 259 Agave umbrophila, 259 Agave uncinata, 185 Agave underwoodii, 259 Agave undulata, 260 Agave undulata [?] glauca, 260, 261 Agave undulata var. strictior, 260 Agave unguiculata, 146 Agave univittata, 157 Agave univittata var. angustifolia, 157 Agave univittata var. brevifolia, 157 Agave univittata var. caerulescens, 157 Agave univittata var. carchariodonta, 279 Agave univittata var. ensifera, 107 Agave univittata var. foliis striatis, 152 Agave univittata var. gracilior, 157 Agave univittata var. heteracantha, 157 Agave univittata var. latifolia, 107 Agave univittata var. lophantha, 157 Agave univittata var. recurvispinis, 152 Agave univittata var. subcanescens, 157 Agave univittata var. tamaulipasana, 152 Agave univittata var. viridis, 152 Agave univittata var. zonata, 152 Agave utahensis, 261 Agave utahensis fa. nuda, 262 Agave utahensis ssp. eborispina, 262 Agave utahensis ssp. kaibabensis, 261 Agave utahensis ssp. nevadensis, 263 Agave utahensis ssp. utahensis, 262 Agave utahensis var. discreta, 262 Agave utahensis var. eborispina, 262 Agave utahensis var. kaibabensis, 261 Agave utahensis var. nevadensis, 262 Agave utahensis var. scaphoidea, 262 Agave utahensis var. utahensis, 262 Agave valenciana, 263 Agave vanderdonckii, 279 Agave vanderdonckii var. horizontalis, 280 Agave vandervinnenii, 40 Agave vangrolae, 146 Agave variegata, 45, 263 Agave vazquezgarciae, 264 Agave ventum-versa, 146 Agave venustuliflora, 265 Agave vera-cruz, 158 Agave vera-cruz var. jacquiniana, 50 Agave verae-crucis, 158 Agave verdensis, 265, 266 Agave verhoekiae, 266 Agave vernae, 158 Agave verschaffeltii, 212 Agave vestita, 230 Agave vexans, 95 Agave vicina, 267 Agave victoriae-reginae, 181, 267 Agave victoriae-reginae cv. compacta, 268 Agave victoriae-reginae fa. compacta, 268 Agave victoriae-reginae fa. dentata, 268

1401 Agave victoriae-reginae fa. ferdinandi-regis, 181, 182 Agave victoriae-reginae fa. latifolia, 268 Agave victoriae-reginae fa. longifolia, 268 Agave victoriae-reginae fa. longispina, 269 Agave victoriae-reginae fa. ornata, 269 Agave victoriae-reginae fa. stolonifera, 269 Agave victoriae-reginae fa. variegata, 268 Agave victoriae-reginae fa. viridis, 226 Agave victoriae-reginae ssp. swobodae, 268 Agave victoriae-reginae ssp. victoriae-reginae, 268 Agave victoriae-reginae var. candida, 268 Agave victoriae-reginae var. compacta, 268 Agave victoriae-reginae var. dentata, 268 Agave victoriae-reginae var. elongata, 268 Agave victoriae-reginae var. latifolia, 268 Agave victoriae-reginae var. laxior, 181 Agave victoriae-reginae var. nigra, 226 Agave victoriae-reginae var. punctata, 268 Agave victoriae-reginae var. stolonifera, 268 Agave victoriae-reginae var. striata, 268 Agave victoriae-reginae var. viridis, 268 Agave villae, 270 Agave villarum, 270 Agave vilmoriniana, 270 Agave virens, 63 Agave virginica, 45, 271, 272 Agave virginica fa. tigrina, 271 Agave virginica ssp. lata, 271 Agave virginica ssp. virginica, 271 Agave virginica var. polyantha, 271 Agave virginica var. tigrina, 271 Agave viridissima, 40 Agave vittata, 157 Agave vivipara, 40, 50, 54, 78, 107, 171, 272, 325, 343 Agave vivipara cv. marginata, 52 Agave vivipara cv. variegata, 53 Agave vivipara var. bermejo, 51 Agave vivipara var. bromeliifolia, 50 Agave vivipara var. cabaiensis, 267 Agave vivipara var. cuebensis, 267 Agave vivipara var. deweyana, 51 Agave vivipara var. letonae, 52 Agave vivipara var. nivea, 53 Agave vivipara var. rubescens, 50 Agave vivipara var. sargentii, 53 Agave vivipara var. woodrowii, 53 Agave vizcainoensis, 273 Agave wallisii, 274 Agave warelliana, 274, 275 Agave washingtonensis, 40 Agave watsonii, 40 Agave weberi, 275 Agave weingartii, 191 Agave weissenburgensis, 40 Agave wendtii, 276 Agave wercklei, 40, 276, 277 Agave whitackeri, 224 Agave wiesenbergensis, 40 Agave wiesenburgensis, 40

1402 Agave wightii, 51 Agave wildringii, 40 Agave willdingii, 277, 278 Agave winteriana, 278 Agave wislizeni, 59 Agave wislizeni ssp. parrasana, 195, 196 Agave wocomahi, 278, 279 Agave woodrowii, 53 Agave wrightii, 230 Agave xalapensis, 185 Agave xylacantha, 279 Agave xylonacantha, 279 Agave xylonacantha [?] mediopicta, 279 Agave xylonacantha [?] minor, 280 Agave xylonacantha [?] vittata, 137 Agave xylonacantha var. horizontalis, 279 Agave xylonacantha var. latifolia, 279 Agave xylonacantha var. macracantha, 279 Agave xylonacantha var. torta, 279 Agave yaquiana, 51 Agave yavapaiensis, 280, 281 Agave yuccifolia, 243 Agave yuccifolia var. caespitosa, 243 Agave yuccifolia var. spicata, 243 Agave yuccifolia var. viridis, 243, 244 Agave yxtli, 50 Agave zapopanensis, 281 Agave zapupe, 51 Agave zebra, 281, 282 Agave zonata, 46 Agave zuccarinii, 40 Agriodendron, 486 Albuca, 1231 Albuca batteniana, 1233 Albuca bracteata, 1233, 1260 Albuca buffelspoortensis, 1234 Albuca clanwilliamae-gloria, 1234 Albuca cremnophila, 1235 Albuca crudenii, 1235 Albuca deaconii, 1235 Albuca dilucula, 1236 Albuca elliotii, 1239 Albuca gariepensis, 1236 Albuca gildenhuysii, 1236 Albuca granulata, 1239 Albuca imbricata, 1237 Albuca juncifolia, 1237 Albuca karachabpoortensis, 1237 Albuca kirstenii, 1237 Albuca lebaensis, 1238 Albuca minima, 1239 Albuca nelsonii, 1238 Albuca pendula, 1238 Albuca scabrocostata, 1239 Albuca shawii, 1239 Albuca spiralis, 1239 Albuca tenuifolia, 1239, 1240 Albuca thermarum, 1240 Albuca trichophylla, 1239

Index Albuca unifoliata, 1240 Albuca viridiflora, 1239 Albucca, 1281 Aletris guineensis, 1367 Aletris hyacinthoides, 1367, 1381 Aletris hyacinthoides var. guineensis, 1367 Aletris hyacinthoides var. zeylanica, 1355, 1382 Aletris zeylanica, 1382 Aletris zeylanica var. β, 1362 Algastoloba, 483 Allardtia, 1107 Allemptauminia, 486 Alleptauminia, 486 Alliaceae, 425 Allibertia, 22 Allibertia intermedia, 271 Aloaceae, 475 Aloe, 485, 490 Aloe aaata, 491, 492 Aloe aageodonta, 492 Aloe abhaica, 492 Aloe abyssicola, 492 Aloe abyssinica, 491, 494, 524, 552 Aloe abyssinica var. peacockii, 552 Aloe abyssinica var. percrassa, 631 Aloe acinacifolia, 753 Aloe acinacifolia var. angustifolia, 753 Aloe acinacifolia var. laetevirens, 753 Aloe aculeata, 493 Aloe acuminata, 577 Aloe acuminate var. major, 577 Aloe acutissima, 493 Aloe acutissima ssp. itampolensis, 494 Aloe acutissima var. acutissima, 493 Aloe acutissima var. antanimorensis, 493 Aloe acutissima var. berevoana, 513 Aloe acutissima var. fiherenensis, 494 Aloe acutissima var. isaloana, 494 Aloe acutissima var. itampoloana, 494 Aloe adigratana, 494 Aloe aestivalis, 583 Aloe aethiopica, 552 Aloe affinis, 494 Aloe africana, 494 Aloe africana var. angustior, 495 Aloe africana var. latifolia, 495 Aloe agavefolia, 491 Aloe agrophila, 516 Aloe ahmarensis, 495 Aloe alaotrensis, 495 Aloe albicans, 794 Aloe albida, 496 Aloe albiflora, 496 Aloe albispina, 491 Aloe albocincta, 660 Aloe albopicta, 525 Aloe albostriata, 496 Aloe albovestita, 497 Aloe aldabrensis, 497

Index Aloe alexandrei, 497 Aloe alfredii, 497 Aloe allochroa, 497 Aloe alooides, 498 Aloe altilinea, 772 Aloe altilinea var. denticulata, 776 Aloe altimatsiatrae, 559 Aloe amanensis, 593 Aloe ambigens, 498 Aloe ambositrae, 498 Aloe ambrensis, 499 Aloe amicorum, 499 Aloe ammophila, 499 Aloe amoena, 562 Aloe ampefyana, 499, 500 Aloe amudatensis, 500 Aloe analavelonensis, 500 Aloe andersonii, 500 Aloe andohahelensis, 665 Aloe andongensis, 500 Aloe andongensis var. andongensis, 500 Aloe andongensis var. repens, 501 Aloe andringitrensis, 501 Aloe angelica, 501 Aloe angiensis, 682 Aloe angiensis var. kitaliensis, 682 Aloe angolensis, 501 Aloe angulata, 757 Aloe angulata var. striata, 752 Aloe angustifolia, 495, 673, 759 Aloe angustifolia var. laevis, 757 Aloe anivoranoensis, 501 Aloe ankabeensis, 491 Aloe ankaranensis, 640 Aloe ankoberensis, 502 Aloe anodonta, 502 Aloe anomala, 771 Aloe anosyana, 502 Aloe ansoultae, 502 Aloe antandroi, 502 Aloe antandroi ssp. toliarana, 502 Aloe antoetrana, 503 Aloe antonii, 503 Aloe antoninii, 490 Aloe antsingyensis, 503 Aloe arabica, 491, 631 Aloe arachnoidea, 774 Aloe arachnoidea var. communis, 774 Aloe arachnoidea var. pellucens, 789 Aloe arachnoidea var. pumila, 789 Aloe arachnoidea var. reticulata, 806 Aloe arachnoidea var. translucens, 789 Aloe arachnoides, 774 Aloe arborea, 491, 503 Aloe arborescens, 503 Aloe arborescens ssp. mzimnyati, 504 Aloe arborescens var. frutescens, 504 Aloe arborescens var. milleri, 504 Aloe arborescens var. natalensis, 504

1403 Aloe arborescens var. pachythyrsa, 504 Aloe arborescens var. ucriae, 491 Aloe arborescens var. viridifolia, 504 Aloe archeri, 504 Aloe archeri ssp. tugenensis, 670 Aloe arenicola, 504, 505 Aloe argenticauda, 505 Aloe argentifolia, 505 Aloe argyrostachys, 505 Aloe aristata, 505, 776 Aloe aristata var. leiophylla, 505 Aloe aristata var. parvifolia, 505 Aloe armatissima, 506 Aloe arneodoi, 506 Aloe aspera, 703, 771 Aloe asperifolia, 506 Aloe asperiuscula, 813 Aloe asperula, 772 Aloe atherstonei, 637 Aloe atrovirens, 789 Aloe attenuata, 776 Aloe attenuata var. clariperla, 777 Aloe audhalica, 672 Aloe aufensis, 507 Aloe aurantiaca, 661 Aloe aurelienii, 507 Aloe ausana, 674 Aloe austroarabica, 507 Aloe austrosudanica, 507 Aloe babatiensis, 507, 508 Aloe bainesii, 510 Aloe bainesii var. barberae, 510 Aloe bakeri, 508 Aloe ballii, 508 Aloe ballii var. ballii, 509 Aloe ballii var. makurupiniensis, 509 Aloe ballyi, 509 Aloe bamangwatensis, 684 Aloe barbadensis, 675 Aloe barbadensis var. chinensis, 675 Aloe barbara-jeppeae, 510 Aloe barberae, 510 Aloe barbertoniae, 510 Aloe barendii, 513 Aloe bargalensis, 510 Aloe barteri, 520, 600, 653 Aloe barteri var. dahomensis, 520 Aloe barteri var. lutea, 653 Aloe barteri var. sudanica, 520 Aloe baumii, 684 Aloe beankaensis, 511 Aloe bedinghausii, 745 Aloe beguinii, 745 Aloe beguinii var. perfectior, 746 Aloe belavenokensis, 511 Aloe belitsakensis, 511 Aloe bella, 512 Aloe bellatula, 512 Aloe beniensis, 541

1404 Aloe benishangulana, 512, 513 Aloe bequaertii, 682 Aloe berevoana, 513 Aloe bergeriana, 513 Aloe berhana, 541 Aloe bernadettae, 513 Aloe bernardii, 513 Aloe bertemariae, 514 Aloe betsileensis, 514 Aloe bicarinata, 709 Aloe bicolor, 755 Aloe bicomitum, 514 Aloe boastii, 531 Aloe boehmii, 593 Aloe boiteaui, 514 Aloe bolusii, 495 Aloe boranensis, 626 Aloe boscawenii, 515 Aloe bosseri, 515 Aloe bowiea, 515 Aloe boylei, 515 Aloe boylei ssp. boylei, 515 Aloe boylei ssp. major, 516 Aloe braamvanwykii, 516 Aloe brachyphylla, 756 Aloe brachystachys, 516 Aloe bradlyana, 789 Aloe branddraaiensis, 517 Aloe brandhamii, 517 Aloe brevifolia, 517, 612 Aloe brevifolia var. brevifolia, 517 Aloe brevifolia var. depressa, 517 Aloe brevifolia var. postgenita, 517 Aloe brevifolia var. serra, 518 Aloe brevis, 795 Aloe breviscapa, 518 Aloe broomii, 518 Aloe broomii var. broomii, 518 Aloe broomii var. tarkaensis, 518 Aloe brownii, 622 Aloe brunneodentata, 518 Aloe brunneo-punctata, 623 Aloe brunneostriata, 518, 519 Aloe brunnthaleri, 632 Aloe bruynsii, 519 Aloe buchananii, 519 Aloe buchlohii, 519 Aloe buettneri, 519 Aloe buhrii, 520 Aloe bukobana, 520 Aloe bulbicaulis, 521 Aloe bulbillifera, 521 Aloe bulbillifera var. bulbillifera, 521 Aloe bulbillifera var. paulianiae, 521 Aloe bullockii, 521 Aloe bullulata, 702 Aloe burgersfortensis, 522 Aloe bussei, 522 Aloe butiabana, 523

Index Aloe buzairiensis, 523 Aloe calcairophila, 523 Aloe calcairophylla, 523 Aloe calidophila, 523 Aloe calliantha, 524 Aloe cameronii, 524 Aloe cameronii var. bondana, 524 Aloe cameronii var. cameronii, 524 Aloe cameronii var. dedzana, 524 Aloe camperi, 524 Aloe camperi ssp. adigratana, 494 Aloe campylosiphon, 593 Aloe canarina, 525 Aloe candelabrum, 557, 667 Aloe candicans, 544, 753 Aloe candicans var. fallax, 544 Aloe canii, 525 Aloe canis, 525 Aloe cannellii, 525 Aloe capitata, 526 Aloe capitata var. angavoana, 526 Aloe capitata var. capitata, 526 Aloe capitata var. cipolinicola, 533 Aloe capitata var. gneissicola, 567 Aloe capitata var. quartziticola, 526 Aloe capitata var. silvicola, 526 Aloe capitata var. trachyticola, 668 Aloe capmanambatoensis, 527 Aloe caricina, 618 Aloe carinata, 757, 758 Aloe carinata var. subglabra, 757 Aloe carnea, 527 Aloe carolineae, 527 Aloe caroliniana, 271 Aloe carowii, 657 Aloe cascadensis, 661 Aloe castanea, 527, 528 Aloe castellorum, 528 Aloe castilloniae, 528 Aloe cataractarum, 528 Aloe catengiana, 529 Aloe cephalophora, 529 Aloe cernua, 526 Aloe chabaudii, 529 Aloe chabaudii var. chabaudii, 529, 530 Aloe chabaudii var. mlanjeana, 530 Aloe chabaudii var. verekeri, 530 Aloe challisii, 530 Aloe charlotteae, 530 Aloe cheranganiensis, 531 Aloe chimanimaniensis, 664 Aloe chinensis, 675 Aloe chloracantha, 779 Aloe chlorantha, 531 Aloe chloroleuca, 491 Aloe chludowii, 746 Aloe chortolirioides, 531 Aloe chortolirioides var. boastii, 531 Aloe chortolirioides var. chortolirioides, 531

Index Aloe chortolirioides var. woolliana, 531 Aloe christianii, 532 Aloe chrysostachys, 532 Aloe ciliaris, 532 Aloe ciliaris fa. flanaganii, 532 Aloe ciliaris fa. gigas, 533 Aloe ciliaris fa. tidmarshii, 533 Aloe ciliaris var. ciliaris, 532 Aloe ciliaris var. flanaganii, 532 Aloe ciliaris var. redacta, 533 Aloe ciliaris var. tidmarshii, 533 Aloe cinnabarina, 491 Aloe cipolinicola, 533 Aloe citrea, 533 Aloe citrina, 533, 534 Aloe clariperla, 777 Aloe clarkei, 534 Aloe classenii, 534 Aloe claviflora, 534 Aloe coarctata, 779 Aloe collenetteae, 535 Aloe collina, 535 Aloe commelinii, 491 Aloe commixta, 535 Aloe commutata, 536, 600 Aloe commutata var. tricolor, 536 Aloe comosa, 536 Aloe comosibracteata, 570 Aloe compacta, 601 Aloe compressa, 536 Aloe compressa var. compressa, 536 Aloe compressa var. paucituberculata, 536 Aloe compressa var. rugosquamosa, 649 Aloe compressa var. schistophila, 536 Aloe comptonii, 612 Aloe concinna, 659, 813 Aloe condyae, 537 Aloe confusa, 537 Aloe congdonii, 537 Aloe congesta, 702 Aloe congolensis, 491 Aloe conifera, 537 Aloe conifera ssp. conifera, 537 Aloe conifera ssp. pervagata, 538 Aloe consobrina, 491 Aloe conspurcata, 759 Aloe conspurcata var. truncata, 759 Aloe conspurcata var. unilateralis, 759 Aloe constricta, 684 Aloe contigua, 579 Aloe cooperi, 538 Aloe cooperi ssp. pulchra, 655 Aloe corallina, 538 Aloe corbisieri, 623 Aloe corderoyi, 538 Aloe cordifolia, 813 Aloe craibii, 538 Aloe crassipes, 539 Aloe cremersii, 578

1405 Aloe cremnophila, 539 Aloe croucheri, 759 Aloe cryptoflora, 539 Aloe cryptopoda, 539, 540 Aloe curta, 773 Aloe cuspidata, 773 Aloe cylindrica, 771 Aloe cylindrica var. rigida, 771 Aloe cymbaefolia, 783 Aloe cymbiformis, 783 Aloe cyrillei, 578 Aloe cyrtophylla, 540 Aloe dabenorisana, 540 Aloe darainensis, 541 Aloe davyana, 570 Aloe davyana var. subolifera, 570 Aloe dawei, 541 Aloe debrana, 541 Aloe decaryi, 542 Aloe decipiens, 762 Aloe decora, 534 Aloe decorsei, 542 Aloe decumbens, 542 Aloe decurva, 542 Aloe decurvidens, 589 Aloe defalcata, 491, 611 Aloe deflexidens, 491 Aloe deinacantha, 542 Aloe delicatifolia, 543 Aloe delphinensis, 543 Aloe deltoidea, 703 Aloe deltoideodonta, 543 Aloe deltoideodonta fa. latifolia, 579 Aloe deltoideodonta fa. longifolia, 579 Aloe deltoideodonta ssp. amboahangyensis, 543 Aloe deltoideodonta ssp. brevifolia, 543 Aloe deltoideodonta ssp. candicans, 544 Aloe deltoideodonta ssp. esomonyensis, 545 Aloe deltoideodonta ssp. fallax, 544 Aloe deltoideodonta subfa. variegata, 579 Aloe deltoideodonta var. brevifolia, 543 Aloe deltoideodonta var. candicans, 543 Aloe deltoideodonta var. contigua, 579 Aloe deltoideodonta var. deltoideodonta, 544 Aloe deltoideodonta var. fallax, 544 Aloe deltoideodonta var. intermedia, 544 Aloe deltoideodonta var. ruffingiana, 544 Aloe deltoideodonta var. typica, 544 Aloe denticulata, 776 Aloe dependens, 631 Aloe depressa, 517 Aloe derbetzei, 746 Aloe descoingsii, 545 Aloe descoingsii ssp. augustina, 545 Aloe descoingsii ssp. descoingsii, 545 Aloe deserti, 545 Aloe dewetii, 546 Aloe dewinteri, 546 Aloe dhalensis, 672

1406 Aloe dhufarensis, 546 Aloe dichotoma, 546 Aloe dichotoma ssp. dichotoma, 546 Aloe dichotoma ssp. pillansii, 634 Aloe dichotoma ssp. ramosissima, 547 Aloe dichotoma var. montana, 546 Aloe dichotoma var. ramosissima, 547 Aloe dictyodes, 755 Aloe dinteri, 547 Aloe diolii, 547, 548 Aloe distans, 612 Aloe disticha, 601, 759, 765 Aloe disticha var. δ, 757 Aloe disticha var. plicatilis, 636 Aloe divaricata, 548 Aloe divaricata ssp. deinacantha, 542 Aloe divaricata ssp. divaricata, 548 Aloe divaricata ssp. tulearensis, 548 Aloe divaricata ssp. vaotsohy, 549 Aloe divaricata var. deinacantha, 542 Aloe divaricata var. rosea, 549 Aloe djiboutiensis, 549 Aloe doddsiorum, 549 Aloe doei, 594 Aloe doei var. lavranosii, 594 Aloe dolomitica, 680 Aloe dominella, 549 Aloe dorotheae, 549 Aloe dorsalis, 491 Aloe downsiana, 550 Aloe drepanophylla Baker, 491 Aloe droseroides, 550 Aloe duckeri, 550 Aloe dumetorum, 553 Aloe dyeri, 550 Aloe echinata, 577 Aloe ecklonis, 551 Aloe edentata, 559 Aloe edouardii, 551 Aloe edulis, 600 Aloe elata, 551 Aloe elegans, 552 Aloe elegantissima, 552 Aloe elgonica, 553 Aloe elizae, 491 Aloe elkerriana, 553 Aloe ellenbeckii, 553 Aloe ellenbergeri, 505 Aloe elongata, 675 Aloe eminens, 553 Aloe engleri, 654 Aloe enotata, 677 Aloe ensifolia, 753 Aloe erecta, 795 Aloe erecta var. laetivirens, 795 Aloe eremophila, 554 Aloe erensii, 554 Aloe ericahenriettae, 554 Aloe ericetorum, 554

Index Aloe erinacea, 554 Aloe eru, 494, 525 Aloe eru fa. erecta, 525 Aloe eru fa. glauca, 525 Aloe eru fa. maculata, 525 Aloe eru fa. parvipunctata, 525 Aloe eru var. cornuta, 525 Aloe erythrophylla, 555 Aloe erythrophylla ssp. erythrophylla, 555 Aloe erythrophylla ssp. major, 555 Aloe esculenta, 555 Aloe estevei, 555 Aloe eumassawana, 556 Aloe excavata, 757 Aloe excelsa, 556 Aloe excelsa var. breviflora, 556 Aloe excelsa var. excelsa, 556 Aloe eximia, 557 Aloe expansa, 771 Aloe expansa var. major, 771 Aloe eylesii, 645 Aloe falcata, 557 Aloe fasciata, 787 Aloe fasciata var. major, 787 Aloe fasciata var. minor, 787 Aloe ferox, 557, 604 Aloe ferox var. erythrocarpa, 557 Aloe ferox var. galpinii, 557 Aloe ferox var. hanburyi, 557 Aloe ferox var. incurvata, 557 Aloe ferox var. subferox, 557 Aloe ferox var. xanthostachys, 604 Aloe fianarantsoae, 551 Aloe fibrosa, 558 Aloe fievetii, 558 Aloe fievetii ssp. fievetii, 558 Aloe fievetii ssp. johannis-baptistei, 559 Aloe fievetii var. altimatsiatrae, 559 Aloe fievetii var. ambatofinandrahanensis, 559 Aloe fimbrialis, 559 Aloe flabelliformis, 636 Aloe flava, 675 Aloe flavispina, 491 Aloe fleurentiniorum, 559 Aloe fleuretteana, 559 Aloe flexilifolia, 560 Aloe floramaculata, 654 Aloe florenceae, 560 Aloe foetida, 329 Aloe foliolosa, 704 Aloe fontainei, 560 Aloe forbesii, 560 Aloe fosteri, 561 Aloe fouriei, 561 Aloe fragilis, 561 Aloe framesii, 562 Aloe francombei, 562 Aloe fraskii, 491 Aloe friisii, 562

Index Aloe frutescens, 504 Aloe fruticosa, 503 Aloe fulgens, 504 Aloe fulleri, 563 Aloe galpinii, 557 Aloe gariepensis, 563 Aloe gariusiana, 563 Aloe gasterioides, 491 Aloe gautieri, 564 Aloe gerstneri, 564 Aloe ghibensis, 564 Aloe gilbertii, 565 Aloe gilbertii ssp. gilbertii, 565 Aloe gilbertii ssp. megalacanthoides, 565 Aloe gillettii, 565 Aloe gillilandii, 650 Aloe glabra, 758 Aloe glabra var. brevifolia, 758 Aloe glabra var. major, 758 Aloe glabrata, 777 Aloe glabrata var. concolor, 771 Aloe glabrata var. perviridis, 772 Aloe glabrescens, 566 Aloe glauca, 566 Aloe glauca var. elatior, 566 Aloe glauca var. glauca, 566 Aloe glauca var. humilior, 566 Aloe glauca var. major, 566 Aloe glauca var. minor, 566 Aloe glauca var. muricata, 566 Aloe glauca var. spinosior, 566 Aloe globuligemma, 566 Aloe gloveri, 576 Aloe gneissicola, 567 Aloe gossweileri, 567 Aloe gracilicaulis, 568 Aloe graciliflora, 568 Aloe gracilis, 535, 568 Aloe gracilis var. decumbens, 542 Aloe grahamii, 491 Aloe graminicola, 593 Aloe graminifolia, 618 Aloe granata, 795 Aloe granata var. major, 795 Aloe granata var. minor, 795 Aloe grandidentata, 568 Aloe graniticola, 569 Aloe grata, 569 Aloe greatheadii, 569 Aloe greatheadii var. davyana, 570 Aloe greatheadii var. greatheadii, 570 Aloe greenii, 491, 678 Aloe greenwayi, 595 Aloe grisea, 571 Aloe guerrae, 571 Aloe guillaumetii, 571 Aloe guineensis, 1367 Aloe guttata, 752 Aloe haemanthifolia, 571

1407 Aloe haggeherensis, 572 Aloe hahnii, 572 Aloe hanburiana, 660 Aloe hardyi, 572 Aloe harlana, 573 Aloe harmsii, 549, 550 Aloe haroniensis, 573 Aloe haworthioides, 573 Aloe haworthioides var. aurantiaca, 573 Aloe haworthioides var. haworthioides, 573 Aloe hazeliana, 573 Aloe hazeliana var. hazeliana, 573 Aloe hazeliana var. howmanii, 574 Aloe hebes, 783 Aloe helenae, 574 Aloe heliderana, 574 Aloe hemmingii, 575 Aloe hendrickxii, 575 Aloe herbacea, 789 Aloe hereroensis, 575 Aloe hereroensis var. hereroensis, 575 Aloe hereroensis var. lutea, 575 Aloe hereroensis var. orpeniae, 575 Aloe heteracantha, 490 Aloe hexapetala, 491 Aloe heybensis, 575 Aloe hijazensis, 528 Aloe hildebrandtii, 576 Aloe hlangapensis, 576 Aloe hlangapies, 576 Aloe hlangapitis, 576 Aloe hoffmannii, 576 Aloe horombensis, 543, 544 Aloe horombensis ssp. amboahangyensis, 543 Aloe horombensis ssp. fallax, 544 Aloe horombensis ssp. ruffingiana, 545 Aloe horrida, 491, 558 Aloe howmanii, 574 Aloe humbertii, 577 Aloe humilis, 577 Aloe humilis subvar. minor, 577 Aloe humilis subvar. semiguttata, 577 Aloe humilis var. acuminata, 577 Aloe humilis var. candollei, 577 Aloe humilis var. echinata, 577 Aloe humilis var. humilis, 577 Aloe humilis var. incurva, 577 Aloe humilis var. macilenta, 491 Aloe humilis var. semiguttata, 577 Aloe humilis var. suberecta, 577 Aloe humilis var. subtuberculata, 577 Aloe huntleyana, 578 Aloe hyacinthoides, 1367 Aloe hyacinthoides var. guineensis, 1367 Aloe hyacinthoides var. zeylanica, 1382 Aloe hybrida, 772 Aloe hybrida var. gloriosa, 746 Aloe ibitiensis, 578 Aloe ibityensis, 578

1408 Aloe ifanadianae, 578 Aloe ikiorum, 579 Aloe imalotensis, 579 Aloe imalotensis var. imalotensis, 579 Aloe imalotensis var. longiracemosa, 579 Aloe imbricata, 705, 747 Aloe imerinensis, 579 Aloe immaculata, 579 Aloe inamara, 580 Aloe inconspicua, 580 Aloe incurva, 577 Aloe indica, 675 Aloe indurata, 813 Aloe inermis, 580 Aloe inexpectata, 580 Aloe inopinata, 581 Aloe integra, 581 Aloe intermedia, 544, 758 Aloe intermedia var. asperrima, 758 Aloe inyangensis, 581 Aloe inyangensis var. inyangensis, 581 Aloe inyangensis var. kimberleyana, 581 Aloe irafensis, 582 Aloe isaloensis, 582 Aloe ithya, 582 Aloe ivakoanyensis, 583 Aloe jacksonii, 583 Aloe jacobseniana, 490 Aloe jawiyon, 583 Aloe jeppeae, 583 Aloe jex-blakeae, 650 Aloe jibisana, 584 Aloe johannis, 584 Aloe johannis-bernardii, 584 Aloe johannis-philippei, 585 Aloe johnstonii, 618 Aloe jucunda, 585 Aloe juddii, 585 Aloe juttae, 632 Aloe juvenna, 585 Aloe kahinii, 585 Aloe kamnelii, 586 Aloe kaokoensis, 586 Aloe karasbergensis, 586 Aloe karasbergensis ssp. hunsbergensis, 586 Aloe karasbergensis ssp. karasbergensis, 586 Aloe keayi, 586 Aloe kedongensis, 587 Aloe kefaensis, 587 Aloe keithii, 627 Aloe ketabrowniorum, 587 Aloe khamiesensis, 588 Aloe kilifiensis, 588 Aloe kirkii, 605 Aloe knersvlakensis, 588 Aloe kniphofioides, 589 Aloe koenenii, 589 Aloe komaggasensis, 589 Aloe komatiensis, 589

Index Aloe kouebokkeveldensis, 590 Aloe krapohliana, 590 Aloe krapohliana var. dumoulinii, 590 Aloe krapohliana var. krapohliana, 590 Aloe kraussii, 496, 591 Aloe kraussii var. minor, 496 Aloe kulalensis, 591 Aloe kwasimbana, 591 Aloe labiaflava, 570 Aloe labworana, 592 Aloe labworana ssp. labworana, 592 Aloe labworana ssp. longifolia, 592 Aloe laeta, 592 Aloe laeta var. laeta, 592 Aloe laeta var. maniaensis, 592 Aloe laetepuncta, 757 Aloe laetevirens, 808 Aloe laevis, 757 Aloe lanata, 592 Aloe lanuriensis, 682 Aloe lanzae, 675 Aloe lapaixii, 746 Aloe lapaixii var. latifolia, 746 Aloe lastii, 516 Aloe latens, 593 Aloe lateritia, 593 Aloe lateritia var. graminicola, 593 Aloe lateritia var. kitaliensis, 682 Aloe lateritia var. lateritia, 593 Aloe latifolia, 602 Aloe lavranosii, 594 Aloe laxiflora, 568 Aloe laxissima, 684 Aloe leachii, 594 Aloe leandrii, 594 Aloe leedalii, 594 Aloe lensayuensis, 595 Aloe lepida, 595 Aloe leptocaulon, 502 Aloe leptophylla, 602 Aloe leptophylla var. stenophylla, 602 Aloe leptosiphon, 595 Aloe lettyae, 596 Aloe leucantha, 491 Aloe liliputana, 596 Aloe limpida, 801 Aloe lindenii, 596 Aloe linearifolia, 596 Aloe lineata, 597 Aloe lineata var. glaucescens, 597 Aloe lineata var. lineata, 597 Aloe lineata var. muirii, 597 Aloe lineata var. viridis, 597 Aloe lingua, 636, 755, 758, 759 Aloe lingua var. angulata, 757 Aloe lingua var. angustifolia, 759 Aloe lingua var. brevifolia, 759 Aloe lingua var. crassifolia, 759 Aloe lingua var. multifaria, 757

Index Aloe linguaefolia, 752 Aloe linguaefolia var. angustifolia, 752 Aloe linguaefolia var. latifolia, 752 Aloe linguaeformis, 636 Aloe linguiformis, 752, 759 Aloe linguiformis var. verrucosa, 758 Aloe littoralis, 597 Aloe lolwensis, 597 Aloe lomatophylloides, 598 Aloe longiaristata, 505 Aloe longibracteata, 598 Aloe longiflora, 491 Aloe longistyla, 598 Aloe luapulana, 598 Aloe lucile-allorgeae, 599 Aloe lugardiana, 684 Aloe lukeana, 599 Aloe luntii, 599 Aloe lusitanica, 589 Aloe lutescens, 599 Aloe lynchii, 746 Aloe macchiata, 752 Aloe maclaughlinii, 607 Aloe macleayi, 600 Aloe macowanii, 661 Aloe macra, 600 Aloe macracantha, 491 Aloe macrocarpa, 600 Aloe macrocarpa ssp. wollastonii, 682 Aloe macrocarpa var. major, 600 Aloe macroclada, 600 Aloe macrosiphon, 601 Aloe maculata, 601, 624, 755 Aloe maculata ssp. ficksburgensis, 601 Aloe maculata ssp. maculata, 601 Aloe maculata var. obliqua, 755 Aloe maculata var. pulchra, 764 Aloe maculate var. ficksburgensis, 601 Aloe maculosa, 601 Aloe madecassa, 544 Aloe madecassa var. lutea, 544 Aloe magnidentata, 608 Aloe mahraensis, 602 Aloe makayana, 602 Aloe manandonae, 603 Aloe mandotoensis, 603 Aloe mandrarensis, 603 Aloe mangeaensis, 603 Aloe maningoryensis, 603 Aloe margaritifera, 793 Aloe margaritifera var. major, 795 Aloe margaritifera var. maxima, 793 Aloe margaritifera var. media, 795 Aloe margaritifera var. minima, 795 Aloe margaritifera var. minor, 795 Aloe marginalis, 642 Aloe marginata, 642, 794 Aloe marlothii, 604 Aloe marlothii ssp. orientalis, 604

1409 Aloe marlothii var. bicolor, 604 Aloe marlothii var. marlothii, 604 Aloe marsabitensis, 654 Aloe marshallii, 589 Aloe martialii, 605 Aloe massawana, 605 Aloe massawana ssp. massawana, 605 Aloe massawana ssp. sakoankenke, 605 Aloe mawii, 606 Aloe mayottensis, 607 Aloe mccoyi, 607 Aloe mcloughlinii, 607 Aloe medishiana, 607 Aloe megalacantha, 607 Aloe megalacantha ssp. alticola, 608 Aloe megalacantha ssp. megalacantha, 608 Aloe megalocarpa, 608 Aloe melanacantha, 608 Aloe melanacantha var. erinacea, 555 Aloe melsetterensis, 664 Aloe menachensis, 608 Aloe mendesii, 609 Aloe menyharthii, 609 Aloe menyharthii ssp. ensifolia, 609 Aloe menyharthii ssp. menyharthii, 610 Aloe meruana, 532 Aloe metallica, 610 Aloe meyeri, 610 Aloe micracantha, 516, 610 Aloe microdonta, 610 Aloe microstigma, 632 Aloe microstigma ssp. framesii, 562 Aloe millotii, 611 Aloe milne-redheadii, 611 Aloe minima, 611, 651 Aloe minima var. blyderivierensis, 611 Aloe minima var. minima, 612 Aloe minor, 795 Aloe mirabilis, 796 Aloe miskatana, 612 Aloe mitis, 491 Aloe mitriformis, 612 Aloe mitriformis ssp. comptonii, 612 Aloe mitriformis ssp. distans, 612 Aloe mitriformis ssp. mitriformis, 613 Aloe mitriformis var. albispina, 491 Aloe mitriformis var. angustior, 612 Aloe mitriformis var. brevifolia, 612 Aloe mitriformis var. commelinii, 491 Aloe mitriformis var. elatior, 613 Aloe mitriformis var. flavispina, 491 Aloe mitriformis var. humilior, 491, 613 Aloe mitriformis var. pachyphylla, 491 Aloe mitriformis var. spinosior, 622 Aloe mitriformis var. spinulosa, 491 Aloe mitriformis var. xanthacantha, 491 Aloe mitsioana, 613 Aloe mketiensis, 623 Aloe mkushiana, 613

1410 Aloe mocamedensis, 614 Aloe modesta, 614 Aloe molederana, 614 Aloe mollis, 760 Aloe momccoyae, 614 Aloe monotropa, 615 Aloe montana, 546 Aloe monteiroae, 627 Aloe monticola, 615 Aloe montis-nabro, 615 Aloe morijensis, 615 Aloe morijensis var. morijensis, 615 Aloe morijensis var. ojonokae, 616 Aloe morogoroensis, 522 Aloe mortolensis, 746 Aloe mossurilensis, 616 Aloe mottramiana, 616 Aloe mubendiensis, 616 Aloe mucronata, 800 Aloe mudenensis, 616 Aloe muirii, 597 Aloe multicolor, 617 Aloe multifaria, 773 Aloe munchii, 617 Aloe muricata, 566, 608 Aloe murina, 617 Aloe musapana, 617 Aloe mutabilis, 618 Aloe mutans, 570 Aloe mutica, 801 Aloe mwanzana, 601 Aloe myriacantha, 618 Aloe myriacantha var. minor, 496 Aloe mzimbana, 618 Aloe namibensis, 619 Aloe namorokaensis, 619 Aloe neilcrouchii, 619 Aloe neoqaharensis, 619 Aloe neosteudneri, 620 Aloe newtonii, 620 Aloe ngobitensis, 623 Aloe ngongensis, 620, 621 Aloe nicholsii, 621 Aloe niebuhriana, 621 Aloe niensiensis, 621, 622 Aloe nigra, 801 Aloe nigricans, 759 Aloe nigricans var. crassifolia, 752 Aloe nigricans var. denticulata, 759 Aloe nigricans var. fasciata, 752 Aloe nigricans var. latifolia, 752 Aloe nigricans var. marmorata, 757 Aloe nigrimontana, 622 Aloe nitens, 649, 753 Aloe nitida, 762 Aloe nitida var. brevifolia, 752 Aloe nitida var. grandipunctata, 762 Aloe nitida var. major, 762 Aloe nitida var. minor, 762

Index Aloe nitida var. obtusa, 762 Aloe nitida var. parvipunctata, 762 Aloe nobilis, 491, 622 Aloe nobilis var. densifolia, 491 Aloe nordaliae, 594 Aloe nowotnyi, 747 Aloe nubigena, 622 Aloe nugalensis, 622 Aloe nuttii, 623 Aloe nyeriensis, 623 Aloe nyeriensis ssp. kedongensis, 587 Aloe nyikaensis, 624 Aloe obliqua, 755, 764 Aloe obliqua var. fallax, 756 Aloe obscura, 491, 632, 759 Aloe obscura var. truncata, 757 Aloe obtusifolia, 760 Aloe occidentalis, 624 Aloe officinalis, 624 Aloe officinalis var. angustifolia, 624 Aloe oligophylla, 624 Aloe oligospila, 631 Aloe omavandae, 624 Aloe omoana, 625 Aloe orientalis, 625 Aloe orlandoi, 625 Aloe orpeniae, 575 Aloe ortholopha, 625 Aloe otallensis, 626 Aloe otallensis var. elongata, 648 Aloe pachydactylos, 626 Aloe pachygaster, 626 Aloe paedogona, 520 Aloe pallancae, 490 Aloe pallescens, 491 Aloe pallida, 789 Aloe pallidiflora, 570 Aloe palmiformis, 626 Aloe paludicola, 520 Aloe paniculata, 660 Aloe panormitana, 490 Aloe papillosa, 789 Aloe papillosa var. semipapillosa, 790 Aloe parallelifolia, 626, 627 Aloe parva, 810 Aloe parvibracteata, 627 Aloe parvibracteata var. zuluensis, 627 Aloe parvicapsula, 627 Aloe parvicoma, 628 Aloe parvidens, 628 Aloe parviflora, 628 Aloe parvispina, 612, 613 Aloe parvula, 628, 633 Aloe patersonii, 628 Aloe pavelkae, 629 Aloe peacockii, 552 Aloe pearsonii, 629 Aloe peckii, 629, 630 Aloe peglerae, 630

Index Aloe pembana, 630 Aloe pendens, 631 Aloe penduliflora, 631 Aloe pentagona, 705 Aloe pentagona var. spiralis, 705 Aloe percrassa, 631, 669 Aloe percrassa var. albopicta, 669 Aloe percrassa var. menachensis, 609 Aloe percrassa var. saganeitiana, 552 Aloe perdita, 632 Aloe perfectior, 746 Aloe perfoliata, 495, 517, 557, 601, 613, 632, 675 Aloe perfoliata var. α, 535 Aloe perfoliata var. γ, 557 Aloe perfoliata var. e, 557 Aloe perfoliata var. ζ, 517, 557 Aloe perfoliata var. η, 503 Aloe perfoliata var. θ, 601 Aloe perfoliata var. κ, 566 Aloe perfoliata var. λ, 601 Aloe perfoliata var. μ, 632 Aloe perfoliata var. ξ, 613, 662 Aloe perfoliata var. africana, 495 Aloe perfoliata var. arborescens, 503 Aloe perfoliata var. brevifolia, 612 Aloe perfoliata var. ferox, 557 Aloe perfoliata var. glauca, 566 Aloe perfoliata var. humilis, 577 Aloe perfoliata var. lineata, 597 Aloe perfoliata var. mitriformis, 613 Aloe perfoliata var. obscura, 491 Aloe perfoliata var. purpurascens, 662 Aloe perfoliata var. serrulata, 491 Aloe perfoliata var. succotrina, 662 Aloe perfoliata var. vera, 675 Aloe perrieri, 632 Aloe perryi, 633 Aloe petricola, 633 Aloe petrophila, 633 Aloe peyrierasii, 634 Aloe philippei, 634 Aloe picta, 632 Aloe picta var. major, 601 Aloe picta var. minor, 632 Aloe pictifolia, 634 Aloe pienaarii, 634 Aloe pillansii, 634 Aloe pirottae, 635 Aloe planifolia, 755, 783 Aloe platylepis, 491 Aloe platyphylla, 684 Aloe plicatilis, 635 Aloe plicatilis var. major, 636 Aloe plowesii, 636 Aloe pluridens, 637 Aloe pluridens var. beckeri, 637 Aloe pluripuncta, 753 Aloe pole-evansii, 541 Aloe polyphylla, 637

1411 Aloe pongolensis, 627 Aloe pongolensis var. zuluensis, 627 Aloe porphyrostachys, 637, 638 Aloe porphyrostachys ssp. koenenii, 589 Aloe postgenita, 517 Aloe powysiorum, 638 Aloe praetermissa, 638 Aloe pratensis, 638, 639 Aloe pretoriensis, 639 Aloe principis, 490 Aloe prinslooi, 639 Aloe procera, 639, 640 Aloe prolifera, 517 Aloe prolifera var. major, 517 Aloe pronkii, 640 Aloe propagulifera, 640 Aloe prorumpens, 747 Aloe prostrata, 640 Aloe prostrata ssp. pallida, 684 Aloe pruinosa, 641 Aloe pseudoafricana, 495 Aloe pseudoferox, 557 Aloe pseudonigricans, 752 Aloe pseudoparvula, 641 Aloe pseudorigida, 772 Aloe pseudorubroviolacea, 641 Aloe pseudotortuosa, 813 Aloe pubescens, 641, 642 Aloe pulcherrima, 642 Aloe pulchra, 512, 764 Aloe pumila, 772, 789, 793 Aloe pumila var. β, 795 Aloe pumila var. γ, 795 Aloe pumila var. e, 789 Aloe pumila var. arachnoidea, 774 Aloe pumila var. margaritifera, 793 Aloe pumilio, 806 Aloe punctata, 674 Aloe pungens, 491 Aloe purpurascens, 662 Aloe purpurea, 642 Aloe pustuligemma, 642, 643 Aloe pycnacantha, 649 Aloe qaharensis, 643 Aloe quadrangularis, 752 Aloe quehlii, 747 Aloe rabaiensis, 643 Aloe racemosa, 758 Aloe radula, 777 Aloe radula var. major, 772 Aloe radula var. media, 772 Aloe radula var. minor, 772 Aloe ramifera, 794 Aloe ramosa, 546 Aloe ramosissima, 547 Aloe rapanarivoi, 643 Aloe rauhii, 643 Aloe rebmannii, 644 Aloe rebutii, 747

1412 Aloe recurva, 812 Aloe recurvifolia, 498 Aloe reinwardtii, 805 Aloe reitzii, 644 Aloe reitzii var. reitzii, 644 Aloe reitzii var. vernalis, 645 Aloe rendilliorum, 645 Aloe reticulata, 806 Aloe retrospiciens, 645 Aloe retusa, 807 Aloe reynoldsii, 645 Aloe rhodacantha, 566 Aloe rhodesiana, 645 Aloe rhodocincta, 660 Aloe ribauensis, 646 Aloe richardsiae, 646 Aloe richaudii, 646 Aloe richtersveldensis, 610 Aloe rigens, 646 Aloe rigens var. glabrescens, 566 Aloe rigens var. mortimeri, 563 Aloe rigida, 771 Aloe rigida var. expansa, 771 Aloe rigida var. paulo-major, 772 Aloe rivae, 647 Aloe rivierei, 647 Aloe rodolphei, 647 Aloe roeoeslii, 647 Aloe rosea, 647 Aloe rossii, 491 Aloe rouxii, 648 Aloe rubescens, 491 Aloe rubriflora, 828 Aloe rubrodonta, 648 Aloe rubrolutea, 597 Aloe rubroviolacea, 648 Aloe ruffingiana, 545 Aloe rufocincta, 642 Aloe rugosa, 772 Aloe rugosa var. laetevirens, 772 Aloe rugosa var. perviridis, 772 Aloe rugosifolia, 648 Aloe rugosquamosa, 649 Aloe rulkensii, 649 Aloe runcinata, 491 Aloe rupestris, 649 Aloe rupicola, 649 Aloe ruspoliana, 650 Aloe ruspoliana var. dracaeniformis, 645 Aloe ruvuensis, 650 Aloe sabaea, 650 Aloe sahundra, 548 Aloe sakarahensis, 684 Aloe sakarahensis ssp. pallida, 684 Aloe sakarahensis ssp. sakarahensis, 684 Aloe sakoankenke, 605 Aloe salm-dyckiana, 490 Aloe salmdyckiana var. fulgens, 504 Aloe sanguinalis, 650

Index Aloe saponaria, 601, 602 Aloe saponaria var. brachyphylla, 602 Aloe saponaria var. ficksburgensis, 601 Aloe saponaria var. luteo-striata, 601 Aloe saponaria var. obscura, 491 Aloe saponaria var. saponaria, 601 Aloe saronarae, 578 Aloe saudiarabica, 651 Aloe saundersiae, 651 Aloe scabra, 809 Aloe scabrifolia, 651 Aloe schelpei, 651, 652 Aloe schilliana, 652 Aloe schimperi, 631, 652 Aloe schinzii, 597 Aloe schlechteri, 534 Aloe schliebenii, 516 Aloe schmidtiana, 538 Aloe schoelleri, 652 Aloe schoenlandii, 652 Aloe schomeri, 652 Aloe schweinfurthii, 552, 653 Aloe schweinfurthii var. labworana, 592 Aloe scobinifolia, 653 Aloe scorpioides, 653 Aloe secundiflora, 653 Aloe secundiflora var. secundiflora, 654 Aloe secundiflora var. sobolifera, 654 Aloe secundiflora var. tweedieae, 671 Aloe seibanica, 654 Aloe semiglabrata, 773 Aloe semimargaritifera, 793 Aloe semimargaritifera var. glabrata, 772 Aloe semimargaritifera var. major, 772 Aloe semimargaritifera var. maxima, 793 Aloe semimargaritifera var. minor, 772 Aloe semimargaritifera var. multipapillosa, 772 Aloe semimargaritifera var. multiperlata, 773 Aloe sempervivoides, 628 Aloe seretii, 655 Aloe sergoitensis, 655 Aloe serra, 517 Aloe serriyensis, 655 Aloe serrulata, 491 Aloe serrulata var. pallescens, 491 Aloe sessiliflora, 659 Aloe setata, 776 Aloe setata var. major, 776 Aloe setata var. media, 776 Aloe setata var. nigricans, 775 Aloe setosa, 776 Aloe shadensis, 655 Aloe sharoniae, 655 Aloe sheilae, 656 Aloe sigmoidea, 504 Aloe silicicola, 656 Aloe simii, 656 Aloe simoniana, 748 Aloe sinana, 656

Index Aloe sinkatana, 656, 657 Aloe sinuata, 662 Aloe sladeniana, 657 Aloe smaragdina, 748 Aloe sobolifera, 654 Aloe soccotorina, 662 Aloe soccotrina, 662 Aloe soccotrina var. purpurascens, 662 Aloe socialis, 657 Aloe socotorina, 557 Aloe solaiana, 593 Aloe somaliensis, 657 Aloe somaliensis var. marmorata, 657 Aloe sordida, 810 Aloe sororia, 491 Aloe soutpansbergensis, 658 Aloe speciosa, 658 Aloe spectabilis, 658, 659 Aloe spicata, 524, 659 Aloe spinitriaggregata, 659 Aloe spinulosa, 491 Aloe spiralis, 705 Aloe spiralis var. pentagona, 705 Aloe spirella, 705 Aloe splendens, 594 Aloe springatei-neumannii, 659 Aloe spuria, 491 Aloe squarrosa, 659 Aloe stans, 491 Aloe steffanieana, 677 Aloe stenophylla, 774 Aloe stephaninii, 650 Aloe steudneri, 660 Aloe stolonifera, 660 Aloe straussii, 491 Aloe striata, 660 Aloe striata ssp. karasbergensis, 586 Aloe striata ssp. komaggasensis, 589 Aloe striata var. oligospila, 660 Aloe striatula, 660 Aloe striatula fa. conimbricensis, 661 Aloe striatula fa. haworthii, 661 Aloe striatula fa. typica, 661 Aloe striatula var. caesia, 661 Aloe striatula var. striatula, 661 Aloe stuhlmannii, 679 Aloe suarezensis, 661 Aloe subacutissima, 544 Aloe subalbicans, 772 Aloe subalbicans var. acuminata, 772 Aloe subalbicans var. laevior, 772 Aloe subattenuata, 772 Aloe subcarinata, 757 Aloe subcarinata var. striata, 752 Aloe suberecta, 577 Aloe suberecta var. acuminata, 577 Aloe subfasciata, 772 Aloe subferox, 557 Aloe subfulta, 662

1413 Aloe subnigricans, 753 Aloe subnigricans var. canaliculata, 752 Aloe subrigida, 772 Aloe subspicata, 662 Aloe subtortuosa, 813 Aloe subtuberculata, 577 Aloe subulata, 772 Aloe subverrucosa, 758 Aloe subverrucosa var. grandipunctata, 758 Aloe subverrucosa var. parvipunctata, 758 Aloe succotrina, 662 Aloe succotrina var. saxigena, 662 Aloe suffulta, 662 Aloe sulcata, 757 Aloe suprafoliata, 663 Aloe suprafoliolata, 663 Aloe supralaevis, 557 Aloe supralaevis var. hanburyi, 604 Aloe suzannae, 663, 664 Aloe swynnertonii, 664 Aloe tartarensis, 664 Aloe tauri, 664 Aloe tegetiformis, 664, 665 Aloe teissieri, 665 Aloe tenuifolia, 491, 513 Aloe tenuior, 665 Aloe tenuior var. decidua, 665 Aloe tenuior var. densiflora, 665 Aloe tenuior var. glaucescens, 665 Aloe tenuior var. rubriflora, 665 Aloe tenuior var. tenuior, 665 Aloe tenuior var. viridifolia, 665 Aloe termetophila, 665, 666 Aloe tessellata, 810 Aloe tewoldei, 666 Aloe thompsoniae, 666 Aloe thorncroftii, 666 Aloe thraskii, 666 Aloe tidmarshii, 533 Aloe tomentosa, 667 Aloe tomentosa fa. viridiflora, 667 Aloe tongaensis, 667 Aloe tormentorii, 667 Aloe tororoana, 668 Aloe torquata, 813 Aloe torrei, 668 Aloe torrei var. wildii, 681 Aloe tortuosa, 813 Aloe tortuosa var. major, 772 Aloe trachyticola, 668 Aloe trachyticola var. multifolia, 668 Aloe trachyticola var. trachyticola, 668 Aloe translucens, 789 Aloe transvaalensis, 669 Aloe transvaalensis var. stenacantha, 684 Aloe triangularis, 813 Aloe trichosantha, 669 Aloe trichosantha ssp. longiflora, 669 Aloe trichosantha ssp. trichosantha, 669

1414 Aloe trichosantha var. menachensis, 609 Aloe tricolor, 536, 812 Aloe trigona, 753 Aloe trigona var. elongata, 753 Aloe trigona var. obtusa, 762 Aloe trigonantha, 670 Aloe tripetala, 636 Aloe tristicha, 757 Aloe trivialis, 653 Aloe trothae, 521 Aloe tsitongambarikana, 670 Aloe tuberculata, 577 Aloe tugenensis, 670 Aloe tulearensis, 548 Aloe turgida, 808 Aloe turkanensis, 670, 671 Aloe tweedieae, 671 Aloe ucriae, 491 Aloe ukambensis, 671 Aloe umbellata, 602 Aloe umfoloziensis, 671 Aloe uncinata, 672 Aloe vacillans, 672 Aloe vahontsohy, 548 Aloe vallaris, 672 Aloe vanbalenii, 672 Aloe vandermerwei, 673 Aloe vanrooyenii, 673 Aloe vaombe, 673 Aloe vaombe var. poissonii, 673 Aloe vaombe var. vaombe, 673 Aloe vaotsanda, 673 Aloe vaotsohy, 549 Aloe vaotsohy var. rosea, 549 Aloe variegata, 631, 674 Aloe variegata var. haworthii, 674 Aloe varimaculata, 674 Aloe venenosa, 491 Aloe venosa, 812 Aloe venusta, 514, 753 Aloe vera, 662, 675 Aloe vera var. aethiopica, 552 Aloe vera var. angustifolia, 624 Aloe vera var. chinensis, 675 Aloe vera var. lanzae, 675 Aloe vera var. littoralis, 675 Aloe vera var. officinalis, 624 Aloe vera var. wratislaviensis, 675 Aloe verdoorniae, 570 Aloe verecunda, 676 Aloe verrucosa, 758 Aloe verrucosa var. latifolia, 758 Aloe verrucosa var. striata, 758 Aloe verrucosospinosa, 577 Aloe versicolor, 676 Aloe versicolor ssp. steffanieana, 677 Aloe versicolor var. steffanieana, 676 Aloe versicolor var. versicolor, 677 Aloe veseyi, 677

Index Aloe viguieri, 677 Aloe virens, 491 Aloe virens var. macilenta, 491 Aloe virescens, 794 Aloe virginica, 271 Aloe virginieae, 677 Aloe viridiana, 678 Aloe viridiflora, 678 Aloe viscosa, 813 Aloe viscosa var. indurata, 813 Aloe viscosa var. major, 813 Aloe viscosa var. minor, 813 Aloe viscosa var. parvifolia, 813 Aloe vituensis, 678 Aloe vivipara, 272, 273 Aloe vogtsii, 678, 679 Aloe volkensii, 679 Aloe volkensii ssp. multicaulis, 679 Aloe volkensii ssp. volkensii, 679 Aloe vossii, 679 Aloe vryheidensis, 679 Aloe vulcanica, 506 Aloe vulgaris, 675 Aloe wanalensis, 680 Aloe welmelensis, 680 Aloe weloensis, 680, 681 Aloe welwitschii, 681 Aloe werneri, 681 Aloe whitcombei, 681 Aloe wickensii, 681 Aloe wickensii var. lutea, 681 Aloe wildii, 681 Aloe wilsonii, 682 Aloe wollastonii, 682 Aloe woodii, 682 Aloe woolliana, 531, 532 Aloe wrefordii, 682 Aloe xanthacantha, 491, 613 Aloe yavellana, 683 Aloe yemenica, 683 Aloe yuccifolia, 354 Aloe zakamisyi, 683 Aloe zanzibarica, 659 Aloe zebrina, 683 Aloe zeyheri, 510, 755 Aloe zeylanica, 1382 Aloe zombitsiensis, 684 Aloe zombitsiensis ssp. pallida, 684 Aloe zombitsiensis ssp. zombitsiensis, 684 Aloe zubb, 684, 685 Aloe zygorabaiensis, 685 Aloestrela, 486 Aloestrela suzannae, 663 Aloiampaloe, 486 Aloiampelos, 486 Aloiampelos ciliaris, 532 Aloiampelos ciliaris var. redacta, 533 Aloiampelos ciliaris var. tidmarshii, 533 Aloiampelos commixta, 535

Index Aloiampelos decumbens, 542 Aloiampelos gracilis, 568 Aloiampelos juddii, 585 Aloiampelos striatula, 661 Aloiampelos striatula var. caesia, 661 Aloiampelos tenuior, 665 Aloidendron, 486 Aloidendron barberae, 510 Aloidendron dichotomum, 546 Aloidendron eminens, 553 Aloidendron pillansii, 634 Aloidendron ramosissimum, 547 Aloidendron sabaea, 650 Aloidendron tongaense, 667 Aloinella haworthioides, 573 Alolirion, 486 Aloloba, 697 Altulista, 486 Altulisteria, 483 Alworthia, 699 Amalia, 1107 Amaryllidaceae, 425 Amaryllis disticha, 429 Amaryllis radula, 434 Amaryllis toxicaria, 430 Amaryllis umbella, 438 Amelina, 1171 Amischophacelus, 1179 Anacyclia, 859 Ananas, 840 Ananas ananassoides, 840 Ananas comosus, 840 Ananas lucidus, 840 Aneilema, 1171 Aneilema floribundum, 1191 Aneilema succulentum, 1172 Aneilema zebrinum, 1172 Anemarrhena, 11 Anemarrhenaceae, 11 Anoiganthus, 435 Anoplophytum, 1107 Anoplophytum duratii, 1125 Anoplophytum xiphioides, 1160 Anthericaceae, 11, 449 Anthericum acuminatum, 453 Anthericum aloifolium, 715 Anthericum alooides, 715, 731 Anthericum altissimum, 716 Anthericum annuum, 716 Anthericum aphyllum, 1259 Anthericum asphodeloides, 716 Anthericum bisulcata, 718 Anthericum blepharophoron, 830 Anthericum bulbosum, 717 Anthericum campestre, 453 Anthericum canaliculatum, 830 Anthericum ciliatum, 830 Anthericum comosum, 452 Anthericum drepanophyllum, 831

1415 Anthericum falcatum, 831 Anthericum favosum, 721 Anthericum filifolium, 1287 Anthericum frutescens, 723 Anthericum fruticosum, 723 Anthericum fugax, 1259 Anthericum glabrum, 831 Anthericum hamatum, 831 Anthericum incurvum, 723 Anthericum inexpectatum, 453 Anthericum intricatum, 1256 Anthericum involucratum, 831 Anthericum lagopus, 725 Anthericum latifolium, 718, 725 Anthericum longifolium, 830 Anthericum longiscapum, 716 Anthericum longisetosum, 453 Anthericum longituberosum, 452 Anthericum maculatum, 831 Anthericum nutans, 730 Anthericum oocarpum, 832 Anthericum palustre, 831 Anthericum pilosiflorum, 831 Anthericum planifolium, 452 Anthericum praemorsum, 731 Anthericum pugioniforme, 718 Anthericum recurvatum, 830 Anthericum rostratum, 723 Anthericum salteri, 832 Anthericum semibarbatum, 734 Anthericum spongiosum, 831 Anthericum sternbergianum, 452 Anthericum succulentum, 716 Anthericum suffruticosum, 453 Anthericum tabulare, 831 Anthericum tortile, 832 Anthericum vallis-trappii, 452 Anthericum vespertinum, 830 Anthericum weissianum, 831 Aphyllanthaceae, 11 Apicra, 701, 769 Apicra albicans, 794 Apicra anomala, 771 Apicra arachnoidea, 774 Apicra aspera, 771 Apicra aspera var. major, 772 Apicra atrovirens, 789 Apicra attenuata, 776 Apicra bicarinata, 709 Apicra bullulata, 702, 709 Apicra congesta, 702 Apicra cymbaefolia, 783 Apicra deltoidea, 703 Apicra deltoidea var. intermedia, 703 Apicra egregia, 702 Apicra fasciata, 787 Apicra foliolosa, 704 Apicra granata, 795 Apicra imbricata, 705

1416 Apicra jacobseniana, 828 Apicra margaritifera, 793 Apicra margaritifera var. major, 795 Apicra margaritifera var. maxima, 793 Apicra maxima, 793 Apicra minor, 795 Apicra mirabilis, 796 Apicra nigra, 801 Apicra patula, 772 Apicra pentagona, 705 Apicra pentagona var. bullulata, 702 Apicra pentagona var. spirella, 705 Apicra pentagona var. torulosa, 705 Apicra pseudorigida, 772 Apicra pumilio, 806 Apicra radula, 777 Apicra recurva, 812 Apicra reticulata, 806 Apicra retusa, 807 Apicra rigida, 771 Apicra rubriflora, 828 Apicra skinneri, 709 Apicra spiralis, 705 Apicra tortuosa, 813 Apicra translucens, 789 Apicra tricolor, 812 Apicra turgida, 703 Apicra viscosa, 813 Aploleia, 1173 Aploleia multiflora, 1175 Apostasiaceae, 1301 Aprica, 769 Apworthia, 709 Araceae, 457 Ardernia, 1231 Ariodendron, 486 Arisaraceae, 457 Aristaloe, 486 Aristaloe aristata, 505 Asparagaceae, 11, 465, 1293, 1316 Asparagales, 1–4 Asparagopsis, 467 Asparagopsis densiflora, 469 Asparagus, 467 Asparagus aethiopicus var. ternifolius, 469 Asparagus asparagoides, 468–469 Asparagus crassicladus, 469 Asparagus cuscutoides, 1257 Asparagus densiflorus, 469 Asparagus densiflorus cv. Meyersii, 469 Asparagus densiflorus cv. Sprengeri, 469 Asparagus draco, 1339 Asparagus erectus, 471 Asparagus juniperoides, 469 Asparagus klinghardtianus, 471 Asparagus kuisibensis, 468 Asparagus medeoloides, 468 Asparagus medeoloides var. angustifolius, 468 Asparagus medeoloides var. falciformis, 468

Index Asparagus meyersii, 469 Asparagus micranthus, 1257 Asparagus mucronatus, 470 Asparagus multituberosus, 470 Asparagus myriocladus, 469 Asparagus officinalis, 466 Asparagus ovatus, 470 Asparagus oxyacanthus, 470–471 Asparagus sarmentosus var. comatus, 469 Asparagus sarmentosus var. densiflorus, 469 Asparagus setaceus, 466 Asparagus sprengeri, 469 Asparagus striatus, 471 Asparagus ternifolius, 469 Asparagus undulata, 467–471 Aspasia, 1281 Asphodelaceae, 1389 Asphodelaceae s.l., 2 Asphodeline, 478 Asphodelus, 478 Asphodelus aestivus, 479 Aspidistraceae, 1315 Astrolista, 709 Astrolista bicarinata, 709 Astroloba, 701 Astroloba aspera, 771 Astroloba aspera var. major, 772 Astroloba bicarinata, 709 Astroloba bullata, 702 Astroloba bullulata, 702 Astroloba congesta, 702 Astroloba corrugata, 703 Astroloba cremnophila, 704 Astroloba deltoidea, 703 Astroloba deltoidea var. intermedia, 703 Astroloba deltoidea var. turgida, 703 Astroloba dodsoniana, 704 Astroloba egregia, 702 Astroloba egregia var. fardeniana, 702 Astroloba foliolosa, 704 Astroloba foliolosa ssp. congesta, 702 Astroloba hallii, 702 Astroloba herrei, 704 Astroloba muricata, 703 Astroloba pentagona, 705 Astroloba pentagona var. spiralis, 705 Astroloba pentagona var. spirella, 705 Astroloba pentagona var. torulosa, 705 Astroloba robusta, 705 Astroloba rubriflora, 828 Astroloba rugosa, 703 Astroloba skinneri, 709 Astroloba smutsiana, 702 Astroloba spiralis, 704, 705 Astroloba spiralis cv. Pentagona, 705 Astroloba spiralis var. imbricata, 705 Astroloba spirella, 705 Astroloba tenax, 706 Astroloba tenax var. moltenoi, 706

Index Astroloba tenax var. tenax, 706 Astroloba turgida, 703 Astroworthia, 709 Astroworthia bicarinata, 709 Astroworthia bicarinata nvar. skinneri, 709 Astroworthia skinneri, 709 Avetraceae, 1199 Avonsera, 1282 Ayensua uaipanensis, 840 B Baeoterpe brevifolia, 1248 Bakerantha, 997 Bakerantha tillandsioides, 1030 Bakeria, 997 Bakeria tillandsioides, 1030 Ballya, 1171 Ballya zebrina, 1172 Barbacenia gracilis, 1330 Basilaea bifolia, 1274 Battandiera, 1231 Battandiera stapfii, 1236 Bauschia, 1171 Baxteriaceae, 1389 Bayerara, 711 Beaucarnea, 1317, 1319 Beaucarnea ameliae, 1324 Beaucarnea bigelovii, 1347 Beaucarnea compacta, 1320 Beaucarnea congesta, 1320 Beaucarnea glassiana, 1320 Beaucarnea glauca, 1326 Beaucarnea goldmanii, 1321 Beaucarnea gracilis, 1321 Beaucarnea guatemalensis, 1322 Beaucarnea hiriartiae, 1322–1323 Beaucarnea hookeri, 1323 Beaucarnea inermis, 1325 Beaucarnea longifolia, 1350 Beaucarnea oedipus, 1321 Beaucarnea olsonii, 1324 Beaucarnea parviflora, 1350 Beaucarnea petenensis, 1324 Beaucarnea pliabilis, 1324 Beaucarnea purpusii, 1325 Beaucarnea recurvata, 1325 Beaucarnea recurvata var. intermedia, 1325 Beaucarnea recurvata var. rubra, 1325 Beaucarnea recurvata var. stricta, 1326 Beaucarnea sanctomariana, 1326 Beaucarnea stricta, 1326 Beaucarnea texana, 1351 Beaucarnea tuberculata, 1325 Behnia, 11 Behniaceae, 11 Belosynapsis, 1179 Belosynapsis kewensis, 1179 Beryllis, 1281

1417 Beschorneria, 12, 14, 313 Beschorneria albiflora, 314 Beschorneria argyrophylla, 318 Beschorneria bracteata, 318 Beschorneria calcicola, 315 Beschorneria californica, 318 Beschorneria chiapensis, 314 Beschorneria cohniana, 316 Beschorneria decosteriana, 318 Beschorneria dekosteriana, 318 Beschorneria dubia, 318 Beschorneria floribunda, 337 Beschorneria galeottii, 314 Beschorneria glauca, 318 Beschorneria hidalgorupicola, 318 Beschorneria mexicana, 318 Beschorneria multiflora, 337 Beschorneria parmentieri, 337 Beschorneria pubescens, 317 Beschorneria pumila, 314 Beschorneria rigida, 315–316 Beschorneria roseana, 316 Beschorneria schlechtendalii, 314 Beschorneria septentrionalis, 316 Beschorneria superba, 318 Beschorneria toneliana, 316 Beschorneria tonelii, 316 Beschorneria tubiflora, 316–317 Beschorneria verlindeniana, 314 Beschorneria viridiflora, 318 Beschorneria wrightii, 317–318 Beschorneria yuccoides, 318, 319 Beschorneria yuccoides ssp. dekosteriana, 318–319 Beschorneria yuccoides ssp. yuccoides, 319–320 Billbergia, 859 Billbergia bicolor, 986 Billbergia brasiliensis, 860 Billbergia distichostachya, 853 Billbergia euphemiae, 861 Billbergia euphemiae var. nudiflora, 861 Billbergia euphemiae var. purpurea, 861 Billbergia euphemiae var. saundersioides, 861 Billbergia ianthina, 860 Billbergia joinvillei, 843 Billbergia kuhlmannii, 860 Billbergia lansbergiana, 843 Billbergia legrelliana, 856 Billbergia legrelloe, 856 Billbergia leopoldii, 860 Billbergia nudicaulis, 854 Billbergia nuptialis, 860 Billbergia nutans, 861 Billbergia nutans fa. rupestris, 861 Billbergia nutans var. schimperiana, 861 Billbergia nutans var. striata, 861 Billbergia polystachya, 853 Billbergia schimperiana, 861 Billbergia variegata, 1036 Billmea, 860

1418 Biltanthus, 860, 863 Blephanthera, 713 Blephanthera depressa, 717 Blephanthera hookeri, 717 Bonapartea, 21, 1107 Bonapartea filamentosa, 109 Bonapartea flagelliformis, 116 Bonapartea glauca, 1330 Bonapartea gracilis, 1330 Bonapartea hystrix, 246 Bonapartea juncea, 116 Bonapartea stricta, 247 Boophane, 429 Boophone, 429 Boophone ciliaris, 430 Boophone disticha, 429–430, 430 Boophone ernesti-ruschii, 430 Boophone haemanthoides, 429–430, 430 Boophone intermedia, 430 Boophone longipedicellata, 430 Boophone toxicaria, 429 Boosia, 1250 Borderea, 1201 Botryosicyos, 1201 Bowiea, 486, 1243 Bowiea africana, 515 Bowiea gariepensis, 1244 Bowiea kilimandscharica, 1244 Bowiea myriacantha, 618 Bowiea volubilis, 1244 Bowiea volubilis ssp. gariepensis, 1244 Brachyscypha, 1265 Branciona, 1231 Bravoa, 21 Bravoa bulliana, 75 Bravoa coetocapnia, 86 Bravoa densiflora, 91 Bravoa geminiflora, 86 Bravoa graminiflora, 86 Bravoa graminifolia, 124 Bravoa kewensis, 180 Bravoa platyphylla, 210 Bravoa sessiliflora, 55 Bravoa singuliflora, 239 Brizophile, 1281 Brocchinia, 840 Brocchinia uaipanensis, 840 Bromelia, 840 Bromelia albobracteata, 986 Bromelia antiacantha, 840, 998 Bromelia balansae, 840 Bromelia bicolor, 986 Bromelia carnea, 1042 Bromelia comata, 853 Bromelia desmetiana, 1025 Bromelia joinvillei, 986 Bromelia lindleyana, 1042 Bromelia linifera, 1036 Bromelia longifolia, 1042

Index Bromelia nudicaulis, 854 Bromelia pinguin, 840 Bromelia pitcairniifolia, 1043 Bromelia urbaniana, 875 Bromelia variegata, 1036 Bromeliaceae, 835 Brunsvigia, 433 Brunsvigia ciliaris, 429 Brunsvigia disticha, 429 Brunsvigia herrei, 433 Brunsvigia radula, 433 Brunsvigia rautanenii, 430 Brunsvigia toxicaria, 429 Brunsvigiaceae, 425 Bulbine, 713 Bulbine abyssinica, 714–715 Bulbine acaulis, 715 Bulbine adnutans, 714 Bulbine alata, 715 Bulbine alba, 723 Bulbine alooides, 715 Bulbine altissima, 716 Bulbine alveolata, 715 Bulbine angustifolia, 715–716 Bulbine annua, 716 Bulbine asphodeloides, 716 Bulbine asphodeloides var. denticulifera, 725 Bulbine asphodeloides var. filifoliodes, 714 Bulbine asphodeloides var. monticola, 714 Bulbine asphodeloides var. otaviensis, 717 Bulbine australis, 717 Bulbine bachmanniana, 730 Bulbine bachmannii, 730 Bulbine bisulcata, 718 Bulbine breviracemosa, 721 Bulbine brunsvigiifolia, 716 Bulbine bruynsii, 717 Bulbine bulbosa, 717 Bulbine caespitosa, 725 Bulbine canaliculata, 720, 830 Bulbine capensis, 717 Bulbine capitata, 717–718 Bulbine caput-medusae, 718 Bulbine cataphyllata, 718 Bulbine caulescens, 723 Bulbine caulescens var. incurva, 723 Bulbine cepacea, 718 Bulbine ciliata, 830 Bulbine circinata, 736 Bulbine coetzeei, 718 Bulbine concinna, 729 Bulbine crassa, 718 Bulbine cremnophila, 719 Bulbine crocea, 716 Bulbine curvifolia, 723 Bulbine dactylopsoides, 719 Bulbine decurvata, 714 Bulbine densiflora, 730 Bulbine dewetii, 719–720

Index Bulbine dielsii, 716 Bulbine diphylla, 720 Bulbine dissimilis, 720 Bulbine dubia, 721 Bulbine elongata, 714 Bulbine ensifolia, 725 Bulbine erectipilosa, 720 Bulbine erumpens, 720 Bulbine esterhuyseniae, 721 Bulbine falcata, 831 Bulbine fallax, 721 Bulbine favosa, 721 Bulbine filifolia, 721 Bulbine fistulosa, 721–722 Bulbine flexicaulis, 722 Bulbine flexuosa, 722 Bulbine floribunda, 734 Bulbine foleyi, 722 Bulbine fragilis, 722 Bulbine francescae, 722 Bulbine fraseri, 717 Bulbine frutescens, 723 Bulbine frutescens var. incurva, 723 Bulbine frutescens var. rostrata, 723 Bulbine glauca, 723 Bulbine graminea, 725 Bulbine hallii, 724 Bulbine hamata, 714 Bulbine hantamensis, 735 Bulbine haworthioides, 724 Bulbine hookeri, 723 Bulbine huilensis, 714 Bulbine inaei, 724 Bulbine inamarxiae, 724 Bulbine incurva, 723 Bulbine inexpectata, 718 Bulbine inflata, 725 Bulbine inops, 722 Bulbine lagopus, 725 Bulbine lamprophylla, 725 Bulbine latibracteata, 730 Bulbine latifolia, 725 Bulbine latifolia var. curvata, 725–726 Bulbine latifolia var. latifolia, 726 Bulbine latitepala, 714 Bulbine lavrani, 726 Bulbine laxiflora, 731 Bulbine lolita, 726–727 Bulbine longifolia, 727 Bulbine longiscapa, 716 Bulbine louwii, 727 Bulbine lydenburgensis, 717 Bulbine macrocarpa, 729 Bulbine macrophylla, 715 Bulbine mallyana, 736 Bulbine margarethae, 727 Bulbine mayorii, 721 Bulbine meiringii, 727–728 Bulbine melanovaginata, 728

1419 Bulbine mesembryanthemoides, 728 Bulbine mesembryanthoides, 728 Bulbine mesembryanthoides ssp. mesembryanthoides, 728 Bulbine mesembryanthoides ssp. namaquensis, 728 Bulbine mettinghii, 716 Bulbine migiurtina, 729 Bulbine minima, 729 Bulbine monophylla, 729 Bulbine muscicola, 729 Bulbine namaensis, 729 Bulbine narcissifolia, 730 Bulbine natalensis, 730 Bulbine navicularifolia, 730 Bulbine nigra, 714 Bulbine nutans, 730 Bulbine ophiophylla, 731 Bulbine orchioides, 728 Bulbine otaviensis, 717 Bulbine pallida, 716 Bulbine parviflora, 718 Bulbine patersoniae, 714 Bulbine pendens, 731 Bulbine pendula, 731 Bulbine platyphylla, 715 Bulbine praemorsa, 731–732 Bulbine pugioniformis, 718 Bulbine pusilla, 732 Bulbine quartzicola, 732 Bulbine ramosa, 732 Bulbine retinens, 732–733 Bulbine rhopalophylla, 733 Bulbine rigidula, 721 Bulbine rostrata, 723 Bulbine rupicola, 733 Bulbine sedifolia, 733 Bulbine semenaliundata, 733–734 Bulbine semibarbata, 734 Bulbine semibarbata fa. gracilescens, 734 Bulbine semibarbata var. depilata, 715 Bulbine setifera, 721 Bulbine spongiosa, 734 Bulbine stenophylla, 717 Bulbine stolonifera, 734 Bulbine striata, 734–735 Bulbine suavis, 723 Bulbine subbarbata, 717 Bulbine succulenta, 735 Bulbine suurbergensis, 735 Bulbine tecta, 735 Bulbine tecta ssp. tecta, 736 Bulbine tenuifolia, 741 Bulbine tetraphylla, 731 Bulbine thomasiae, 736 Bulbine torsiva, 736 Bulbine torta, 736 Bulbine tortifolia, 716 Bulbine transvaalensis, 725 Bulbine trichophylla, 721 Bulbine triebneri, 723

1420 Bulbine truncata, 736 Bulbine tuberosa, 718 Bulbine undulata, 733 Bulbine urgineoides, 731 Bulbine vagans, 737 Bulbine vesicularis, 729 Bulbine vitrea, 737 Bulbine vittatifolia, 737 Bulbine wiesei, 738 Bulbine witputzii, 714 Bulbine xanthobotrys, 714 Bulbine zeyheri, 731 Bulbinella, 478, 714, 741 Bulbinella calcicola, 741 Bulbinopsis, 713 Bulbinopsis alata, 715 Bulbinopsis bulbosa, 717 Bulbinopsis semibarbata, 734 Bulbinopsis semibarbata var. depilata, 715 Bulbinopsis terrae-victoriae, 723 Buonapartea, 1107 Buphane, 429 Buphane fischeri, 1364 Buphone, 429 Busipho, 486 C Cabbage tree, 1296 Caesia comosa, 452 Caladiaceae, 457 Caladium zamiifolium, 462 Calectasiaceae, 1389 Calibanus, 1319 Calibanus caespitosus, 1323 Calibanus glassianus, 1320, 1320–1321 Calibanus hookeri, 1323 Callaceae, 457 Callisia, 1173 Callisia elegans, 1174 Callisia fragrans, 1174 Callisia fragrans cv. Melnickoff, 1174 Callisia gentlei, 1174, 1176 Callisia gentlei var. elegans, 1174 Callisia glandulosa, 1190 Callisia grandiflora, 1190 Callisia ionantha, 1190 Callisia martensiana, 1175 Callisia micrantha, 1174 Callisia multiflora, 1174–1175 Callisia navicularis, 1175 Callisia repens, 1175–1176 Callisia repens var. ciliata, 1176 Callisia repens var. mandonii, 1176 Callisia tehuantepecana, 1176 Callisia tripogandra, 1191 Callisia warszewicziana, 1176–1177 Calodracon, 1295 Camassia, 13

Index Campelia, 1183 Caraguata, 1107 Cartonemataceae, 1167 Caruelia, 1281 Catevala, 769 Catevala arachnoidea, 774 Catevala arborescens, 503 Catevala atroviridis, 789 Catevala humilis, 577 Catevala retusa, 807 Cathissa, 1281 Catopsis, 840 Catopsis delicatula, 840 Catopsis juncifolia, 840 Catopsis morreniana, 840 Catopsis penduliflora, 991 Celsia, 1281 Chamaealoe, 486 Chamaealoe africana, 515 Chamaeloba, 697 Chamaeteria, 745 Charlwoodia, 1295 Charybdis, 1250 Charybdis aphylla, 1259 Charybdis hesperia, 1259 Charybdis maritima, 1259 Charybdis maura, 1259 Charybdis numidica, 1259 Charybdis pancration, 1259 Charybdis tazensis, 1260 Charybdis undulata, 1259 Chevaliera, 851 Chevaliera grandiceps, 854 Chevalieria, 851 Chloriza, 1265 Chlorogalaceae, 9 Chlorogalum, 10, 13 Chlorophytum beniense, 452 Chlorophytum brevipes, 452 Chlorophytum bukobense, 452 Chlorophytum bukobense var. kilimandscharicum, 452 Chlorophytum bulbinifolium, 452 Chlorophytum burchellii, 452 Chlorophytum cremnophilum, 453 Chlorophytum delagoense, 452 Chlorophytum drepanophyllum, 831 Chlorophytum elatulum, 452 Chlorophytum elatum var. burchelii, 452 Chlorophytum elgonense, 452 Chlorophytum gazense, 452 Chlorophytum glaucidulum, 452 Chlorophytum glaucidulum var. pauper, 452 Chlorophytum inopinum, 452 Chlorophytum ituriense, 452 Chlorophytum kirkii, 452 Chlorophytum limurense, 452 Chlorophytum longum, 452 Chlorophytum macrophyllum var. angustifolium, 452 Chlorophytum magnum, 452

Index Chlorophytum miserum, 452 Chlorophytum nemorosum, 452 Chlorophytum paludicola, 452 Chlorophytum polyphyllum, 453 Chlorophytum ramiferum, 452 Chlorophytum rhizomatosum, 453 Chlorophytum rhizopendulum, 453 Chlorophytum rugosum, 452 Chlorophytum sparsiflorum, 452 Chlorophytum sternbergianum, 452 Chlorophytum suffruticosum, 451–453 Chlorophytum turritum, 452 Chlorophytum usambarense, 452 Chortolirion, 486 Chortolirion angolense, 681 Chortolirion bergerianum, 513 Chortolirion latifolium, 583 Chortolirion stenophyllum, 513 Chortolirion subspicatum, 662 Chortolirion tenuifolium, 513 Chrysodracon, 1338 Clistoyucca, 363 Clistoyucca arborescens, 378 Clistoyucca brevifolia, 378 Clivia, 426 Clivia mirabilis, 426 Coburgia radula, 434 Codonocrinum, 323 Codonocrinum agavoides, 325 Coelanthus, 1265 Coetocapnia, 21 Coetocapnia geminiflora, 86 Cohnia, 1295 Coilonox, 1231 Coilonox diluculum, 1236 Coilonox scabrocostatum, 1239 Coilonox unifoliatum, 1240 Colocasiaceae, 457 Commelina floribunda, 1191 Commelina multiflora, 1175 Commelina speciosa, 1180 Commelinaceae, 1167–1169 Convallaria racemosa, 1364 Convallariaceae, 1315, 1316 Cordyline, 370, 1293, 1295, 1317, 1353 Cordyline australis, 1295 Cordyline parviflora, 1350 Cordyline roxburghiana, 1377 Cordyline rumphii, 1338 Cordyline vivipara, 452 Cottendorfia albicans, 990 Cottendorfia weddelliana, 993 Cremobotrys, 859 Crinum angustifolium, 48 Crinum obliquum, 438 Cryptanthopsis, 1051 Cryptanthopsis navioides, 1104 Cryptanthus, 865, 1053, 1102 Cryptanthus acaulis var. bromelioides, 868

1421 Cryptanthus acaulis var. diversifolius, 868 Cryptanthus bahianus, 867 Cryptanthus bahiensis, 867 Cryptanthus boanovensis, 867 Cryptanthus bromelioides, 868 Cryptanthus bromelioides var. tricolor, 868 Cryptanthus caracensis, 868 Cryptanthus carnosus, 868 Cryptanthus crassifolius, 868 Cryptanthus diversifolius, 868 Cryptanthus fosterianus, 868 Cryptanthus glaziovii, 867, 869 Cryptanthus incrassatus, 869 Cryptanthus leopoldo-horstii, 869 Cryptanthus microglazioui, 869, 1067 Cryptanthus micrus, 870 Cryptanthus praetextus, 870 Cryptanthus schwackeanus, 870 Cryptanthus seidelianus, 871 Cryptanthus sergipensis, 871 Cryptanthus suaveolens, 868 Cryptanthus warasii, 871 Cryptanthus warren-loosei, 871 Cryptbergia, 860, 863 Cryptbergia meadii, 863 Cryptbergia rubra, 863 Cryptocorynaceae, 457 Cryptostephanus herrei, 436 Cummingara, 743 Cuthbertia, 1173 Cyanotis, 1179 Cyanotis gryphaea, 1181 Cyanotis kewensis, 1179 Cyanotis kewoides, 1179 Cyanotis nodiflora, 1181 Cyanotis nodiflora var. madagascarica, 1180 Cyanotis somaliensis, 1180 Cyanotis speciosa, 1180 Cyanotis speciosa ssp. bulbosa, 1180 Cyanotis speciosa ssp. madagascarica, 1180 Cyanotis speciosa ssp. speciosa, 1180 Cymbispatha, 1183 Cyperaceae, 1195 Cyperus, 1195 Cyperus bulbosus, 1195 Cyperus esculentus, 1195 Cyperus esculentus var. sativus, 1195 Cyperus rotundus, 1195 Cypripediaceae, 1301 Cyrtanthaceae, 425 Cyrtanthus, 435 Cyrtanthus falcatus, 435–436 Cyrtanthus flammosus, 436 Cyrtanthus flanaganii, 436 Cyrtanthus herrei, 436–437 Cyrtanthus inaequalis, 437 Cyrtanthus junodii, 437 Cyrtanthus labiatus, 437 Cyrtanthus montanus, 438

1422 Cyrtanthus obliquus, 438 Cyrtanthus salmonoides, 438 Cyrtanthus sanguineus, 438 Cyrtanthus sanguineus ssp. ballyi, 438 Cyrtanthus sanguineus ssp. minor, 438 Cyrtanthus sanguineus ssp. salmonoides, 438 Cyrtanthus sanguineus ssp. sanguineus, 439 Cyrtanthus sanguineus ssp. wakefieldii, 439 Cyrtanthus sanguineus var. wakefieldii, 439 D Danthonia pumila, 1311 Dasylirion, 1329 Dasylirion acrotrichum, 1330 Dasylirion bigelovii, 1347 Dasylirion caespitosum, 1323 Dasylirion durangense, 1333 Dasylirion flexile, 1323 Dasylirion glaucophyllum, 1330 Dasylirion glaucum, 1330 Dasylirion gracile, 1321, 1330 Dasylirion graminifolium, 1330 Dasylirion hartwegianum, 1323 Dasylirion hookeri, 1323 Dasylirion humboldtii, 1350 Dasylirion inerme, 1325 Dasylirion juncifolium, 1331 Dasylirion laxiflorum, 1332 Dasylirion leiophyllum, 1331 Dasylirion longifolium, 1350 Dasylirion longissimum, 1332 Dasylirion pitcairniifolium, 1006 Dasylirion pliabile, 1324 Dasylirion quadrangulatum, 1332–1333 Dasylirion recurvatum, 1325 Dasylirion robustum, 1330 Dasylirion serratifolium, 1332 Dasylirion stewartii, 1331 Dasylirion strictum, 1326 Dasylirion wheeleri, 1333 Dasylirion wheeleri [?] wislizeni, 1333 Dasylirion wheeleri var. durangense, 1333 Dasylirion wheeleri var. wislizeni, 1333 Dasylirium, 1329 Dasypogonaceae, 1389 Dasystachys polyphylla, 453 Deinacanthon, 875 Deinacanthon urbanianum, 875 Delpinoa, 22 Delpinoa gracillima, 213, 214 Dendrobium, 1300, 1301 Dendropogon, 1107 Descantaria, 1189 Descantaria cumanensis, 1191 Descantaria glandulosa, 1190 Descantaria ionantha, 1190 Descantaria multiflora, 1190 Descantaria pflanzii, 1190

Index Descantaria procumbens, 1191 Descantaria radiata, 1190 Desertia, 1273 Desertia etesionamibensis, 1275 Desertia luteovirens, 1273 Deuterocairnia, 877 Deuterocohnia, 879 Deuterocohnia abstrusa, 880 Deuterocohnia bracteosa, 888 Deuterocohnia brevifolia, 877, 880–881, 1045 Deuterocohnia brevispicata, 881–882, 891 Deuterocohnia carmineoviridiflora, 885 Deuterocohnia chrysantha, 882 Deuterocohnia digitata, 882–883 Deuterocohnia divaricata, 885 Deuterocohnia gableana, 883 Deuterocohnia glandulosa, 883 Deuterocohnia haumanii, 883 Deuterocohnia longipetala, 884 Deuterocohnia longipetala fa. uberrima, 884 Deuterocohnia lorentziana, 881, 891, 1045 Deuterocohnia lotteae, 884 Deuterocohnia meziana, 884, 891 Deuterocohnia meziana ssp. carmineoviridiflora, 885 Deuterocohnia meziana ssp. meziana, 885–886 Deuterocohnia meziana ssp. pedicellata, 886 Deuterocohnia meziana ssp. vogtii, 886 Deuterocohnia meziana var. carmineoviridiflora, 885 Deuterocohnia paraguariensis, 885 Deuterocohnia pedicellata, 886 Deuterocohnia recurvipetala, 886 Deuterocohnia sanctae-crucis, 886–887 Deuterocohnia scapigera, 887 Deuterocohnia scapigera ssp. sanctae-crucis, 886 Deuterocohnia schreiteri, 887, 891, 995, 1071 Deuterocohnia seramisiana, 887 Deuterocohnia strobilifera, 888 Deuterocohnia strobilifera var. inermis, 888 Deuterocohnia strobilifera var. strobilifera, 888 Diacles, 441 Diaphoranthema, 1107 Diaphoranthema capillaris, 1116 Diaphoranthema virescens, 1159 Dichorisandra warszewicziana, 1176 Dilanthes, 829 Dioscorea, 1199, 1201 Dioscorea astrostigma, 1205 Dioscorea axilliflora, 1204 Dioscorea basiclavicaulis, 1202–1203 Dioscorea billbergiana, 1205 Dioscorea brevipes, 1206 Dioscorea composita, 1205 Dioscorea cylindrostachya, 1204 Dioscorea deamii, 1205 Dioscorea deppei, 1205 Dioscorea elephantipes, 1203–1204 Dioscorea elephantopus, 1204 Dioscorea fastigiata, 1204–1205 Dioscorea gayi, 1204

Index Dioscorea geissei, 1204 Dioscorea hederifolia, 1207 Dioscorea hemicrypta, 1205 Dioscorea leiboldiana, 1205 Dioscorea macrophylla, 1205 Dioscorea macrostachya, 1205 Dioscorea macrostachya var. spiculiflora, 1205 Dioscorea marlothii, 1206 Dioscorea mexicana, 1205 Dioscorea mexicana var. sessiliflora, 1205 Dioscorea montana, 1206 Dioscorea montana var. duemmeri, 1207 Dioscorea montana var. glauca, 1207 Dioscorea montana var. lobata, 1206 Dioscorea montana var. paniculata, 1207 Dioscorea montana var. sagittata, 1206 Dioscorea palmeri, 1205 Dioscorea paniculata, 1207 Dioscorea paupera, 1204 Dioscorea rehmannii, 1207 Dioscorea silvatica, 1206 Dioscorea strydomiana, 1205–1206 Dioscorea sylvatica, 1206 Dioscorea sylvatica ssp. lydenbergensis, 1206 Dioscorea sylvatica var. brevipes, 1206 Dioscorea sylvatica var. multiflora, 1206 Dioscorea sylvatica var. paniculata, 1206–1207 Dioscorea sylvatica var. rehmannii, 1207 Dioscorea sylvatica var. sylvatica, 1207 Dioscorea thinophila, 1204 Dioscorea tuerckheimii, 1205 Dioscoreaceae, 1199 Dipcadi, 1247 Dipcadi brevifolium, 1248 Dipcadi hyacinthoides, 1248 Dipcadi spirale, 1248 Disgrega, 1189 Disquamia, 851 Disteganthus, 840 Donnellia, 1189 Donnellia grandiflora, 1190 Doryanthaceae, 1211 Doryanthes, 1213 Doryanthes excelsa, 1213 Doryanthes excelsa var. guilfoylei, 1213 Doryanthes excelsa var. palmeri, 1213 Doryanthes guilfoylei, 1213 Doryanthes larkinii, 1213 Doryanthes palmeri, 1213 Doryanthes palmeri var. larkinii, 1213 Dracaena, 1337, 1354 Dracaena aethiopica, 1355 Dracaena aletriformis, 1338 Dracaena angolensis, 1360 Dracaena arborescens, 1356 Dracaena ascendens, 1356 Dracaena aubrytiana, 1356 Dracaena australis, 1296 Dracaena bacularis, 1357

1423 Dracaena bagamoyensis, 1357 Dracaena ballyi, 1357 Dracaena bella, 1358 Dracaena boscii, 116 Dracaena bugandana, 1369 Dracaena burdettii, 1358 Dracaena burmanica, 1359 Dracaena canaliculata, 1359 Dracaena caulescens, 1359 Dracaena cinnabari, 1338–1339 Dracaena conspicua, 1360 Dracaena dawei, 1361 Dracaena dentata, 642 Dracaena dooneri, 1361 Dracaena downsii, 1361 Dracaena draco, 1339 Dracaena draco ssp. ajgal, 1339 Dracaena draco ssp. caboverdeana, 1339 Dracaena draco ssp. draco, 1340 Dracaena dumetescens, 1362 Dracaena ebracteata, 1362 Dracaena ehrenbergii, 394 Dracaena eilensis, 1362 Dracaena ellenbeckiana, 1340 Dracaena ellenbeckii, 1341 Dracaena ensifolia, 393, 394 Dracaena erecta, 471 Dracaena erythraeae, 1363 Dracaena fasciata, 1364 Dracaena filamentosa, 116 Dracaena fintelmannii, 394 Dracaena forskaoliana, 1364 Dracaena francisii, 1365 Dracaena frequens, 1365 Dracaena gracillima, 1365 Dracaena hallii, 1366 Dracaena hargeisana, 1366 Dracaena hookeriana, 1338 Dracaena humiflora, 1366 Dracaena hyacinthoides, 1367 Dracaena itumei, 1367 Dracaena jayniana, 1341 Dracaena latifolia, 1338 Dracaena lenneana, 372, 394 Dracaena lennei, 394 Dracaena liberica, 1369 Dracaena longiflora, 1370 Dracaena longiflora var. fernandopoensis, 1370 Dracaena longistyla, 1370 Dracaena lunatifolia, 1370 Dracaena malawiana, 1364 Dracaena marginata, 642 Dracaena masoniana, 1370 Dracaena newtoniana, 1371 Dracaena nilotica, 1371 Dracaena nitida, 1372 Dracaena ombet, 1341 Dracaena ombet ssp. ombet, 1341 Dracaena ombet ssp. schizantha, 1341

1424 Dracaena parva, 1372 Dracaena parviflora, 1350 Dracaena patens, 1372 Dracaena pearsonii, 1373 Dracaena pedicellata, 1373 Dracaena perrotii, 1373 Dracaena petenensis, 1324 Dracaena pethera, 1368 Dracaena pethera var. pulchra, 1368 Dracaena pfisteri, 1374 Dracaena phillipsiae, 1374 Dracaena pinguicula, 1374 Dracaena pinguicula ssp. disticha, 1374 Dracaena pinguicula ssp. nana, 1374 Dracaena powellii, 1375 Dracaena powysii, 1376 Dracaena raffillii, 1376 Dracaena rhabdophylla, 1342 Dracaena roxburghiana, 1377 Dracaena rumphii, 1338 Dracaena sambiranensis, 1377 Dracaena schizantha, 1342 Dracaena scimitariformis, 1378 Dracaena senegambica, 1378 Dracaena serpenta, 1365 Dracaena serpenta var. humbertiana, 1381 Dracaena serrulata, 1342 Dracaena serrulata ssp. dhofarica, 1342 Dracaena serrulata ssp. mccoyorum, 1342 Dracaena serrulata ssp. serrulata, 1343 Dracaena singularis, 1364 Dracaena sinus-simiorum, 1378 Dracaena sordida, 1379 Dracaena spathulata, 1360 Dracaena striata, 471 Dracaena stricta, 471 Dracaena stuckyi, 1379 Dracaena subspicata, 1380 Dracaena subtilis, 1380 Dracaena suffruticosa, 1380 Dracaena suffruticosa var. longituba, 1380 Dracaena tamaranae, 1343 Dracaena tayfunii, 1337 Dracaena testudinea, 1358 Dracaena transvaalensis, 1338 Dracaena trifasciata, 1381 Dracaena undulata, 471 Dracaena varians, 1381 Dracaena volkensii, 1381 Dracaena yuccoides, 394 Dracaena zebra, 1371 Dracaena zebra var. nyasica, 1371 Dracaena zeylanica, 1382 Dracaenaceae, 1315 Dracaenoideae, 1316–1317 Draco hookeriana, 1338 Draco ombet, 1341 Dracontiaceae, 457 Dregeochloa, 1308, 1311

Index Dregeochloa pumila, 1311 Drimia, 1249 Drimia acarophylla, 1251 Drimia albiflora, 1251 Drimia altissima, 1251 Drimia anomala, 1252 Drimia arenicola, 1252 Drimia barteri, 1251 Drimia chalumnensis, 1253 Drimia cochlearis, 1253 Drimia convallarioides, 1253 Drimia cooperi, 1270 Drimia cremnophila, 1254 Drimia cryptopoda, 1254 Drimia cuscutoides, 1257 Drimia cyanelloides, 1254 Drimia delagoensis, 1254 Drimia edwardsii, 1255 Drimia ensifolia, 1270 Drimia fasciata, 1255 Drimia flagellaris, 1255 Drimia fugax, 1259 Drimia gigantea, 1251 Drimia haworthioides, 1256 Drimia incerta, 1252 Drimia intricata, 1256 Drimia intricata var. intricata, 1256 Drimia intricata var. visagieae, 1258 Drimia karooica, 1258 Drimia loedolffiae, 1259 Drimia ludwigii, 1270 Drimia maritima, 1259 Drimia maritima ssp. maura, 1259 Drimia media, 1260 Drimia multifolia, 1260 Drimia mzimvubuensis, 1261 Drimia namibensis, 1261 Drimia paolii, 1252 Drimia platyphylla, 1261 Drimia rotunda, 1262 Drimia secunda, 1262 Drimia sigmoidea, 1262 Drimia uitenhagensis, 1251 Drimia undata, 1260 Drimia undata ssp. caeculi, 1259 Drimia uniflora, 1263 Drimiopsis, 1269 Drimya, 1249 Duthiea, 1250 Dyckcohnia, 891 Dyckia, 893 Dyckia acrosoma, 895 Dyckia acutiflora, 895–896 Dyckia affinis, 896 Dyckia agudensis, 896 Dyckia alba, 896 Dyckia altissima, 895 Dyckia apensis, 926 Dyckia areniticola, 897

Index Dyckia argentea, 932, 1007 Dyckia atratiflora, 898 Dyckia aurea, 898 Dyckia barthlottii, 898–899 Dyckia beateae, 899 Dyckia biflora, 967 Dyckia boliviensis, 926 Dyckia brachyphylla, 899 Dyckia brachystachya, 899 Dyckia bracteata, 900 Dyckia brasiliana, 900 Dyckia braunii, 900 Dyckia brevifolia, 901 Dyckia burchellii, 901 Dyckia burle-marxii, 901 Dyckia cabrerae, 902 Dyckia cangaphila, 902 Dyckia catharinensis, 911 Dyckia catharinensis var. dentata, 911 Dyckia chaguar, 915 Dyckia choristaminea, 902–903 Dyckia cinerea, 904 Dyckia coccinea, 956 Dyckia coccinea var. deltoidea, 906 Dyckia commixta, 903 Dyckia conceicionensis, 904 Dyckia consimilis, 904 Dyckia conspicua, 926 Dyckia coximensis, 904–905 Dyckia crassifolia, 905 Dyckia crocea, 905 Dyckia dawsonii, 905 Dyckia decomposita, 884 Dyckia delicata, 905–906 Dyckia deltoidea, 906 Dyckia densiflora, 907 Dyckia desmetiana, 1025 Dyckia dissitiflora, 907 Dyckia dissitiflora var. bracteata, 900 Dyckia distachya, 907 Dyckia distachya fa. inducta, 907 Dyckia divaricata, 907–908 Dyckia domfelicianensis, 908–909 Dyckia duarteana, 952 Dyckia duckei, 909 Dyckia dusenii, 909 Dyckia edwardii, 909–910 Dyckia elata, 910 Dyckia elisabethae, 910 Dyckia elongata, 910 Dyckia eminens, 911 Dyckia encholirioides, 911 Dyckia encholirioides var. encholirioides, 911 Dyckia encholirioides var. rubra, 911 Dyckia espiritosantensis, 911–912 Dyckia estevesii, 891, 912–913 Dyckia excelsa, 913 Dyckia exserta, 913

1425 Dyckia ferox, 914 Dyckia ferox fa. australis, 914 Dyckia ferox fa. hamosa, 914 Dyckia ferox fa. vulgaris, 914 Dyckia ferox ssp. hamosa, 914 Dyckia ferrisincola, 914 Dyckia ferruginea, 914–915 Dyckia floribunda, 915 Dyckia formosensis, 915 Dyckia fosteriana, 891, 916, 1069 Dyckia fosteriana var. fosteriana, 916 Dyckia fosteriana var. robustior, 916 Dyckia frigida, 916 Dyckia gemellaria, 901 Dyckia gigantea, 943 Dyckia gilliesii, 915 Dyckia glabrifolia, 916–917 Dyckia glandulosa, 917 Dyckia glaziovii, 1036 Dyckia goehringii, 917–918 Dyckia goiana, 918 Dyckia gouveiana, 918–919 Dyckia gracilis, 919 Dyckia grandidentata, 919 Dyckia grandiflora, 923 Dyckia grandifolia, 950 Dyckia granmogulensis, 920 Dyckia grisebachii, 880 Dyckia hamosa, 914 Dyckia hassleri, 926 Dyckia hassleri fa. gracilis, 926 Dyckia hassleri ssp. basispina, 926 Dyckia hassleri var. arenosa, 926 Dyckia hassleri var. montana, 926 Dyckia hassleri var. typica, 926 Dyckia hatschbachii, 920 Dyckia hebdingii, 920 Dyckia heloisae, 971 Dyckia hilaireana, 949 Dyckia hohenbergioides, 920 Dyckia horridula, 921 Dyckia ibicuiensis, 921 Dyckia ibiramensis, 922 Dyckia incana, 922–923 Dyckia inflexifolia, 923 Dyckia insignis, 923–924 Dyckia insignis [?] flaviflora, 923 Dyckia insignis var. macrantha, 923 Dyckia insignis var. obtusiflora, 923 Dyckia interrupta, 907 Dyckia irmgardiae, 924 Dyckia irwinii, 924 Dyckia joanae-marcioi, 924–925 Dyckia jonesiana, 925 Dyckia julianae, 925 Dyckia kranziana, 925 Dyckia kuntzeana, 955 Dyckia lagoensis, 926 Dyckia laxiflora, 895

1426 Dyckia lemaireana, 895 Dyckia leptostachia, 926 Dyckia leptostachya, 926–927 Dyckia limae, 927 Dyckia lindaiana, 895 Dyckia lindevaldae, 927 Dyckia linearifolia, 927 Dyckia longifolia, 926 Dyckia longipetala, 884 Dyckia lunaris, 928 Dyckia lutziana, 928 Dyckia macedoi, 891, 928–929 Dyckia machrisiana, 929 Dyckia macracantha, 950 Dyckia macropoda, 929 Dyckia maracasensis, 929 Dyckia maritima, 929 Dyckia marnier-lapostollei, 891, 930 Dyckia marnier-lapostollei var. estevesii, 930 Dyckia marnier-lapostollei var. marnier-lapostollei, 930 Dyckia martinellii, 931–932 Dyckia mauriziae, 931–932 Dyckia mello-barretoi, 932 Dyckia meziana, 914 Dyckia mezii, 932 Dyckia micracantha, 956 Dyckia microcalyx, 932 Dyckia microcalyx var. inermis, 933 Dyckia microcalyx var. micrantha, 933 Dyckia microcalyx var. microcalyx, 933 Dyckia microcalyx var. ostenii, 933 Dyckia milagrensis, 933 Dyckia minarum, 933 Dyckia minutiflora, 933 Dyckia mirandana, 934–935 Dyckia missionum, 937 Dyckia missionum var. breviflora, 937 Dyckia mitis, 935 Dyckia montevidensis, 946 Dyckia montevidensis var. tandilensis, 947 Dyckia montezumensis, 935 Dyckia monticola, 935–936 Dyckia morreniana, 955 Dyckia myriostachya, 950 Dyckia nana, 936 Dyckia nervata, 936 Dyckia niederleinii, 937 Dyckia nigrospinulata, 937 Dyckia nobilis, 937–938 Dyckia odorata, 938 Dyckia oligantha, 949 Dyckia oligantha var. cristallina, 949 Dyckia orobanchoides, 938 Dyckia paraensis, 938 Dyckia pauciflora, 939 Dyckia paucispina, 939 Dyckia pectinata, 939–940 Dyckia pedicellata, 975

Index Dyckia pernambucana, 940 Dyckia piauiensis, 940 Dyckia piracanjubensis, 941 Dyckia platyphylla, 941, 1069 Dyckia polyclada, 942 Dyckia pontesii, 942 Dyckia pottiorum, 942–943 Dyckia princeps, 943 Dyckia pseudococcinea, 943–944 Dyckia pulquinensis, 944 Dyckia pumila, 944 Dyckia racemosa, 944 Dyckia racinae, 944 Dyckia ragonesei, 945 Dyckia ramosa, 895 Dyckia rariflora, 945 Dyckia rariflora var. cunninghamii, 947 Dyckia rariflora var. montevidensis, 946 Dyckia rariflora var. remotiflora, 946 Dyckia rauhii, 930 Dyckia regalis, 916 Dyckia reitzii, 891, 945 Dyckia remotiflora, 946 Dyckia remotiflora var. angustior, 946 Dyckia remotiflora var. montevidensis, 946 Dyckia remotiflora var. remotiflora, 946 Dyckia remotiflora var. tandilensis, 947 Dyckia retardata, 947 Dyckia retroflexa, 947 Dyckia richardii, 947 Dyckia rigida, 948 Dyckia rojasii, 926 Dyckia rondonopolitana, 948 Dyckia rubra, 911 Dyckia rupestris, 940 Dyckia saxatilis, 948–949 Dyckia schreiteri, 895 Dyckia schwackeana, 949 Dyckia scrutor, 978 Dyckia secunda, 949 Dyckia secundifolia, 949 Dyckia selloa, 950–951 Dyckia sellowiana, 951 Dyckia sickii, 951 Dyckia silvae, 951 Dyckia simulans, 951–952 Dyckia sordida, 952 Dyckia spectabilis, 978 Dyckia spinulosa, 952 Dyckia stenophylla, 952 Dyckia stolonifera, 953 Dyckia strehliana, 953 Dyckia subinermis, 953 Dyckia subsecunda, 979 Dyckia sulcata, 954 Dyckia sulphurea, 901 Dyckia tenebrosa, 954 Dyckia tenuis, 955 Dyckia tobatiensis, 955

Index Dyckia tomentella, 955 Dyckia tomentosa, 930 Dyckia trichostachya, 955 Dyckia tuberosa, 956 Dyckia tuberosa var. deltoidea, 906 Dyckia tubifilamentosa, 956–957 Dyckia tweediei, 957 Dyckia uleana, 957 Dyckia ursina, 957 Dyckia vaginosa, 947 Dyckia velascana, 957–958 Dyckia velloziifolia, 958 Dyckia vestita, 958 Dyckia vicentensis, 958–959 Dyckia virgata, 959 Dyckia waechteri, 959 Dyckia walteriana, 959 Dyckia warmingii, 960 Dyckia weddelliana, 893–960 Dyckipu, 1069 Dycktia, 963 E Ebertia, 1250 Elephantodon, 1201 Eliokarmos, 1281 Eliokarmos caudatum, 1233 Eliokarmos craibii, 1285 Eliokarmos geniculatus, 1283 Eliokarmos naviculus, 1285 Eliokarmos neomaculatus, 1285 Eliokarmos pendens, 1285 Eliokarmos thunbergianus, 1284 Elsiea, 1282 Encholirion, 965 Encholirium, 965 Encholirium agavoides, 966 Encholirium ascendens, 966 Encholirium bahianum, 978 Encholirium belemii, 967 Encholirium biflorum, 967 Encholirium brachypodum, 967 Encholirium bracteatum, 968 Encholirium bradeanum, 968 Encholirium carmineoviridiflorum, 978 Encholirium catharinense, 911 Encholirium crassiscapum, 975 Encholirium ctenophyllum, 968 Encholirium densiflorum, 978 Encholirium diamantinae, 966 Encholirium diamantinum, 969 Encholirium disjunctum, 969 Encholirium eddie-estevesii, 969 Encholirium erectiflorum, 970 Encholirium fragae, 970 Encholirium garrelii, 911 Encholirium glaziovii, 979 Encholirium gracile, 970

1427 Encholirium harleyi, 978 Encholirium heloisae, 971 Encholirium hoehneanum, 978 Encholirium horridum, 971, 983 Encholirium inerme, 978 Encholirium irwinii, 972 Encholirium josinoi-narcisae, 972 Encholirium kranzianum, 973 Encholirium longiflorum, 973 Encholirium lutzii, 978 Encholirium luxor, 973 Encholirium lymanianum, 974 Encholirium magalhaesii, 974 Encholirium maximum, 975 Encholirium paraibae, 978 Encholirium patens, 978 Encholirium pedicellatum, 975 Encholirium pernambucanum, 978 Encholirium pierre-braunii, 976 Encholirium piresianum, 973 Encholirium pulchrum, 977 Encholirium reflexum, 977 Encholirium rupestre, 978 Encholirium sabinae, 966 Encholirium sanguinolentum, 966 Encholirium sazimae, 971 Encholirium scrutor, 977 Encholirium spectabile, 978 Encholirium splendidum, 979 Encholirium subsecundum, 979 Encholirium suzannae, 975 Encholirium viride, 980 Encholirium viridicentrum, 980 Encholirium vogelii, 980 Enchotia, 983 Ephemeraceae, 1167 Epidendroideae, 1301 Epipetrum, 1201 Eratobotrys, 1269 Eremurus, 478 Eriospermaceae, 1217, 1316 Eriospermum, 1217, 1219 Eriospermum arenicola, 1220 Eriospermum armianum, 1220 Eriospermum bowieanum, 1220 Eriospermum coralliferum, 1220 Eriospermum cylindricum, 1220 Eriospermum paradoxum, 1220 Eriospermum strachaniae, 1220 Eriospermum titanopsoides, 1220 Eriospermum vallisgratiae, 1220 Eriostax, 851 Erythrotis beddomei, 1179 Ethesia, 1281 Eucallias, 859 Eucomidaceae, 1225 Eucomis bifolia, 1274 Eustachys, 1282 Eustrephaceae, 1293–1294

1428 F Falconera, 1231 Falconera spiralis, 1239 Fascicularia, 985 Fascicularia bicolor, 986 Fascicularia bicolor ssp. bicolor, 986 Fascicularia bicolor ssp. canaliculata, 986 Fascicularia kirchhoffiana, 986 Fascicularia litoralis, 1043 Fascicularia micrantha, 986 Fascicularia parviflora, 986 Fascicularia pitcairniifolia, 1043 Fenelonia, 1231 Fenelonia bracteata, 1233 Forzzaea, 865 Forzzaea leopoldo-horstii, 869 Forzzaea micra, 870 Forzzaea warasii, 871 Fosterella, 989 Fosterella albicans, 990 Fosterella chiquitana, 991 Fosterella cotacajensis, 990 Fosterella elata, 992 Fosterella fuentesii, 990 Fosterella latifolia, 991 Fosterella nowickii, 993 Fosterella penduliflora, 991 Fosterella rexiae, 991 Fosterella rusbyi, 992 Fosterella vasquezii, 992 Fosterella weddelliana, 993 Fosterella windischii, 993 Fosterella yuvinkae, 994 Fourcraea, 323 Fourcroea, 323 Fourcroya, 323 Fourcroya bedinghausii, 337 Fourcroya roezlii, 337 Fourcroya tuberosa, 342 Fourcroya vivipara, 343 Funium, 323 Funium pitiferum, 329 Funkiaceae, 9 Furcraea, 12, 14, 15, 323 Furcraea acaulis, 325–326 Furcraea agavephylla, 332 Furcraea aitonii, 344 Furcraea albispina, 341 Furcraea altissima, 325, 329, 332 Furcraea andina, 332 Furcraea antillana, 326–327 Furcraea aspera, 332 Furcraea atroviridis, 329 Furcraea australis, 1213 Furcraea barillettii, 329 Furcraea bedinghausii, 337 Furcraea boliviensis, 327 Furcraea cabuya, 327–328 Furcraea cabuya var. integra, 329

Index Furcraea cahum, 328 Furcraea cantala, 78 Furcraea commelynii, 343 Furcraea cubensis, 325, 332 Furcraea cubensis var. inermis, 329 Furcraea deledevantii, 332 Furcraea delevantii, 332 Furcraea demouliniana, 325 Furcraea depauperata, 328–329 Furcraea elegans, 332 Furcraea flaccida, 337 Furcraea flavoviridis, 341 Furcraea foetida, 329–330 Furcraea foetida cv. mediopicta, 329 Furcraea foetida cv. striata, 330 Furcraea geminispina, 330–331 Furcraea ghiesbreghtii, 332 Furcraea gigantea, 146, 329 Furcraea gigantea [?] mediopicta, 329 Furcraea gigantea [?] variegata, 329 Furcraea gigantea var. mediopicta, 329 Furcraea gigantea var. watsoniana, 329 Furcraea gigantea var. willemetiana, 343 Furcraea guatemalensis, 331–332 Furcraea guerrerensis, 338 Furcraea hexapetala, 332–333 Furcraea humboldtiana, 325 Furcraea interrupta, 343 Furcraea lindenii, 341 Furcraea lipsiensis, 343 Furcraea longa, 333 Furcraea longaeva, 333–334 Furcraea longaeva ssp. bedinghausii, 337 Furcraea macdougallii, 334 Furcraea macra, 329 Furcraea macrophylla, 332 Furcraea madagascariensis, 329 Furcraea martinezii, 334–335 Furcraea melanodonta, 331 Furcraea nana, 330 Furcraea niquivilensis, 335–336 Furcraea noackii, 205 Furcraea occidentalis, 336 Furcraea parmentieri, 336–338 Furcraea pubescens, 338–339, 344 Furcraea pugioniformis, 332 Furcraea quicheensis, 339–340 Furcraea rigida, 50, 325 Furcraea roezlii, 325 Furcraea roezlii var. atropurpurea, 325 Furcraea samalana, 340–341 Furcraea selloa, 341 Furcraea selloana, 341–342 Furcraea selloana fa. edentata, 341 Furcraea selloana var. edentata, 341 Furcraea selloana var. marginata, 341 Furcraea sisalana, 240 Furcraea sobolifera, 325 Furcraea spinosa, 343

Index Furcraea stratiotes, 325 Furcraea stricta, 342 Furcraea tuberosa, 146, 329, 341, 342–344, 343 Furcraea tuberosa var. geminispina, 330 Furcraea undulata, 344 Furcraea valleculata, 332 Furcraea variegata, 329 Furcraea viridis, 329 Furcraea watsoniana, 329 Furcroea, 323 Furcroya, 323 Furcroya cubensis, 332 Furcroya gigantea, 329 Furcroya tubiflora, 316 Fusifilum, 1249 G Galanthaceae, 425 Galtonia, 1282 Garrelia, 893 Garrelia encholirioides, 911 Gaslauminia, 745 Gasteraloe, 745 Gasteraloe bedinghausii, 745 Gasteraloe beguinii, 745 Gasteraloe beguinii nvar. beguinii, 745 Gasteraloe beguinii nvar. chludowii, 746 Gasteraloe beguinii nvar. perfectior, 746 Gasteraloe derbetzei, 746 Gasteraloe gloriosa, 746 Gasteraloe lapaixii, 746 Gasteraloe lapaixii nvar. lapaixii, 746 Gasteraloe lapaixii nvar. latifolia, 746 Gasteraloe lynchii, 746 Gasteraloe mortolensis, 746–747 Gasteraloe nowotnyi, 747 Gasteraloe peacockii, 747 Gasteraloe pethamensis, 747 Gasteraloe pfrimmeri, 747 Gasteraloe prorumpens, 747 Gasteraloe quehlii, 747 Gasteraloe radlii, 747 Gasteraloe rebutii, 747 Gasteraloe sculptilis, 747 Gasteraloe simoniana, 748 Gasteraloe smaragdina, 748 Gasterhaworthia, 749 Gasteria, 490, 751 Gasteria acinacifolia, 753 Gasteria acinacifolia var. acinacifolia, 753 Gasteria acinacifolia var. ensifolia, 753 Gasteria acinacifolia var. nitens, 753 Gasteria acinacifolia var. pluripuncta, 753 Gasteria acinacifolia var. spathulata, 753 Gasteria acinacifolia var. venusta, 753 Gasteria angulata, 757 Gasteria angulata var. truncata, 753 Gasteria angustiarum, 757

1429 Gasteria angustifolia, 759 Gasteria antandroi, 502 Gasteria apicroides, 767 Gasteria armstrongii, 753 Gasteria barbae, 754 Gasteria batesiana, 754 Gasteria batesiana var. batesiana, 754 Gasteria batesiana var. dolomitica, 755 Gasteria baylissiana, 755 Gasteria beckeri, 762 Gasteria bicolor, 755 Gasteria bicolor var. bicolor, 755 Gasteria bicolor var. fallax, 756 Gasteria bicolor var. liliputana, 756 Gasteria biformis, 756 Gasteria bijliae, 757 Gasteria brachyphylla, 756 Gasteria brachyphylla var. bayeri, 756 Gasteria brachyphylla var. brachyphylla, 757 Gasteria brevifolia, 753 Gasteria brevifolia var. laetevirens, 753 Gasteria brevifolia var. perviridis, 753 Gasteria caespitosa, 755 Gasteria candicans, 753 Gasteria carinata, 757 Gasteria carinata var. carinata, 757 Gasteria carinata var. falcata, 757 Gasteria carinata var. glabra, 758 Gasteria carinata var. kewensis, 757 Gasteria carinata var. latifolia, 757 Gasteria carinata var. parva, 757 Gasteria carinata var. retusa, 765 Gasteria carinata var. strigata, 757 Gasteria carinata var. thunbergii, 765 Gasteria carinata var. verrucosa, 758 Gasteria chamaegigas, 755 Gasteria colubrina, 755 Gasteria conspurcata, 759 Gasteria crassifolia, 753 Gasteria croucheri, 758 Gasteria croucheri ssp. croucheri, 759 Gasteria croucheri ssp. pendulifolia, 759 Gasteria croucheri ssp. pondoensis, 759 Gasteria decipiens, 762 Gasteria denticulata, 759 Gasteria dicta, 753 Gasteria disticha, 759 Gasteria disticha var. angulata, 757 Gasteria disticha var. angustifolia, 759 Gasteria disticha var. conspurcata, 759 Gasteria disticha var. disticha, 759 Gasteria disticha var. langebergensis, 760 Gasteria disticha var. major, 760 Gasteria disticha var. minor, 760 Gasteria disticha var. natalensis, 759 Gasteria disticha var. robusta, 760 Gasteria doreeniae, 760 Gasteria ellaphieae, 760 Gasteria elongata, 753

1430 Gasteria ensifolia, 753 Gasteria ernesti-ruschii, 763 Gasteria excavata, 757 Gasteria excelsa, 760 Gasteria fasciata, 752 Gasteria fasciata var. laxa, 753 Gasteria fasciata var. polyspila, 753 Gasteria formosa, 753 Gasteria fuscopunctata, 760 Gasteria glabra, 758 Gasteria glauca, 761 Gasteria glomerata, 761 Gasteria gracilis, 753 Gasteria herreana, 755 Gasteria humilis, 757 Gasteria huttoniae, 760 Gasteria inexpectata, 753 Gasteria intermedia, 758 Gasteria intermedia var. asperrima, 758 Gasteria intermedia var. laevior, 758 Gasteria intermedia var. longior, 758 Gasteria joubertii, 757 Gasteria kirsteana, 756 Gasteria koenii, 761 Gasteria laetepuncta, 757 Gasteria laevis, 753 Gasteria latifolia, 759 Gasteria latifolia var. multifaria, 757 Gasteria liliputana, 756 Gasteria linita, 753 Gasteria loedolffiae, 762 Gasteria loeriensis, 756 Gasteria longiana, 755 Gasteria longibracteata, 756 Gasteria longifolia, 759 Gasteria lutzii, 760 Gasteria maculata, 755 Gasteria maculata var. dregeana, 755 Gasteria maculata var. fallax, 756 Gasteria maculata var. liliputana, 756 Gasteria maculata var. maculata, 755 Gasteria marmorata, 755 Gasteria mollis, 760 Gasteria multiplex, 756 Gasteria neliana, 763 Gasteria nigricans, 759 Gasteria nigricans var. crassifolia, 759 Gasteria nigricans var. fasciata, 752 Gasteria nigricans var. guttata, 752 Gasteria nigricans var. marmorata, 757 Gasteria nigricans var. platyphylla, 757 Gasteria nigricans var. polyspila, 753 Gasteria nigricans var. subnigricans, 753 Gasteria nitens, 753 Gasteria nitida, 762 Gasteria nitida var. armstrongii, 753 Gasteria nitida var. grandipunctata, 762 Gasteria nitida var. parvipunctata, 762 Gasteria obliqua, 755 Gasteria obtusa, 762 Gasteria obtusifolia, 760

Index Gasteria pallescens, 757 Gasteria parva, 757 Gasteria parvifolia, 757 Gasteria patentissima, 758 Gasteria peacockii, 747 Gasteria pendulifolia, 759 Gasteria pethamensis, 747 Gasteria picta, 755 Gasteria picta var. formosa, 753 Gasteria pillansii, 763 Gasteria pillansii var. ernesti-ruschii, 763 Gasteria pillansii var. hallii, 763 Gasteria pillansii var. pillansii, 763 Gasteria planifolia, 755 Gasteria pluripuncta, 753 Gasteria poellnitziana, 764 Gasteria polita, 763 Gasteria porphyrophylla, 757 Gasteria prolifera, 753 Gasteria pseudonigricans, 752 Gasteria pseudonigricans var. canaliculata, 753 Gasteria pseudonigricans var. glabrior, 753 Gasteria pulchra, 764 Gasteria radulosa, 758 Gasteria rawlinsonii, 764 Gasteria repens, 758 Gasteria retata, 755 Gasteria retusa, 765 Gasteria salmdyckiana, 756 Gasteria schweickerdtiana, 757 Gasteria spiralis, 755 Gasteria spiralis var. tortulata, 755 Gasteria stayneri, 762 Gasteria strigata, 757 Gasteria subcarinata, 757 Gasteria subnigricans, 753 Gasteria subnigricans var. canaliculata, 752 Gasteria subnigricans var. glabrior, 753 Gasteria subverrucosa, 758 Gasteria subverrucosa var. grandipunctata, 758 Gasteria subverrucosa var. marginata, 758 Gasteria subverrucosa var. parvipunctata, 758 Gasteria sulcata, 757 Gasteria thunbergii, 765 Gasteria transvaalensis, 753 Gasteria triebneriana, 757 Gasteria trigona, 753 Gasteria tukhelensis, 765 Gasteria variolosa, 755 Gasteria venusta, 753 Gasteria verrucosa var. asperrima, 758 Gasteria verrucosa var. latifolia, 758 Gasteria verrucosa var. scaberrima, 758 Gasteria verrucosa var. striata, 758 Gasteria vlaaktensis, 757 Gasteria vlokii, 766 Gasteria zeyheri, 755 Gasterlirion, 745 Gastonialoe, 745 Gastrolea, 745 Gastrolea bedinghausii, 745

Index Gastrolea beguinii, 745 Gastrolea beguinii nvar. chludowii, 746 Gastrolea beguinii var. perfectior, 746 Gastrolea chludowii, 746 Gastrolea derbetzei, 746 Gastrolea imbricata, 747 Gastrolea lapaixii, 746 Gastrolea lynchii, 746 Gastrolea mortolensis, 746 Gastrolea nowotnyi, 747 Gastrolea peacockii, 747 Gastrolea perfectior, 746 Gastrolea pethamensis, 747 Gastrolea pfrimmeri, 747 Gastrolea prorumpens, 747 Gastrolea quehlii, 747 Gastrolea rebutii, 747 Gastrolea sculptilis, 747 Gastrolea simoniana, 748 Gastrolea smaragdina, 748 Gastrolirion, 745 Gastroloba, 767 Gastroloba apicroides, 767 Gastronema sanguineum, 438 Gastulista, 745 Gasworthia, 749 Gasworthiopsis, 749 Geschollia, 1250 Geschollia anomala, 1252 Gethyllidaceae, 425 Ghiesbreghtia, 22 Ghiesbreghtia dentata, 63 Ghiesbreghtia mollis, 62 Gonatopus, 458 Gonialoe, 486 Gonialoe dinteri, 547 Gonialoe sladeniana, 657 Gonialoe variegata, 674 Gonimara, 486, 487 Gonimara corderoyi, 538 Gramineae, 1307 Gravisia, 851 Guillauminia, 486 Guillauminia albiflora, 496 Guillauminia bakeri, 508 Guillauminia bellatula, 512 Guillauminia calcairophila, 523 Guillauminia descoingsii, 545 Guillauminia rauhii, 643 Guzmania, 840 Guzmania lingulata, 840 Guzmania monostachia, 840 H Habenaria repens, 1299 Hadrodemas, 1173 Hadrodemas warszewiczianum, 1176 Haemanthaceae, 425 Haemanthus, 426, 441 Haemanthus albiflos, 442

1431 Haemanthus albiflos var. burchellii, 442 Haemanthus albiflos var. brachyphyllus, 442 Haemanthus albomaculatus, 442 Haemanthus avasimontanus, 442 Haemanthus baurii, 442 Haemanthus ciliaris, 429, 442 Haemanthus deformis, 442 Haemanthus distichus, 429 Haemanthus intermedius, 442 Haemanthus lemairei, 430 Haemanthus leucanthus, 442 Haemanthus mackenii, 442 Haemanthus pauculifolius, 443 Haemanthus pubescens, 442 Haemanthus robustus, 430 Haemanthus sinuatus, 430 Haemanthus toxicarius, 429 Haemanthus virescens var. albiflos, 442 Haemanthus virescens var. intermedius, 442 Hamatris, 1201 Hapalanthus repens, 1176 Hartwegia comosa, 452 Hawiopsis, 769 Haworthia, 769 Haworthia acuminata, 804 Haworthia adelaidensis, 780 Haworthia aegrota, 790 Haworthia affinis, 772 Haworthia agavoides, 810 Haworthia agnis, 803 Haworthia albanensis, 774 Haworthia albertinensis, 805 Haworthia albicans, 794 Haworthia albicans var. virescens, 794 Haworthia albispina, 802 Haworthia allomadis, 800 Haworthia altilinea, 772 Haworthia altilinea fa. acuminata, 772 Haworthia altilinea fa. inconfluens, 800 Haworthia altilinea fa. inermis, 778 Haworthia altilinea fa. minor, 772 Haworthia altilinea fa. subglauca, 772 Haworthia altilinea fa. typica, 801 Haworthia altilinea var. bicarinata, 772 Haworthia altilinea var. brevisetata, 772 Haworthia altilinea var. denticulata, 776 Haworthia altilinea var. inermis, 778 Haworthia altilinea var. limpida, 801 Haworthia altilinea var. morrisiae, 800 Haworthia altilinea var. mucronata, 800 Haworthia altilinea var. polyphylla, 773 Haworthia altilinea var. setulifera, 772 Haworthia altilinea var. typica, 772 Haworthia altissima, 774 Haworthia amethysta, 785 Haworthia angiras, 776 Haworthia angolensis, 681 Haworthia angustata, 784 Haworthia angustifolia, 774 Haworthia angustifolia fa. baylissii, 774 Haworthia angustifolia fa. grandis, 774

1432 Haworthia angustifolia fa. paucifolia, 774 Haworthia angustifolia var. albanensis, 774 Haworthia angustifolia var. altissima, 774 Haworthia angustifolia var. angustifolia, 774 Haworthia angustifolia var. baylissii, 774 Haworthia angustifolia var. denticulifera, 779 Haworthia angustifolia var. grandis, 774 Haworthia angustifolia var. liliputana, 779 Haworthia angustifolia var. paucifolia, 774 Haworthia angustifolia var. subfalcata, 797 Haworthia anna, 774 Haworthia ao-onii, 793 Haworthia apta, 775 Haworthia aquamarina, 779 Haworthia arabesqua, 782 Haworthia arachnoidea, 774 Haworthia arachnoidea ssp. pearsonii, 773 Haworthia arachnoidea ssp. setata, 776 Haworthia arachnoidea ssp. tenera, 783 Haworthia arachnoidea var. angiras, 776 Haworthia arachnoidea var. arachnoidea, 774 Haworthia arachnoidea var. aranea, 775 Haworthia arachnoidea var. bolusii, 778 Haworthia arachnoidea var. calitzdorpensis, 800 Haworthia arachnoidea var. gigas, 776 Haworthia arachnoidea var. gracilis, 781 Haworthia arachnoidea var. joubertii, 775 Haworthia arachnoidea var. laxa, 775 Haworthia arachnoidea var. limbata, 775 Haworthia arachnoidea var. marumiana, 794 Haworthia arachnoidea var. minor, 774 Haworthia arachnoidea var. namaquensis, 775 Haworthia arachnoidea var. nigricans, 775 Haworthia arachnoidea var. pearsonii, 773 Haworthia arachnoidea var. pehlemanniae, 803 Haworthia arachnoidea var. scabrispina, 775 Haworthia arachnoidea var. semiviva, 810 Haworthia arachnoidea var. setata, 776 Haworthia arachnoidea var. tenera, 783 Haworthia arachnoidea var. translucens, 789 Haworthia arachnoidea var. xiphiophylla, 786 Haworthia aranea, 775 Haworthia aranea var. candida, 776 Haworthia araneoides, 775 Haworthia arcana, 791 Haworthia archeri, 794 Haworthia archeri var. archeri, 794 Haworthia archeri var. dimorpha, 794 Haworthia argyrostigma, 772 Haworthia aristata, 776 Haworthia aristata ssp. helmiae, 773 Haworthia aristata var. helmiae, 773 Haworthia armata, 801 Haworthia armstrongii, 789 Haworthia asema, 799 Haworthia aspera, 703, 772 Haworthia aspera var. major, 772 Haworthia asperata, 805 Haworthia asperiuscula, 813

Index Haworthia asperiuscula var. patagiata, 813 Haworthia asperiuscula var. subintegra, 813 Haworthia asperula, 772 Haworthia atrofusca, 796 Haworthia atrofusca var. enigma, 804 Haworthia atrovirens, 789 Haworthia atroviridis, 789 Haworthia attenuata, 776 Haworthia attenuata fa. britteniana, 777 Haworthia attenuata fa. caespitosa, 777 Haworthia attenuata fa. clariperla, 777 Haworthia attenuata var. argyrostigma, 772 Haworthia attenuata var. attenuata, 777 Haworthia attenuata var. britteniana, 777 Haworthia attenuata var. caespitosa, 777 Haworthia attenuata var. clariperla, 777 Haworthia attenuata var. deltoidea, 777 Haworthia attenuata var. glabrata, 777 Haworthia attenuata var. inusitata, 777 Haworthia attenuata var. linearis, 777 Haworthia attenuata var. minissima, 777 Haworthia attenuata var. odonoghueana, 777 Haworthia attenuata var. radula, 777 Haworthia attenuata var. uitewaaliana, 777 Haworthia audens, 793 Haworthia auraria, 786 Haworthia azurea, 782 Haworthia baccata, 772 Haworthia badia, 796 Haworthia badia var. bobii, 799 Haworthia badia var. joleneae, 799 Haworthia batesiana, 794 Haworthia batesiana var. lepida, 784 Haworthia batesiana var. reddii, 795 Haworthia bathylis, 782 Haworthia batteniae, 778 Haworthia baviens, 799 Haworthia bayeri, 777 Haworthia bayeri var. hayashii, 777 Haworthia bayeri var. jadea, 777 Haworthia bayeri var. scabrifolia, 777 Haworthia baylissii, 774 Haworthia beanii, 813 Haworthia beanii var. minor, 813 Haworthia bella, 781 Haworthia beukmanii, 796 Haworthia bicarinata, 709 Haworthia bijliana, 773 Haworthia bijliana var. joubertii, 800 Haworthia bilineata, 773 Haworthia bilineata var. affinis, 772 Haworthia bilineata var. gracilidelineata, 773 Haworthia blackbeardiana, 778 Haworthia blackbeardiana var. calaensis, 778 Haworthia blackbeardiana var. major, 778 Haworthia blackbeardiana var. specksii, 778 Haworthia blackburniae, 778, 787 Haworthia blackburniae ssp. graminifolia, 778 Haworthia blackburniae var. blackburniae, 778

Index Haworthia blackburniae var. derustensis, 778 Haworthia blackburniae var. graminifolia, 778 Haworthia blackburniana, 778 Haworthia blinkia, 784 Haworthia bobii, 799 Haworthia bolusii, 778 Haworthia bolusii var. aranea, 775 Haworthia bolusii var. blackbeardiana, 778 Haworthia bolusii var. bolusii, 778 Haworthia bolusii var. floccosa, 786 Haworthia bolusii var. pringlei, 779 Haworthia bolusii var. semiviva, 810 Haworthia borealis, 795 Haworthia brandea, 781 Haworthia breueri, 787 Haworthia brevicula, 806 Haworthia brevis, 795 Haworthia britteniana, 777 Haworthia bronkhorstii, 799 Haworthia browniana, 787 Haworthia bruynsii, 779 Haworthia caerulea, 782 Haworthia caerulea var. pallens, 781 Haworthia caesia, 785 Haworthia caespitosa, 808 Haworthia caespitosa fa. subplana, 808 Haworthia caespitosa fa. subproliferans, 808 Haworthia caespitosa var. consanguinea, 797 Haworthia calaensis, 778 Haworthia calcarea, 808 Haworthia calitzdorpensis, 800 Haworthia calitzensis, 800 Haworthia calliantha, 797 Haworthia calva, 781 Haworthia cana, 784 Haworthia candida, 776 Haworthia cangoensis, 776 Haworthia cangoensis var. kogmanensis, 776 Haworthia cangoensis var. royalis, 776 Haworthia cangoensis var. tradouwensis, 801 Haworthia capillaris, 779 Haworthia carrissoi, 788 Haworthia cassytha, 773 Haworthia chalwinii, 780 Haworthia chibita, 794 Haworthia chloracantha, 779 Haworthia chloracantha ssp. angustifolia, 774 Haworthia chloracantha ssp. baylissii, 774 Haworthia chloracantha ssp. variegata, 812 Haworthia chloracantha var. angustifolia, 774 Haworthia chloracantha var. chloracantha, 779 Haworthia chloracantha var. denticulifera, 779 Haworthia chloracantha var. floribunda, 788 Haworthia chloracantha var. mclarenii, 801 Haworthia chloracantha var. monticola, 799 Haworthia chloracantha var. pulchella, 803 Haworthia chloracantha var. serrata, 808 Haworthia chloracantha var. subglauca, 779 Haworthia chloracantha var. variegata, 812

1433 Haworthia chloracantha var. zantneriana, 814 Haworthia chromutica, 807 Haworthia ciliata, 782 Haworthia cineraria, 781 Haworthia clariperla, 777 Haworthia coarctata, 779 Haworthia coarctata fa. bellula, 780 Haworthia coarctata fa. chalwinii, 780 Haworthia coarctata fa. conspicua, 780 Haworthia coarctata fa. greenii, 779 Haworthia coarctata fa. major, 780 Haworthia coarctata fa. pseudocoarctata, 780 Haworthia coarctata ssp. adelaidensis, 780 Haworthia coarctata var. adelaidensis, 780 Haworthia coarctata var. coarctata, 780 Haworthia coarctata var. greenii, 780 Haworthia coarctata var. haworthii, 780 Haworthia coarctata var. kraussii, 780 Haworthia coarctata var. sampaiana, 773 Haworthia coarctata var. tenuis, 780 Haworthia coarctatoides, 773 Haworthia cocksia, 774 Haworthia columnaris, 773 Haworthia compacta, 784 Haworthia compressa, 781 Haworthia comptoniana, 786 Haworthia comptoniana fa. brevifolia, 786 Haworthia comptoniana fa. major, 786 Haworthia concava, 783 Haworthia concinna, 813 Haworthia confluens, 801 Haworthia confusa, 773 Haworthia congesta, 702 Haworthia consanguinea, 797 Haworthia cooperi, 780 Haworthia cooperi fa. dielsiana, 781 Haworthia cooperi fa. pilifera, 782 Haworthia cooperi ssp. batteniae, 778 Haworthia cooperi var. cooperi, 780 Haworthia cooperi var. cymbiformoides, 781 Haworthia cooperi var. dielsiana, 781 Haworthia cooperi var. doldii, 781 Haworthia cooperi var. gordoniana, 781 Haworthia cooperi var. gracilis, 781 Haworthia cooperi var. hisui, 779 Haworthia cooperi var. isabellae, 782 Haworthia cooperi var. leightonii, 782 Haworthia cooperi var. minima, 783 Haworthia cooperi var. picturata, 782 Haworthia cooperi var. pilifera, 782 Haworthia cooperi var. tenera, 783 Haworthia cooperi var. truncata, 783 Haworthia cooperi var. venusta, 783 Haworthia cooperi var. viridis, 783 Haworthia cordifolia, 813 Haworthia coriacea, 811 Haworthia correcta, 787 Haworthia correcta var. lucida, 787 Haworthia corrugata, 703

1434 Haworthia corticosa, 799 Haworthia crausii, 811 Haworthia crinita, 785 Haworthia crousii, 811 Haworthia crystallina, 800 Haworthia cummingii, 783 Haworthia cuprina, 782 Haworthia curta, 773 Haworthia cuspidata, 773 Haworthia cyanea, 785 Haworthia cyanea var. amethysta, 785 Haworthia cyanea var. ianthina, 785 Haworthia cyanea var. succinea, 785 Haworthia cymbiformis, 783 Haworthia cymbiformis fa. gracilidelineata, 773 Haworthia cymbiformis fa. multifolia, 811 Haworthia cymbiformis fa. obesa, 785 Haworthia cymbiformis fa. planifolia, 783 Haworthia cymbiformis fa. ramosa, 785 Haworthia cymbiformis fa. subarmata, 784 Haworthia cymbiformis subvar. subarmata, 784 Haworthia cymbiformis var. angustata, 784 Haworthia cymbiformis var. brevifolia, 811 Haworthia cymbiformis var. compacta, 784 Haworthia cymbiformis var. cymbiformis, 783 Haworthia cymbiformis var. incurvula, 784 Haworthia cymbiformis var. multifolia, 811 Haworthia cymbiformis var. obesa, 785 Haworthia cymbiformis var. obtusa, 784 Haworthia cymbiformis var. planifolia, 783 Haworthia cymbiformis var. ramosa, 785 Haworthia cymbiformis var. reddii, 795 Haworthia cymbiformis var. setulifera, 785 Haworthia cymbiformis var. transiens, 811 Haworthia cymbiformis var. translucens, 811 Haworthia cymbiformis var. umbraticola, 784 Haworthia dasylis, 782 Haworthia davidii, 782 Haworthia decipiens, 785 Haworthia decipiens var. cyanea, 785 Haworthia decipiens var. decipiens, 785 Haworthia decipiens var. exilis, 785 Haworthia decipiens var. incrassa, 785 Haworthia decipiens var. minor, 785 Haworthia decipiens var. pringlei, 779 Haworthia decipiens var. scottiana, 785 Haworthia decipiens var. virella, 785 Haworthia decipiens var. xiphiophylla, 786 Haworthia dekenahii, 804 Haworthia dekenahii var. argenteo-maculosa, 804 Haworthia delicata, 786 Haworthia deltoidea, 703 Haworthia deltoidea var. intermedia, 703 Haworthia deltoidea var. turgida, 703 Haworthia dentata, 788 Haworthia denticulata, 776 Haworthia denticulifera, 779 Haworthia depauperata, 798 Haworthia derustensis, 778

Index Haworthia devriesii, 802 Haworthia diaphana, 811 Haworthia dielsiana, 781 Haworthia dimorpha, 794 Haworthia distincta, 812 Haworthia divergens, 799 Haworthia divergens var. meiringii, 798 Haworthia diversicolor, 797 Haworthia diversifolia, 802 Haworthia dodsoniana, 704 Haworthia doldii, 781 Haworthia dura, 776 Haworthia egregia, 702 Haworthia egregia var. fardeniana, 702 Haworthia eilyae, 789 Haworthia eilyae var. poellnitziana, 789 Haworthia eilyae var. zantneriana, 789 Haworthia elegans, 781 Haworthia elizeae, 808 Haworthia emelyae, 786 Haworthia emelyae var. beukmanii, 796 Haworthia emelyae var. comptoniana, 786 Haworthia emelyae var. emelyae, 787 Haworthia emelyae var. major, 787 Haworthia emelyae var. multifolia, 787 Haworthia emeralda, 784 Haworthia eminens, 786 Haworthia engleri, 811 Haworthia enigma, 804 Haworthia eonigra, 802 Haworthia erecta, 795 Haworthia erii, 800 Haworthia ernstii, 800 Haworthia esterhuizenii, 805, 808 Haworthia euchlora, 795 Haworthia exilis, 785 Haworthia expansa, 771 Haworthia fallax, 780 Haworthia fasciata, 787 Haworthia fasciata fa. browniana, 787 Haworthia fasciata fa. major, 787 Haworthia fasciata fa. ovato-lanceolata, 788 Haworthia fasciata fa. sparsa, 788 Haworthia fasciata fa. subconfluens, 788 Haworthia fasciata fa. vanstaadensis, 788 Haworthia fasciata fa. variabilis, 788 Haworthia fasciata var. browniana, 787 Haworthia fasciata var. caespitosa, 777 Haworthia fasciata var. major, 787 Haworthia fasciata var. subconfluens, 788 Haworthia fatreddii, 795 Haworthia fergusoniae, 773 Haworthia ferox, 773 Haworthia ferox var. armata, 773 Haworthia flaccida, 790 Haworthia flavens, 796 Haworthia flavida, 786 Haworthia floccosa, 786 Haworthia florens, 782

Index Haworthia floribunda, 788 Haworthia floribunda var. dentata, 788 Haworthia floribunda var. floribunda, 788 Haworthia floribunda var. major, 788 Haworthia florida, 782 Haworthia fluffa, 786 Haworthia foeda, 781 Haworthia foliolosa, 704 Haworthia formosa, 775 Haworthia fouchei, 807 Haworthia fraseri, 794 Haworthia fukuyae, 773 Haworthia fulva, 780 Haworthia fusca, 804 Haworthia gelatina, 781 Haworthia geraldii, 807 Haworthia gideonii, 792 Haworthia giftbergensis, 803 Haworthia gigantea, 791 Haworthia gigas, 776 Haworthia gilva, 775 Haworthia glabella, 799 Haworthia glabrata, 777 Haworthia glabrata var. concolor, 772 Haworthia glabrata var. perviridis, 772 Haworthia glauca, 788 Haworthia glauca fa. armstrongii, 789 Haworthia glauca fa. jacobseniana, 789 Haworthia glauca fa. jonesiae, 789 Haworthia glauca var. glauca, 788 Haworthia glauca var. herrei, 788 Haworthia glaucophylla, 792 Haworthia globifera, 803 Haworthia globosiflora, 802 Haworthia gordoniana, 781 Haworthia gracilidelineata, 773 Haworthia gracilis, 781 Haworthia gracilis var. isabellae, 782 Haworthia gracilis var. minor, 785 Haworthia gracilis var. picturata, 782 Haworthia gracilis var. tenera, 783 Haworthia gracilis var. viridis, 783 Haworthia graminifolia, 778 Haworthia graminifolia var. derustensis, 778 Haworthia granata, 795 Haworthia granata var. polyphylla, 795 Haworthia grandis, 774 Haworthia granulata, 789 Haworthia greenii, 780 Haworthia greenii fa. bakeri, 780 Haworthia greenii fa. minor, 780 Haworthia greenii fa. pseudocoarctata, 780 Haworthia greenii var. silvicola, 780 Haworthia grenieri, 789 Haworthia groenewaldii, 801, 810 Haworthia guttata, 806 Haworthia gweneana, 705 Haworthia haageana, 807 Haworthia haageana var. subreticulata, 807

1435 Haworthia habdomadis, 800 Haworthia habdomadis var. habdomadis, 800 Haworthia habdomadis var. inconfluens, 800 Haworthia habdomadis var. morrisiae, 800 Haworthia hallii, 702 Haworthia hamata, 783 Haworthia hamata var. subhamata, 783 Haworthia hammeri, 796 Haworthia harlandiana, 704 Haworthia harryi, 781 Haworthia hastata, 779 Haworthia hayashii, 777 Haworthia heidelbergensis, 797 Haworthia heidelbergensis var. minor, 808 Haworthia heidelbergensis var. scabra, 799 Haworthia heidelbergensis var. toonensis, 799 Haworthia helensis, 811 Haworthia helmiae, 773 Haworthia hemicrypta, 812 Haworthia henda, 788 Haworthia henriquesii, 773 Haworthia herbacea, 789 Haworthia herbacea var. flaccida, 790 Haworthia herbacea var. herbacea, 790 Haworthia herbacea var. lupula, 790 Haworthia herbacea var. paynei, 790 Haworthia heroldia, 803 Haworthia herrei, 788 Haworthia herrei var. depauperata, 789 Haworthia herrei var. poellnitzii, 789 Haworthia hilliana, 784 Haworthia hisui, 779 Haworthia hogsia, 778 Haworthia horrida, 800 Haworthia hurlingii, 806 Haworthia hurlingii var. ambigua, 806 Haworthia hybrida, 772 Haworthia ianthina, 785 Haworthia icosiphylla, 773 Haworthia ikra, 783 Haworthia imbricata, 705 Haworthia imperialis, 782 Haworthia inconfluens, 800 Haworthia inconfluens var. crystallina, 800 Haworthia inconfluens var. habdomadis, 800 Haworthia inconfluens var. lockwoodii, 792 Haworthia inconfluens var. morrisiae, 800 Haworthia incrassa, 785 Haworthia incurvula, 784 Haworthia indigoa, 777 Haworthia indigoa var. truterorum, 777 Haworthia indurata, 813 Haworthia inermis, 778 Haworthia ingens, 784 Haworthia inspida, 814 Haworthia integra, 801 Haworthia integra var. standeri, 800 Haworthia intermedia, 798 Haworthia intermedia var. livida, 793

1436 Haworthia intermedia var. maculata, 793 Haworthia intermedia var. notabilis, 798 Haworthia ionandra, 781 Haworthia ionides, 786 Haworthia irmiae, 812 Haworthia isabellae, 782 Haworthia isabellae var. arabesqua, 782 Haworthia isabellae var. bella, 781 Haworthia isabellae var. ligulata, 781 Haworthia isomorpha, 775 Haworthia jacobseniana, 789 Haworthia jadea, 777 Haworthia jakubii, 798 Haworthia janseana, 773 Haworthia jansenvillensis, 786 Haworthia jansenvillensis var. crinita, 785 Haworthia jansenvillensis var. eminens, 786 Haworthia jansenvillensis var. flavida, 786 Haworthia jansenvillensis var. fluffa, 786 Haworthia jansenvillensis var. mollis, 786 Haworthia jansenvillensis var. regina, 786 Haworthia janvlokii, 787 Haworthia jeffreis, 781 Haworthia joeyae, 781 Haworthia johanii, 809 Haworthia joleneae, 799 Haworthia jonesiae, 789 Haworthia joubertii, 774 Haworthia kammaensis, 786 Haworthia keithii, 792 Haworthia kemari, 786 Haworthia kingiana, 790 Haworthia klipensis, 811 Haworthia koelmaniorum, 790 Haworthia koelmaniorum var. koelmaniorum, 791 Haworthia koelmaniorum var. mcmurtryi, 791 Haworthia kogmanensis, 776 Haworthia kondoi, 788 Haworthia kotei, 800 Haworthia krausiana, 773 Haworthia krausii, 773 Haworthia kromia, 782 Haworthia kubusie, 782 Haworthia kuromisa, 775 Haworthia lachnosa, 782 Haworthia laeta, 777 Haworthia laetevirens, 808 Haworthia laevis, 794 Haworthia lanceata, 786 Haworthia lapis, 776 Haworthia lapis var. rava, 776 Haworthia lateganiae, 809 Haworthia lateritia, 803 Haworthia lavrani, 810 Haworthia laxa, 775 Haworthia lazulis, 779 Haworthia leightonii, 782 Haworthia leightonii var. davidii, 782 Haworthia leightonii var. doldii, 781

Index Haworthia lepida, 784 Haworthia ligulata, 781 Haworthia limbata, 775 Haworthia limifolia, 791 Haworthia limifolia fa. diploidea, 792 Haworthia limifolia fa. major, 792 Haworthia limifolia fa. marlothiana, 792 Haworthia limifolia fa. pimentelii, 792 Haworthia limifolia fa. schuldtiana, 792 Haworthia limifolia fa. tetraploidea, 792 Haworthia limifolia fa. variegata, 792 Haworthia limifolia ssp. koelmaniorum, 791 Haworthia limifolia var. arcana, 791 Haworthia limifolia var. diploidea, 792 Haworthia limifolia var. gigantea, 791 Haworthia limifolia var. glaucophylla, 792 Haworthia limifolia var. keithii, 792 Haworthia limifolia var. limifolia, 792 Haworthia limifolia var. marlothiana, 792 Haworthia limifolia var. schuldtiana, 792 Haworthia limifolia var. stolonifera, 792 Haworthia limifolia var. striata, 792 Haworthia limifolia var. tetraploidea, 792 Haworthia limifolia var. ubomboensis, 792 Haworthia limpida, 801 Haworthia lisbonensis, 773 Haworthia livida, 793 Haworthia lockwoodii, 792 Haworthia longiana, 792 Haworthia longiana var. albinota, 792 Haworthia longiaristata, 786 Haworthia longibracteata, 807 Haworthia longifolia, 773 Haworthia lupula, 790 Haworthia luri, 783 Haworthia luteorosea, 790 Haworthia macmurtryi, 791 Haworthia maculata, 793 Haworthia maculata var. intermedia, 798 Haworthia maculata var. livida, 793 Haworthia maculata var. maculata, 793 Haworthia magnifica, 797 Haworthia magnifica var. acuminata, 804 Haworthia magnifica var. asperula, 772 Haworthia magnifica var. atrofusca, 796 Haworthia magnifica var. dekenahii, 804 Haworthia magnifica var. magnifica, 797 Haworthia magnifica var. major, 787 Haworthia magnifica var. maraisii, 797 Haworthia magnifica var. meiringii, 798 Haworthia magnifica var. notabilis, 798 Haworthia magnifica var. obserata, 797 Haworthia magnifica var. paradoxa, 798 Haworthia magnifica var. pseudomutica, 804 Haworthia magnifica var. splendens, 799 Haworthia major, 795 Haworthia malvina, 778 Haworthia mantelii, 793 Haworthia maraisii, 797

Index Haworthia maraisii var. livida, 793 Haworthia maraisii var. magnifica, 797 Haworthia maraisii var. major, 787 Haworthia maraisii var. meiringii, 798 Haworthia maraisii var. notabilis, 798 Haworthia maraisii var. paradoxa, 798 Haworthia maraisii var. pubescens, 797 Haworthia maraisii var. schuldtiana, 797 Haworthia maraisii var. toonensis, 799 Haworthia margaritifera, 793 Haworthia margaritifera subvar. acuminata, 772 Haworthia margaritifera subvar. laetivirens, 795 Haworthia margaritifera subvar. laevior, 772 Haworthia margaritifera subvar. major, 772 Haworthia margaritifera subvar. maxima, 793 Haworthia margaritifera subvar. minor, 795 Haworthia margaritifera subvar. multipapillosa, 772 Haworthia margaritifera subvar. multiperlata, 773 Haworthia margaritifera subvar. polyphylla, 795 Haworthia margaritifera var. corallina, 796 Haworthia margaritifera var. erecta, 795 Haworthia margaritifera var. granata, 795 Haworthia margaritifera var. maxima, 793 Haworthia margaritifera var. minima, 795 Haworthia margaritifera var. minor, 795 Haworthia margaritifera var. semimargaritifera, 793 Haworthia margaritifera var. subalbicans, 772 Haworthia marginata, 794 Haworthia marginata var. laevis, 794 Haworthia marginata var. ramifera, 794 Haworthia marginata var. virescens, 794 Haworthia marmorata, 795 Haworthia marumiana, 794 Haworthia marumiana var. archeri, 794 Haworthia marumiana var. batesiana, 794 Haworthia marumiana var. dimorpha, 794 Haworthia marumiana var. marmorata, 795 Haworthia marumiana var. marumiana, 795 Haworthia marumiana var. reddii, 795 Haworthia marumiana var. viridis, 795 Haworthia marxii, 795 Haworthia matjiesta, 775 Haworthia maughanii, 811 Haworthia maxima, 793 Haworthia mclarenii, 801 Haworthia mcmurtryi, 791 Haworthia mediata, 811 Haworthia meiringii, 798 Haworthia minima, 783, 795 Haworthia minima var. confusa, 773 Haworthia minima var. major, 776 Haworthia minima var. minima, 796 Haworthia minima var. poellnitziana, 796 Haworthia minor, 795 Haworthia minutissima, 811 Haworthia mirabilis, 796 Haworthia mirabilis fa. beukmanii, 796 Haworthia mirabilis fa. napierensis, 798 Haworthia mirabilis fa. rubrodentata, 798

1437 Haworthia mirabilis fa. sublineata, 799 Haworthia mirabilis ssp. badia, 796 Haworthia mirabilis ssp. mundula, 798 Haworthia mirabilis var. atrofusca, 796 Haworthia mirabilis var. badia, 796 Haworthia mirabilis var. beukmanii, 796 Haworthia mirabilis var. calcarea, 808 Haworthia mirabilis var. consanguinea, 797 Haworthia mirabilis var. depauperata, 798 Haworthia mirabilis var. heidelbergensis, 797 Haworthia mirabilis var. magnifica, 797 Haworthia mirabilis var. maraisii, 797 Haworthia mirabilis var. meiringii, 797 Haworthia mirabilis var. mirabilis, 798 Haworthia mirabilis var. mundula, 798 Haworthia mirabilis var. notabilis, 798 Haworthia mirabilis var. paradoxa, 798 Haworthia mirabilis var. scabra, 799 Haworthia mirabilis var. splendens, 799 Haworthia mirabilis var. sublineata, 799 Haworthia mirabilis var. toonensis, 799 Haworthia mirabilis var. triebneriana, 798 Haworthia modesta, 812 Haworthia mollis, 786 Haworthia montagua, 801 Haworthia montana, 803 Haworthia monticola, 799 Haworthia monticola var. asema, 799 Haworthia monticola var. bronkhorstii, 799 Haworthia monticola var. monticola, 799 Haworthia morrisiae, 809 Haworthia mortonii, 796 Haworthia mucronata, 800 Haworthia mucronata fa. acuminata, 772 Haworthia mucronata fa. inconfluens, 800 Haworthia mucronata fa. inermis, 778 Haworthia mucronata fa. minor, 772 Haworthia mucronata fa. subglauca, 772 Haworthia mucronata fa. typica, 801 Haworthia mucronata ssp. lockwoodii, 792 Haworthia mucronata var. altilinea, 772 Haworthia mucronata var. bicarinata, 772 Haworthia mucronata var. calitzdorpensis, 800 Haworthia mucronata var. habdomadis, 800 Haworthia mucronata var. inconfluens, 800 Haworthia mucronata var. limpida, 801 Haworthia mucronata var. morrisiae, 800 Haworthia mucronata var. mucronata, 801 Haworthia mucronata var. nortieri, 802 Haworthia mucronata var. polyphylla, 773 Haworthia mucronata var. rooibergensis, 800 Haworthia mucronata var. rycroftiana, 801 Haworthia mucronata var. setulifera, 772 Haworthia multifaria, 773 Haworthia multifolia, 787 Haworthia multifolia var. breueri, 787 Haworthia multifolia var. major, 787 Haworthia multifolia var. sandkraalensis, 787 Haworthia multilineata, 807

1438 Haworthia mundula, 798 Haworthia mundula var. calcarea, 808 Haworthia musculina, 780 Haworthia mutabilis, 796 Haworthia mutica, 801 Haworthia mutica var. nigra, 807 Haworthia namaquensis, 775 Haworthia neritica, 781 Haworthia nigra, 801 Haworthia nigra fa. angustata, 802 Haworthia nigra fa. nana, 802 Haworthia nigra var. angustata, 802 Haworthia nigra var. diversifolia, 801 Haworthia nigra var. elongata, 802 Haworthia nigra var. nigra, 802 Haworthia nigra var. pusilla, 802 Haworthia nigra var. schmidtiana, 802 Haworthia nigra var. suberecta, 802 Haworthia nigrata, 775 Haworthia nigricans, 762 Haworthia nitidula, 798 Haworthia nitidula var. opaca, 798 Haworthia nortieri, 802 Haworthia nortieri var. agnis, 803 Haworthia nortieri var. albispina, 802 Haworthia nortieri var. devriesii, 802 Haworthia nortieri var. giftbergensis, 803 Haworthia nortieri var. globosiflora, 802 Haworthia nortieri var. montana, 803 Haworthia nortieri var. nortieri, 802 Haworthia nortieri var. pehlemanniae, 803 Haworthia notabilis, 798 Haworthia notabilis var. diversicolor, 797 Haworthia nudata, 794 Haworthia obesa, 785 Haworthia obrata, 796 Haworthia obscura, 773 Haworthia obserata, 797 Haworthia obtusa, 784 Haworthia obtusa fa. acuminata, 783 Haworthia obtusa fa. truncata, 783 Haworthia obtusa var. columnaris, 773 Haworthia obtusa var. dielsiana, 781 Haworthia obtusa var. gordoniana, 781 Haworthia obtusa var. pilifera, 782 Haworthia obtusa var. salina, 783 Haworthia obtusa var. stayneri, 782 Haworthia oculata, 782 Haworthia odetteae, 778 Haworthia odetteae var. odyssei, 779 Haworthia odyssei, 778 Haworthia ohkuwae, 793 Haworthia olivacea, 805 Haworthia olivettiana, 709 Haworthia ollasonii, 773 Haworthia opalina, 796 Haworthia otzenii, 801 Haworthia outeniquensis, 803 Haworthia oxygona, 806

Index Haworthia pallens, 781 Haworthia pallida, 789 Haworthia pallida var. flaccida, 790 Haworthia pallida var. paynei, 790 Haworthia pallidifolia, 807 Haworthia papillaris, 812 Haworthia papillosa, 789 Haworthia papillosa var. semipapillosa, 789 Haworthia paradoxa, 798 Haworthia paradoxa var. jakubii, 798 Haworthia parksiana, 803 Haworthia parksiana var. dentata, 788 Haworthia parva, 810 Haworthia patriae, 782 Haworthia paucifolia, 774 Haworthia paynei, 790 Haworthia peacockii, 780 Haworthia pearsonii, 773 Haworthia pectinis, 776 Haworthia pehlemanniae, 803 Haworthia pehlemanniae var. albispina, 802 Haworthia pellucens, 789 Haworthia pellucens var. delicatula, 773 Haworthia pellucida, 786 Haworthia pentagona, 705 Haworthia pentagona var. spiralis, 705 Haworthia pentagona var. spirella, 705 Haworthia pentagona var. torulosa, 705 Haworthia perplexa, 773 Haworthia petrophila, 808 Haworthia picta, 787 Haworthia picta var. comptoniana, 786 Haworthia picta var. janvlokii, 787 Haworthia picta var. tricolor, 787 Haworthia picturata, 782 Haworthia picturata var. pusilla, 781 Haworthia pilifera, 782 Haworthia pilifera fa. acuminata, 783 Haworthia pilifera var. columnaris, 773 Haworthia pilifera var. dielsiana, 781 Haworthia pilifera var. gordoniana, 781 Haworthia pilifera var. salina, 782 Haworthia pilifera var. stayneri, 782 Haworthia pilosa, 782 Haworthia pilosa var. ciliata, 782 Haworthia pilosa var. lachnosa, 782 Haworthia planifolia, 783 Haworthia planifolia fa. agavoides, 784 Haworthia planifolia fa. alta, 784 Haworthia planifolia fa. calochlora, 784 Haworthia planifolia fa. olivacea, 784 Haworthia planifolia fa. robusta, 784 Haworthia planifolia subvar. agavoides, 784 Haworthia planifolia var. exulata, 784 Haworthia planifolia var. incrassata, 784 Haworthia planifolia var. longifolia, 784 Haworthia planifolia var. poellnitziana, 784 Haworthia planifolia var. setulifera, 785 Haworthia planifolia var. sublaevis, 784

Index Haworthia planifolia var. transiens, 811 Haworthia platinosa, 773 Haworthia plena, 784 Haworthia plettens, 809 Haworthia poellnitziana, 796 Haworthia polyphylla, 773 Haworthia pringlei, 779 Haworthia pringlei var. hastata, 779 Haworthia pseuda, 808 Haworthia pseudogranulata, 773 Haworthia pseudorigida, 772 Haworthia pseudotessellata, 811 Haworthia pseudotortuosa, 813 Haworthia pubescens, 803 Haworthia pubescens var. livida, 793 Haworthia pulchella, 803 Haworthia pulchella var. globifera, 803 Haworthia pulchella var. pulchella, 803 Haworthia pumila, 789 Haworthia pumila ssp. attenuata, 776 Haworthia pumila ssp. fasciata, 787 Haworthia pumila ssp. longiana, 792 Haworthia pumila ssp. minima, 795 Haworthia pumila ssp. radula, 777 Haworthia pumila var. kingiana, 790 Haworthia pumila var. smitii, 809 Haworthia pungens, 804 Haworthia pusilla, 781 Haworthia pygmaea, 804 Haworthia pygmaea fa. crystallina, 805 Haworthia pygmaea fa. major, 805 Haworthia pygmaea var. acuminata, 804 Haworthia pygmaea var. argenteo-maculosa, 804 Haworthia pygmaea var. dekenahii, 804 Haworthia pygmaea var. esterhuizenii, 805 Haworthia pygmaea var. fusca, 804 Haworthia pygmaea var. pygmaea, 804 Haworthia pygmaea var. vincentii, 805 Haworthia quimutica, 807 Haworthia radula, 777 Haworthia radula var. asperior, 773 Haworthia radula var. laevior, 773 Haworthia radula var. magniperlata, 773 Haworthia radula var. pluriperlata, 777 Haworthia ramifera, 794 Haworthia ramosa, 785 Haworthia rava, 776 Haworthia recurva, 812 Haworthia reddii, 795 Haworthia reflexa, 808 Haworthia regalis, 786 Haworthia regens, 775 Haworthia regina, 786 Haworthia reinwardtii, 805 Haworthia reinwardtii fa. brevicula, 806 Haworthia reinwardtii fa. chalumnensis, 805 Haworthia reinwardtii fa. diminuta, 806 Haworthia reinwardtii fa. kaffirdriftensis, 805 Haworthia reinwardtii fa. olivacea, 805

1439 Haworthia reinwardtii fa. zebrina, 805 Haworthia reinwardtii ssp. coarctata, 779 Haworthia reinwardtii ssp. glauca, 788 Haworthia reinwardtii var. adelaidensis, 780 Haworthia reinwardtii var. archibaldiae, 806 Haworthia reinwardtii var. bellula, 780 Haworthia reinwardtii var. brevicula, 805 Haworthia reinwardtii var. chalumnensis, 805 Haworthia reinwardtii var. chalwinii, 780 Haworthia reinwardtii var. coarctata, 779 Haworthia reinwardtii var. committeesensis, 780 Haworthia reinwardtii var. conspicua, 780 Haworthia reinwardtii var. diminuta, 806 Haworthia reinwardtii var. fallax, 780 Haworthia reinwardtii var. glauca, 788 Haworthia reinwardtii var. grandicula, 806 Haworthia reinwardtii var. greenii, 780 Haworthia reinwardtii var. haworthii, 806 Haworthia reinwardtii var. herrei, 788 Haworthia reinwardtii var. huntsdriftensis, 780 Haworthia reinwardtii var. kaffirdriftensis, 805 Haworthia reinwardtii var. major, 806 Haworthia reinwardtii var. minor, 773 Haworthia reinwardtii var. olivacea, 805 Haworthia reinwardtii var. peddiensis, 806 Haworthia reinwardtii var. pseudocoarctata, 780 Haworthia reinwardtii var. pulchra, 806 Haworthia reinwardtii var. reinwardtii, 806 Haworthia reinwardtii var. riebeekensis, 780 Haworthia reinwardtii var. tenuis, 780 Haworthia reinwardtii var. triebneri, 806 Haworthia reinwardtii var. valida, 806 Haworthia reinwardtii var. zebrina, 805 Haworthia resendeana, 773 Haworthia reticulata, 806 Haworthia reticulata ssp. batesiana, 794 Haworthia reticulata var. acuminata, 806 Haworthia reticulata var. attenuata, 806 Haworthia reticulata var. hurlingii, 806 Haworthia reticulata var. integra, 801 Haworthia reticulata var. reticulata, 806 Haworthia reticulata var. subregularis, 807 Haworthia retusa, 807 Haworthia retusa fa. acuminata, 804 Haworthia retusa fa. argenteo-maculosa, 804 Haworthia retusa fa. fouchei, 807 Haworthia retusa fa. geraldii, 807 Haworthia retusa fa. longibracteata, 807 Haworthia retusa fa. multilineata, 807 Haworthia retusa ssp. asperula, 772 Haworthia retusa ssp. emelyae, 786 Haworthia retusa var. acuminata, 804 Haworthia retusa var. asperula, 772 Haworthia retusa var. bruynsii, 779 Haworthia retusa var. comptoniana, 786 Haworthia retusa var. dekenahii, 804 Haworthia retusa var. densiflora, 807 Haworthia retusa var. emelyae, 786 Haworthia retusa var. fouchei, 807

1440 Haworthia retusa var. fusca, 804 Haworthia retusa var. heidelbergensis, 797 Haworthia retusa var. longibracteata, 807 Haworthia retusa var. magnifica, 797 Haworthia retusa var. mirabilis, 796 Haworthia retusa var. multilineata, 807 Haworthia retusa var. mutica, 801 Haworthia retusa var. nigra, 807 Haworthia retusa var. quimutica, 807 Haworthia retusa var. retusa, 807 Haworthia retusa var. solitaria, 807 Haworthia retusa var. springbokvlakensis, 810 Haworthia retusa var. suberecta, 807 Haworthia retusa var. turgida, 808 Haworthia revendettii, 773 Haworthia rigida, 771 Haworthia rigida var. expansa, 771 Haworthia rodinii, 808 Haworthia rooibergensis, 800 Haworthia rooibergensis var. erii, 800 Haworthia rooivleiensis, 808 Haworthia rosea, 784 Haworthia rossouwii, 808 Haworthia rossouwii var. calcarea, 808 Haworthia rossouwii var. elizeae, 808 Haworthia rossouwii var. minor, 808 Haworthia rossouwii var. petrophila, 808 Haworthia rossouwii var. rossouwii, 808 Haworthia rossouwii var. scabra, 799 Haworthia rossouwii var. serrata, 808 Haworthia royalis, 776 Haworthia rubriflora, 828 Haworthia rubriflora var. jacobseniana, 828 Haworthia rubrobrunnea, 773 Haworthia rugosa, 772 Haworthia rugosa var. perviridis, 772 Haworthia rycroftiana, 801 Haworthia ryderiana, 773 Haworthia ryneveldii, 802 Haworthia sabita, 782 Haworthia sabrina, 783 Haworthia sakaii, 801 Haworthia salina, 783 Haworthia salina var. venusta, 783 Haworthia sampaiana, 773 Haworthia sapphaia, 779 Haworthia sarcoidea, 785 Haworthia saundersiae, 513 Haworthia scabra, 809 Haworthia scabra ssp. granulata, 789 Haworthia scabra ssp. sordida, 810 Haworthia scabra ssp. starkiana, 809 Haworthia scabra var. johanii, 809 Haworthia scabra var. lateganiae, 809 Haworthia scabra var. lavrani, 810 Haworthia scabra var. morrisiae, 809 Haworthia scabra var. scabra, 809 Haworthia scabra var. sordida, 810 Haworthia scabra var. starkiana, 809

Index Haworthia scabra var. tuberculata, 809 Haworthia scabrida, 799 Haworthia scabrispina, 775 Haworthia schmidtiana, 802 Haworthia schmidtiana fa. nana, 802 Haworthia schmidtiana var. angustata, 802 Haworthia schmidtiana var. diversifolia, 802 Haworthia schmidtiana var. elongata, 802 Haworthia schmidtiana var. pusilla, 802 Haworthia schmidtiana var. suberecta, 802 Haworthia schoemanii, 789 Haworthia schuldtiana, 797 Haworthia schuldtiana var. erecta, 798 Haworthia schuldtiana var. maculata, 793 Haworthia schuldtiana var. major, 787 Haworthia schuldtiana var. minor, 797 Haworthia schuldtiana var. robertsonensis, 797 Haworthia schuldtiana var. simplicior, 797 Haworthia schuldtiana var. sublaevis, 797 Haworthia schuldtiana var. subtuberculata, 797 Haworthia schuldtiana var. unilineata, 797 Haworthia schuldtiana var. whitesloaneana, 797 Haworthia scottii, 775 Haworthia semiglabrata, 773 Haworthia semimargaritifera, 793 Haworthia semimargaritifera var. major, 772 Haworthia semimargaritifera var. maxima, 793 Haworthia semimargaritifera var. minor, 772 Haworthia semimargaritifera var. multiperlata, 773 Haworthia semiviva, 809 Haworthia serrata, 808 Haworthia sessiliflora, 773 Haworthia setata, 776 Haworthia setata subvar. joubertii, 800 Haworthia setata var. bijliana, 773 Haworthia setata var. gigas, 776 Haworthia setata var. joubertii, 800 Haworthia setata var. major, 776 Haworthia setata var. media, 776 Haworthia setata var. nigricans, 775 Haworthia setata var. subinermis, 773 Haworthia setata var. xiphiophylla, 786 Haworthia setulifera, 785 Haworthia shieldsiana, 703 Haworthia silviae, 804 Haworthia silviae var. nigra, 807 Haworthia silvicola, 781 Haworthia similis, 773 Haworthia simofuri, 799 Haworthia skinneri, 709 Haworthia smitii, 809 Haworthia solitaria, 807 Haworthia sordida, 810 Haworthia sordida var. agavoides, 810 Haworthia sordida var. lavrani, 810 Haworthia sordida var. sordida, 810 Haworthia sparsa, 793 Haworthia speciosa, 778 Haworthia specksii, 778

Index Haworthia sphaeroidea, 810 Haworthia spiralis, 705 Haworthia spirella, 705 Haworthia splendens, 799 Haworthia splendens var. hansii, 797 Haworthia splendens var. ingoi, 799 Haworthia splendens var. masai, 797 Haworthia springbokvlakensis, 810 Haworthia standeri, 800 Haworthia starkiana, 809 Haworthia starkiana var. lateganiae, 809 Haworthia starkiana var. starkiana, 809 Haworthia stayneri, 782 Haworthia stayneri var. salina, 782 Haworthia stenophylla, 513 Haworthia stewarta, 786 Haworthia stiemiei, 773 Haworthia striata, 792 Haworthia studis, 785 Haworthia subattenuata, 772 Haworthia suberecta, 807 Haworthia suberecta var. pallidifolia, 807 Haworthia subfasciata, 772 Haworthia subfasciata var. kingiana, 790 Haworthia subglauca, 779 Haworthia subhamata, 783 Haworthia sublimpidula, 797 Haworthia sublineata, 799 Haworthia submaculata, 790 Haworthia subregularis, 807 Haworthia subretusa, 807 Haworthia subspicata, 662 Haworthia subularis, 781 Haworthia subulata, 772 Haworthia succinea, 785 Haworthia swanea, 783 Haworthia tarkasia, 795 Haworthia tauteae, 810 Haworthia taylorii, 809 Haworthia tenera, 783 Haworthia tenera var. confusa, 773 Haworthia tenera var. cummingii, 783 Haworthia tenera var. doldii, 781 Haworthia tenera var. major, 776 Haworthia tenmari, 785 Haworthia tenuifolia, 513 Haworthia tenuis, 780 Haworthia teres, 782 Haworthia tessellata, 810 Haworthia tessellata fa. brevior, 811 Haworthia tessellata fa. longior, 811 Haworthia tessellata fa. major, 811 Haworthia tessellata var. coriacea, 811 Haworthia tessellata var. elongata, 811 Haworthia tessellata var. engleri, 811 Haworthia tessellata var. inflexa, 810 Haworthia tessellata var. luisieri, 811 Haworthia tessellata var. minutissima, 811 Haworthia tessellata var. obesa, 811

1441 Haworthia tessellata var. palhinhiae, 811 Haworthia tessellata var. parva, 810 Haworthia tessellata var. simplex, 811 Haworthia tessellata var. stepheniana, 811 Haworthia tessellata var. tuberculata, 811 Haworthia tessellata var. velutina, 811 Haworthia tisleyi, 773 Haworthia tomentosa, 781 Haworthia toonensis, 799 Haworthia tooris, 785 Haworthia torquata, 813 Haworthia tortella, 773 Haworthia tortuosa, 813 Haworthia tortuosa var. curta, 773 Haworthia tortuosa var. major, 772 Haworthia tortuosa var. pseudorigida, 772 Haworthia tortuosa var. tortella, 773 Haworthia tradouwensis, 801 Haworthia transiens, 811 Haworthia transiens var. florens, 782 Haworthia translucens, 789 Haworthia translucens ssp. tenera, 783 Haworthia translucens var. delicatula, 773 Haworthia tretyrensis, 776 Haworthia tretyrensis var. pectinis, 776 Haworthia tricolor, 787 Haworthia triebneriana, 798 Haworthia triebneriana var. depauperata, 798 Haworthia triebneriana var. diversicolor, 797 Haworthia triebneriana var. lanceolata, 774 Haworthia triebneriana var. multituberculata, 798 Haworthia triebneriana var. napierensis, 798 Haworthia triebneriana var. nitida, 774 Haworthia triebneriana var. pulchra, 798 Haworthia triebneriana var. rubrodentata, 798 Haworthia triebneriana var. sublineata, 799 Haworthia triebneriana var. subtuberculata, 798 Haworthia triebneriana var. turgida, 798 Haworthia truncata, 811 Haworthia truncata fa. crassa, 812 Haworthia truncata fa. normalis, 812 Haworthia truncata fa. tenuis, 812 Haworthia truncata var. maughanii, 811 Haworthia truncata var. minor, 812 Haworthia truncata var. tenuis, 812 Haworthia truncata var. truncata, 812 Haworthia truterorum, 777 Haworthia tuberculata, 809 Haworthia tuberculata var. acuminata, 809 Haworthia tuberculata var. angustata, 809 Haworthia tuberculata var. subexpansa, 809 Haworthia tuberculata var. sublaevis, 809 Haworthia turcosa, 781 Haworthia turgida, 808 Haworthia turgida fa. caespitosa, 808 Haworthia turgida fa. pallidifolia, 807 Haworthia turgida fa. suberecta, 807 Haworthia turgida var. longibracteata, 807 Haworthia turgida var. pallidifolia, 807

1442 Haworthia turgida var. suberecta, 807 Haworthia turgida var. subtuberculata, 807 Haworthia ubomboensis, 792 Haworthia uitewaaliana, 774, 796 Haworthia umbraticola, 784 Haworthia umbraticola var. hilliana, 784 Haworthia umbraticola var. obesa, 785 Haworthia unicolor, 801 Haworthia unicolor var. helmiae, 773 Haworthia unicolor var. venteri, 775 Haworthia uniondalensis, 777 Haworthia variabilis, 813 Haworthia variegata, 812 Haworthia variegata var. hemicrypta, 812 Haworthia variegata var. modesta, 812 Haworthia variegata var. petrophila, 808 Haworthia variegata var. variegata, 812 Haworthia velutina, 783 Haworthia venetia, 781 Haworthia venetia var. calva, 781 Haworthia venetia var. jeffreis, 781 Haworthia venetia var. teres, 782 Haworthia venosa, 812 Haworthia venosa ssp. granulata, 789 Haworthia venosa ssp. nigra, 801 Haworthia venosa ssp. recurva, 812 Haworthia venosa ssp. tessellata, 810 Haworthia venosa ssp. woolleyi, 814 Haworthia venosa var. oertendahlii, 812 Haworthia venosa var. tessellata, 810 Haworthia venteri, 775 Haworthia venusta, 783 Haworthia vernalis, 798 Haworthia victoria, 810 Haworthia villosa, 778 Haworthia vincentii, 805 Haworthia violacea, 778 Haworthia virella, 785 Haworthia virescens, 794 Haworthia virescens var. minor, 794 Haworthia virginea, 785 Haworthia viridis, 795 Haworthia viridula, 774 Haworthia viscosa, 813 Haworthia viscosa fa. asperiuscula, 813 Haworthia viscosa fa. beanii, 813 Haworthia viscosa fa. pseudotortuosa, 813 Haworthia viscosa fa. subobtusa, 813 Haworthia viscosa fa. torquata, 813 Haworthia viscosa ssp. dereki-clarki, 813 Haworthia viscosa ssp. nigra, 801 Haworthia viscosa var. caespitosa, 813 Haworthia viscosa var. concinna, 813 Haworthia viscosa var. cougaensis, 813 Haworthia viscosa var. indurata, 813 Haworthia viscosa var. major, 813 Haworthia viscosa var. minor, 813 Haworthia viscosa var. parvifolia, 813 Haworthia viscosa var. pseudotortuosa, 813

Index Haworthia viscosa var. quaggaensis, 813 Haworthia viscosa var. subobtusa, 813 Haworthia viscosa var. torquata, 813 Haworthia viscosa var. variabilis, 813 Haworthia viscosa var. viridissima, 813 Haworthia vitris, 774 Haworthia vittata, 781 Haworthia vlokii, 814 Haworthia walmsleyi, 774 Haworthia whitesloaneana, 797 Haworthia willowmorensis, 777, 798 Haworthia wimii, 787 Haworthia wittebergensis, 814 Haworthia woolleyi, 814 Haworthia xiphiophylla, 786 Haworthia yarrowa, 774 Haworthia yocans, 781 Haworthia zantneriana, 814 Haworthia zantneriana var. minor, 814 Haworthia zantneriana var. zantneriana, 814 Haworthia zenigata, 790 Haworthio-gasteria, 749 Haworthiopsis, 769 Haworthiopsis attenuata, 777 Haworthiopsis attenuata var. glabrata, 777 Haworthiopsis attenuata var. radula, 777 Haworthiopsis bruynsii, 779 Haworthiopsis cassytha, 773 Haworthiopsis coarctata, 779 Haworthiopsis coarctata var. adelaidensis, 780 Haworthiopsis coarctata var. tenuis, 780 Haworthiopsis curta, 773 Haworthiopsis expansa, 771 Haworthiopsis fasciata, 787 Haworthiopsis fasciata var. browniana, 787 Haworthiopsis glauca, 788 Haworthiopsis glauca var. herrei, 788 Haworthiopsis granulata, 789 Haworthiopsis granulata var. schoemanii, 789 Haworthiopsis hybrida, 772 Haworthiopsis koelmaniorum, 791 Haworthiopsis koelmaniorum var. mcmurtryi, 791 Haworthiopsis limifolia, 791 Haworthiopsis limifolia var. arcana, 791 Haworthiopsis limifolia var. gigantea, 791 Haworthiopsis limifolia var. glaucophylla, 792 Haworthiopsis limifolia var. striata, 792 Haworthiopsis limifolia var. ubomboensis, 792 Haworthiopsis lisbonensis, 773 Haworthiopsis longiana, 792 Haworthiopsis major, 772 Haworthiopsis mcmurtryi, 791 Haworthiopsis nigra, 801 Haworthiopsis nigra var. diversifolia, 802 Haworthiopsis nigra var. elongata, 802 Haworthiopsis pseudorigida, 772 Haworthiopsis pungens, 804 Haworthiopsis reinwardtii, 805 Haworthiopsis reinwardtii fa. chalumnensis, 805

Index Haworthiopsis reinwardtii fa. kaffirdriftensis, 805 Haworthiopsis reinwardtii fa. olivacea, 805 Haworthiopsis reinwardtii var. brevicula, 806 Haworthiopsis reinwardtii var. coarctata, 779 Haworthiopsis reinwardtii var. greenii, 780 Haworthiopsis reinwardtii var. olivacea, 805 Haworthiopsis reinwardtii var. tenuis, 780 Haworthiopsis resendeana, 773 Haworthiopsis revendettii, 773 Haworthiopsis rigida, 771 Haworthiopsis rubrobrunnea, 773 Haworthiopsis sampaiana, 773 Haworthiopsis scabra, 809 Haworthiopsis scabra var. johanii, 809 Haworthiopsis scabra var. lateganiae, 809 Haworthiopsis scabra var. morrisiae, 809 Haworthiopsis scabra var. smitii, 809 Haworthiopsis scabra var. starkiana, 809 Haworthiopsis sordida, 810 Haworthiopsis sordida var. agavoides, 810 Haworthiopsis sordida var. lavrani, 810 Haworthiopsis subrigida, 772 Haworthiopsis tauteae, 810 Haworthiopsis tessellata, 810 Haworthiopsis tessellata var. crausii, 811 Haworthiopsis tessellata var. crousii, 811 Haworthiopsis tortella, 773 Haworthiopsis tortuosa, 813 Haworthiopsis variabilis, 813 Haworthiopsis venosa, 812 Haworthiopsis venosa var. granulata, 789 Haworthiopsis venosa var. woolleyi, 814 Haworthiopsis viscosa, 813 Haworthiopsis viscosa var. asperiuscula, 813 Haworthiopsis viscosa var. beanii, 813 Haworthiopsis viscosa var. variabilis, 813 Haworthiopsis woolleyi, 814 Hecatris, 467 Hecatris asparagoides, 468 Hechcohnia, 995 Hechtia, 985, 997 Hechtia aquamarina, 998 Hechtia argentea, 1007 Hechtia besseriana, 1028 Hechtia bracteata, 998 Hechtia caerulea, 1030 Hechtia capituligera, 1007 Hechtia carlsoniae, 999 Hechtia carnea, 843 Hechtia caudata, 999 Hechtia caulescens, 1000 Hechtia chichinautzensis, 1000 Hechtia colossa, 1000 Hechtia complanata, 1001 Hechtia confusa, 1001 Hechtia conzattiana, 1002 Hechtia cordylinoides, 998 Hechtia deceptrix, 1002 Hechtia desmetiana, 1025

1443 Hechtia dichroantha, 1003 Hechtia edulis, 1003 Hechtia ellemeetii, 1042 Hechtia elliptica, 1004 Hechtia epigyna, 963, 1004 Hechtia flexilifolia, 1004 Hechtia fosteriana, 1005 Hechtia fragilis, 1005 Hechtia galeottii, 1005 Hechtia gamopetala, 1007 Hechtia gayii, 1018 Hechtia gayorum, 1018 Hechtia ghiesbreghtiana, 1007 Hechtia ghiesbreghtii, 1007 Hechtia glabra, 1028 Hechtia glauca, 1006 Hechtia glomerata, 995, 1006 Hechtia gracilis, 843 Hechtia guatemalensis, 1007 Hechtia hernandez-sandovalii, 1008 Hechtia hintoniana, 1008 Hechtia huamelulaensis, 1009 Hechtia iltisii, 1009 Hechtia integerrima, 1030 Hechtia isthmusiana, 1010 Hechtia ixtlanensis, 1010 Hechtia jaliscana, 1011 Hechtia joinvillei, 843, 986 Hechtia laevis, 1011 Hechtia lanata, 1012 Hechtia laxissima, 1013 Hechtia lepidophylla, 1007 Hechtia liebmannii, 1013 Hechtia lindmanioides, 1023 Hechtia longifolia, 998 Hechtia lundelliorum, 1013 Hechtia lyman-smithii, 1014 Hechtia macdougallii, 1025 Hechtia macrophylla, 1019 Hechtia malvernii, 1014 Hechtia mapimiana, 1015 Hechtia marnier-lapostollei, 1016 Hechtia matudae, 1016 Hechtia melanocarpa, 1017 Hechtia mexicana, 1017 Hechtia meziana, 1025 Hechtia michoacana, 1024 Hechtia montana, 1018 Hechtia montana cv. Burgundy, 1018 Hechtia montis-frigidi, 1018 Hechtia mooreana, 1019 Hechtia morreniana, 1007 Hechtia myriantha, 1019 Hechtia nivea, 1020 Hechtia nuusaviorum, 1020 Hechtia oaxacana, 1020 Hechtia pedicellata, 1021 Hechtia perotensis, 1013 Hechtia pitcairniifolia, 1043

1444 Hechtia podantha, 1021 Hechtia pretiosa, 1022 Hechtia pringlei, 1022 Hechtia pueblensis, 998, 1029 Hechtia pumila, 1023 Hechtia purhepecha, 1023 Hechtia purpusii, 1023 Hechtia reflexa, 1024 Hechtia reticulata, 1025 Hechtia roezlii, 1025 Hechtia rosea, 983, 1025 Hechtia roseana, 1026 Hechtia rubicunda, 1027 Hechtia santanae, 1027 Hechtia scariosa, 1030 Hechtia schottii, 1027 Hechtia sphaeroblasta, 1028 Hechtia stenopetala, 1028 Hechtia striata, 998 Hechtia suaveolens, 998 Hechtia subalata, 1029 Hechtia tehuacana, 1029 Hechtia texensis, 1030 Hechtia tillandsioides, 1030 Hechtia zacatecae, 1030 Hechtia zamudioi, 1031 Helicodea, 859 Helicodea leopoldii, 860 Helmia, 1201 Hemerocallidaceae, 1389 Heminema, 1189 Heminema multiflora, 1190 Hepetis consimilis, 1083 Hepetis ferruginea, 1083 Hepetis lorentziana, 881 Herreria, 11 Herreriaceae, 11 Hesperaloe, 12, 14, 349 Hesperaloe bechtoldii, 354 Hesperaloe campanulata, 350–351 Hesperaloe chiangii, 351 Hesperaloe davyi, 352 Hesperaloe engelmannii, 351–352 Hesperaloe funifera, 352–353 Hesperaloe funifera ssp. chiangii, 351 Hesperaloe malacophylla, 353 Hesperaloe nocturna, 353 Hesperaloe parviflora, 354 Hesperaloe parviflora ssp. bechtoldii, 354 Hesperaloe parviflora ssp. parviflora, 354–355 Hesperaloe parviflora var. engelmannii, 351 Hesperaloe tenuifolia, 355 Hesperaloe yuccifolia, 354 Hesperaloe yuccoides, 354 Hesperocallidaceae, 9, 10 Hesperocallis, 10, 13 Hesperoyucca, 12, 14, 357 Hesperoyucca newberryi, 358 Hesperoyucca peninsularis, 359

Index Hesperoyucca whipplei, 358–360 Hesperoyucca whipplei var. graminifolia, 358 Higher Asparagales, 2 Higinbothamia, 1201 Himas, 1265 Hippeastrum calyptratum, 426 Hohenbergia distichantha, 853 Hohenbergia legrelliana, 855 Hohenbergia nudicaulis, 854 Hohenbergia strobilacea, 850 Hoiriri, 851 Hoiriri polystachia, 853 Hollia comosa, 452 Honorius, 1281 Hoplocryptanthus, 865 Hoplocryptanthus caracensis, 868 Hoplocryptanthus glaziovii, 869 Hoplocryptanthus schwackeanus, 871 Hoplophytum, 851 Hoplophytum distichanthum, 853 Hoplophytum nudicaule, 854 Hosta, 10, 12, 14 Hostaceae, 9, 10 Hyacinthaceae, 1225 Hyacinthus brevifolius, 1248 Hyacinthus convallarioides, 1253 Hyacinthus cryptopodus, 1254 Hyacinthus medius, 1260 I Idothea, 1249 Idothea altissima, 1251 Idothea barteri, 1251 Idothea ludwigii, 1270 Idothea media, 1260 Idothearia, 1249 Igidia, 1231 Indurgia, 1250 Iuka, 363 J Jodrellia, 713 Jodrellia fistulosa, 721 Jodrellia macrocarpa, 729 Jodrellia migiurtina, 729 Jonghea, 859 K Kingia, 1389 Kingiacaceae, 1389 Kniphofia, 819 Kniphofia typhoides, 820–821 Kumalista, 486 Kumalista corderoyi, 538 Kumara, 485 Kumara disticha, 636

Index Kumara haemanthifolia, 571 Kumara plicatilis, 636 L Lachenalia, 1265, 1266 Lachenalia albida, 1266 Lachenalia aloides, 1266 Lachenalia angustifolia, 1266 Lachenalia contaminata, 1266 Lachenalia fragrans, 1266 Lachenalia graminifolia, 1248 Lachenalia hyacinthina, 1266 Lachenalia hyacinthoides, 1266 Lachenalia moniliformis, 1266 Lachenalia patula, 1266 Lachenalia polypodantha, 1267 Lachenalia polypodantha ssp. eburnea, 1267 Lachenalia polypodantha ssp. polypodantha, 1267 Lachenalia succulenta, 1266 Lachenaliaceae, 1225 Lamprococcus, 851 Lamprodithyros, 1171 Lapanthus, 866, 1052 Lasiaceae, 457 Laxmanniaceae, 1293 Ledebouria, 1269 Ledebouria concolor, 1270 Ledebouria corrugata, 1270 Ledebouria cremnophila, 1270 Ledebouria cryptopoda, 1254 Ledebouria ensifolia, 1270 Ledebouria loedolffiae, 1270 Ledebouria megaphylla, 1271 Ledebouria ovatifolia, 1271 Ledebouria ovatifolia ssp. ovatifolia, 1271 Ledebouria ovatifolia ssp. scabrida, 1271 Ledebouria socialis, 1271 Ledebouria venteri, 1272 Ledebouria violacea, 1272 Ledurgia, 1250 Leichtlinia, 22 Leichtlinia commutata, 128 Leichtlinia protuberans, 128 Lemeea, 486 Lemeea boiteaui, 514 Lemeea haworthioides, 573 Lemeea parvula, 628 Lemnaceae, 457 Lepicaulon, 829 Leptaloe, 486 Leptaloe albida, 496 Leptaloe blyderivierensis, 611 Leptaloe minima, 611 Leptaloe myriacantha, 618 Leptaloe saundersiae, 651 Leptocallisia, 1173 Leptocallisia multiflora, 1175 Leptorhoeo, 1189

1445 Leptorhoeo floribunda, 1191 Leucodesmis, 441 Leucojaceae, 425 Liliaceae, 1 Limodoraceae, 1299 Lindmania albicans, 990 Lindmania brevifolia, 880 Lindmania chlorantha, 881 Lindmania penduliflora, 991 Lindmania rusbyi, 992 Lindmania weddelliana, 993 Liparidaceae, 1299 Liriothamnus, 829 Liriothamnus adamsonii, 830 Liriothamnus involucratus, 831 Litanthus, 1250 Litanthus pusillus, 1263 Littaea, 21 Littaea geminiflora, 116 Littaea gracilis, 1330 Littaea juncea, 116 Littaea roezlii, 230 Lomandraceae, 1293 Lomataloe, 486 Lomateria, 745 Lomateria gloriosa, 746 Lomatoloba, 697 Lomatophyllum, 485 Lomatophyllum aldabrense, 497 Lomatophyllum aloiflorum, 642 Lomatophyllum anivoranoense, 501, 502 Lomatophyllum antsingyense, 503 Lomatophyllum belavenokense, 511 Lomatophyllum borbonicum, 642 Lomatophyllum citreum, 533 Lomatophyllum lomatophylloides, 598 Lomatophyllum macrum, 600 Lomatophyllum marginatum, 642 Lomatophyllum namorokaense, 619 Lomatophyllum occidentale, 624 Lomatophyllum oligophyllum, 624 Lomatophyllum orientale, 625 Lomatophyllum pembanum, 630 Lomatophyllum peyrierasii, 634 Lomatophyllum propaguliferum, 640 Lomatophyllum prostratum, 640 Lomatophyllum purpureum, 642 Lomatophyllum roseum, 647 Lomatophyllum rufocinctum, 642 Lomatophyllum sociale, 657 Lomatophyllum tormentorii, 668 Lomatophyllum viviparum, 652 Loncomelos, 1281 Lower Asparagales, 1 M Macrochordion, 851 Macrochordium, 851

1446 Macrochordium recurvatum, 855 Madvigia, 865 Manfreda, 12, 22 Manfreda alibertii, 271 Manfreda angustifolia, 96 Manfreda brachystachys, 228 Manfreda brunnea, 73 Manfreda bulbulifera, 74 Manfreda chamelensis, 82 Manfreda conduplicata, 271 Manfreda elongata, 123 Manfreda fusca, 113 Manfreda galvaniae, 114 Manfreda guerrerensis, 126 Manfreda guttata, 128 Manfreda hauniensis, 131 Manfreda huetamensis, 259 Manfreda insignis, 131 Manfreda involuta, 141 Manfreda jaliscana, 144 Manfreda justosierrana, 145 Manfreda littoralis, 154 Manfreda longibracteata, 155 Manfreda longiflora, 155 Manfreda longistaminata, 239 Manfreda maculata, 244 Manfreda maculosa, 161 Manfreda malinaltenangensis, 228, 229 Manfreda occidentalis, 187 Manfreda oliveriana, 228 Manfreda paniculata, 194 Manfreda parva, 200, 201 Manfreda petskinil, 207, 208 Manfreda planifolia, 209 Manfreda potosina, 213 Manfreda pringlei, 96 Manfreda pubescens, 217 Manfreda revoluta, 220 Manfreda riosramirii, 129, 210 Manfreda rubescens, 215 Manfreda santanamichelii, 227 Manfreda scabra, 228 Manfreda sessiliflora, 228 Manfreda sileri, 238 Manfreda singuliflora, 239 Manfreda tamazunchalensis, 264 Manfreda tigrina, 271 Manfreda tlatlayensis, 131 Manfreda umbrophila, 259 Manfreda undulata, 260 Manfreda valsequillana, 129 Manfreda variegata, 264 Manfreda variegata var. sileri, 238, 239 Manfreda verhoekiae, 266, 267 Manfreda virginica, 271 Manfreda virginica ssp. lata, 271 Manfreda virginica var. tigrina, 271 Manfreda xilitlensis, 264 Mangave, 12, 22

Index Mangave gonzaloi, 122 Massonia, 1273 Massonia bifolia, 1274 Massonia brachypus, 1274 Massonia depressa, 1274 Massonia etesionamibensis, 1275 Massonia grandiflora, 1274 Massonia latifolia, 1274 Massonia namaquensis, 1274 Massonia obovata, 1274 Massonia sanguinea, 1274 Massonia triflora, 1274 Maysara, 823 Medeola angustifolia, 468 Medeola asparagoides, 468 Melanthium massoniifolium, 1274 Melicho, 441 Melomphis, 1281 Merione, 1201 Merwilla, 1277 Merwilla dracomontana, 1277 Merwilla kraussii, 1278 Merwilla natalensis, 1278 Merwilla plumbea, 1278, 1289 Meziothamnus, 879 Meziothamnus brevifolius, 880 Misandra, 1107 Monocotyledons, 1 Monoestes, 1265 Monotassa, 1231 Monsteraceae, 457 Mucinaea, 1250 Myanthe, 1282 Myogalum, 1281 Myrsiphyllum, 467 Myrsiphyllum angustifolium, 468 Myrsiphyllum asparagoides, 468 Myrsiphyllum falciforme, 468 Myrsiphyllum juniperoides, 469 Myrsiphyllum multituberosum, 470 Myrsiphyllum ovatum, 470 Myrsiphyllum striatum, 471 Myrsiphyllum undulatum, 471 N Namophila, 1274, 1279 Namophila urotepala, 1279 Nanarepenta, 1201 Narcissaceae, 425 Navia, 840 Navia brevifolia, 880 Navia splendens, 840 Nemampsis, 1337 Nemopogon, 713 Nemopogon glaucum, 723 Neobakeria, 1273 Neobergia, 860, 1033 Neodonnellia, 1189

Index Neodonnellia grandiflora, 1190 Neoglaziovia, 1035, 1102 Neoglaziovia burle-marxii, 1035 Neoglaziovia concolor, 1035 Neoglaziovia variegata, 1036, 1047 Neopatersonia, 1282 Neophytum, 1039, 1102 Neophytum lymanii, 1039 Neoregelia, 867, 1102 Neottiaceae, 1299 Neuwiediaceae, 1299 New Zealand Cabbage tree, 1296 Nicipe, 1281 Nicipe britteniae, 1283 Nicipe capillifolia, 1284 Nicipe comptonii, 1283 Nicipe filifolia, 1287 Nicipe juncifolia, 1284 Nicipe lithopsoides, 1284 Nicipe perdurans, 1286 Nicipe petraea, 1284 Nicipe sardienii, 1286 Nicipe tortuosa, 1287 Nidularium, 1053 Nidularium caeruleum, 843 Nidularium hydrophorum, 853 Nidumea, 852 Niveophyllum, 997 Niveophyllum caeruleum, 1030 Nolina, 1345 Nolina altamiranoana, 1350 Nolina azureogladiata, 1346 Nolina beldingii [?] deserticola, 1346 Nolina beldingii var. deserticola, 1346 Nolina bigelovii, 1346, 1349 Nolina bigelovii ssp. parryi, 1349 Nolina bigelovii ssp. wolfii, 1349 Nolina bigelovii var. parryi, 1349 Nolina bigelovii var. wolfii, 1349 Nolina cismontana, 1347 Nolina elegans, 1350 Nolina erumpens [?] compacta, 1351 Nolina erumpens var. compacta, 1351 Nolina excelsa, 1347 Nolina gracilis, 1321 Nolina guatemalensis, 1322 Nolina hibernica, 1347–1348 Nolina histrix, 1321 Nolina hookeri, 1323 Nolina interrata, 1348 Nolina longifolia, 1350 Nolina matapensis, 1348 Nolina nelsonii, 1349 Nolina parryi, 1349 Nolina parryi ssp. parryi, 1349 Nolina parryi ssp. wolfii, 1349 Nolina parviflora, 1349 Nolina pliabilis, 1324 Nolina pollyjeanneae, 1350

1447 Nolina recurvata, 1325 Nolina stricta, 1326 Nolina texana, 1351 Nolina texana var. compacta, 1351 Nolina tuberculata, 1325 Nolina wolfii, 1349 Nolinaceae, 1315 Nolinoideae, 1317, 1317–1318 Notosceptrum, 819 Notosceptrum alooides, 498 Notosceptrum natalense, 820 O Obsitila, 829 Ochagavia, 1041 Ochagavia andina, 1042 Ochagavia carnea, 1042 Ochagavia chamissonis, 1042 Ochagavia elegans, 1042 Ochagavia grandiflora, 1042 Ochagavia leiboldiana, 1042 Ochagavia lindleyana, 1043 Ochagavia litoralis, 1042–1043 Oechmea, 851 Oncorhiza, 1201 Oncus, 1201 Ophiobostryx, 1243 Ophiobostryx volubilis, 1244 Ophiopogonaceae, 1315 Ophrydaceae, 1299 Oporanthaceae, 425 Orchidaceae, 1, 1299–1302 Orchiops, 1265 Ornithogalaceae, 1225 Ornithogalum, 1281 Ornithogalum altissimum, 1251 Ornithogalum anomalum, 1252 Ornithogalum aurantiacum, 1286 Ornithogalum battenianum, 1233 Ornithogalum bracteatum, 1233 Ornithogalum brevifolium, 1284 Ornithogalum britteniae, 1283 Ornithogalum capillifolia, 1284 Ornithogalum capillifolium, 1284 Ornithogalum caudatum, 1233 Ornithogalum cepaceum, 718 Ornithogalum circinatum, 1239 Ornithogalum clanwilliamae-gloria, 1234 Ornithogalum comptonii, 1283 Ornithogalum comptum, 1287 Ornithogalum cremnophilum, 1235 Ornithogalum crudenii, 1235 Ornithogalum diluculum, 1236 Ornithogalum epigeum, 1284 Ornithogalum geniculatum, 1283 Ornithogalum giganteum, 1251 Ornithogalum gildenhuysii, 1236 Ornithogalum griseum, 1284

1448 Ornithogalum hyacinthoides, 1248 Ornithogalum imbricatum, 1237 Ornithogalum insigne, 1285 Ornithogalum juncifolium, 1284 Ornithogalum juncifolium var. emmsii, 1284 Ornithogalum juncifolium var. juncifolium, 1284 Ornithogalum karachabpoortense, 1237 Ornithogalum kirstenii, 1237 Ornithogalum langebergense, 1284 Ornithogalum lebaense, 1238 Ornithogalum leptophyllum, 1284 Ornithogalum limosum, 1284 Ornithogalum lithopsoides, 1284 Ornithogalum longibracteatum, 1233 Ornithogalum maculatum, 1284 Ornithogalum maculatum var. speciosum, 1284 Ornithogalum maculatum var. splendens, 1285 Ornithogalum magnificum, 1285 Ornithogalum maritimum, 1259 Ornithogalum massonii, 1233 Ornithogalum multifolium, 1286 Ornithogalum naviculum, 1285 Ornithogalum nelsonii, 1238 Ornithogalum nematophyllum, 1240 Ornithogalum notatum, 1284 Ornithogalum oliganthum, 1284 Ornithogalum paradoxum, 1220 Ornithogalum pendens, 1285 Ornithogalum pendulum, 1238 Ornithogalum perdurans, 1285 Ornithogalum perpulchrum, 1286 Ornithogalum petraeum, 1284 Ornithogalum ranunculoides, 1286 Ornithogalum richtersveldensis, 1286 Ornithogalum roussouwii, 1285 Ornithogalum rupestre, 1286 Ornithogalum sardienii, 1286 Ornithogalum scabrocostatum, 1239 Ornithogalum scilloides, 1233 Ornithogalum setifolium, 1284 Ornithogalum shawii, 1239 Ornithogalum speciosum, 1284 Ornithogalum splendens, 1285 Ornithogalum stapfii, 1236 Ornithogalum stenostachyum, 1284 Ornithogalum subulatum, 1284 Ornithogalum tenuipes, 1284 Ornithogalum thermarum, 1240 Ornithogalum thunbergianulum, 1287 Ornithogalum thunbergianum, 1284 Ornithogalum thunbergianum var. concolor, 1284 Ornithogalum tortuosum, 1287 Ornithogalum tuberosum, 718 Ornithogalum unifoliatum, 1240 Ornithogalum volutare, 1239 Orontiaceae, 457 Ortgiesia, 851 Ortgiesia cylindrata, 853 Ortgiesia gracilis, 854

Index Ortgiesia legrelliana, 855 Ortgiesia palleolata, 856 Ortgiesia recurvata, 855 Ortgiesia recurvata var. benrathii, 855 Ortgiesia recurvata var. ortgiesii, 855 Ortgiesia tillandsioides, 855 Ortgiesia tillandsioides var. subexserta, 855 Orthocohnia, 1045 Orthoglaziovia, 1047, 1102 Ortholarium, 1053 Orthomea, 1049, 1053, 1102 Orthophytum, 852, 1051 Orthophytum albopictum, 1047, 1102 Orthophytum argentum, 1053–1054 Orthophytum braunii, 1054 Orthophytum burle-marxii, 1103 Orthophytum burle-marxii var. seabrae, 1103 Orthophytum diamantinense, 1054–1055 Orthophytum disjunctum, 1062 Orthophytum eddie-estevesii, 1055 Orthophytum estevesii, 1045, 1061 Orthophytum graomogolense, 1055, 1058 Orthophytum gurkenii, 1062 Orthophytum harleyi, 1055–1056 Orthophytum hatschbachii, 1104 Orthophytum jacaraciense, 1056–1057 Orthophytum lemei, 1057 Orthophytum leprosum, 1062 Orthophytum magalhaesii, 1062 Orthophytum maracasense, 1057, 1059 Orthophytum mello-barretoi, 1057–1058 Orthophytum navioides, 1049, 1067, 1104 Orthophytum piranianum, 1058 Orthophytum pseudovagans, 1064 Orthophytum riocontense, 1058–1059 Orthophytum roseum, 1103 Orthophytum rubrum, 1059 Orthophytum saxicola, 1056 Orthophytum saxicola ssp. rubra, 1067 Orthophytum schulzianum, 1059–1060 Orthophytum striatifolium, 1060–1061 Orthophytum toscanoi, 1061–1062 Orthophytum toscanoi ssp. atropurpureum, 1053 Orthophytum triunfense, 1062 Orthophytum vagans, 1049, 1064 Orthophytum vasconcelosianum, 1062–1063 Orthophytum zanonii, 1063–1064 Orthotanthus, 1067, 1102 Osmyne, 1231 P Pachidendron, 486 Pachidendron africanum, 495 Pachidendron africanum var. angustum, 495 Pachidendron africanum var. latum, 495 Pachidendron angustifolium, 495 Pachidendron ferox, 557 Pachidendron pseudoferox, 557

Index Pachidendron supralaeve, 557 Pachydendron, 486 Pallastema, 1231 Pancratiaceae, 425 Papilista, 751 Peliosanthaceae, 1315 Pelourdea, 366 Periboea, 1265 Periboea brevifolia, 1248 Perihema, 441 Peripetasma, 1201 Perosanthera, 1171 Phaedranassa megistophylla, 445 Phaedranassa multiflora, 445 Phaeocles, 1282 Phalangium asphodeloides, 716 Phalangium bulbosum, 717 Phalangium canaliculatum, 830 Phalangium comosum, 452 Phalangium longifolium, 830 Phalangium semibarbatum, 734 Phalangium tenuifolium, 741 Phalangium vespertinum, 830 Phalangium viviparum, 452 Philodendraceae, 457 Philodendron warszewiczii, 458 Pholidophyllum, 865 Phylloglottis, 1219 Phylloma, 486 Phylloma aloiflorum, 642 Phylloma macrum, 600 Phylloma rufocinctum, 642 Phyodina, 1173 Phyodina micrantha, 1174 Phyodina navicularis, 1175 Phyodina warszewicziana, 1176 Physodia, 1249 Phytarrhiza, 1107 Phytarrhiza circinalis, 1125 Phytarrhiza crocata, 1121 Phytarrhiza duratii, 1125 Phytarrhiza xiphioides, 1160 Pilasia, 1249 Pincenectitia, 1319 Pincenectitia glauca, 1326 Pincenectitia gracilis, 1321 Pincenectitia tuberculata, 1325 Pistiaceae, 457 Pitcairnia, 840 Pitcairnia alpestris, 1076 Pitcairnia andreana, 840 Pitcairnia asterotricha, 1083 Pitcairnia balsaminea, 840 Pitcairnia brevifolia, 880 Pitcairnia burle-marxii, 840 Pitcairnia chilensis, 1079 Pitcairnia chlorantha, 881 Pitcairnia chrysantha, 882 Pitcairnia coarctata, 1079

1449 Pitcairnia coerulea, 1080 Pitcairnia consimilis, 1083 Pitcairnia cotahuasiana, 1083 Pitcairnia cristalinensis, 840 Pitcairnia dyckioides, 1082 Pitcairnia echinotricha, 1083 Pitcairnia encholirioides, 840 Pitcairnia ferruginea, 1083 Pitcairnia grandiflora, 1092 Pitcairnia herrerae, 1083 Pitcairnia imperialis, 1083 Pitcairnia lanata, 1088 Pitcairnia laresiana, 1083 Pitcairnia latibracteata, 1083 Pitcairnia lorentziana, 881 Pitcairnia megastachya, 1094 Pitcairnia mirabilis, 1090 Pitcairnia mirabilis var. tucumana, 1090 Pitcairnia philippii, 1081 Pitcairnia sphaerocephala, 1097 Pitcairnia venusta, 1097 Pitcairnia violacea, 1081 Pitcairnia viridis, 1083 Pitcairnia volker-schaedlichii, 841 Pitcairnia weberbaueri, 1083 Pitcairnia weddelliana, 1098 Pityrophyllum, 1107 Pityrophyllum erubescens, 1135 Placseptalia, 1041 Placseptalia rebecae, 1043 Platyaechmea, 851 Platyaechmea distichantha, 853 Platyaechmea distichantha var. glaziovii, 854 Platyaechmea distichantha var. schlumbergeri, 854 Platyaechmea distichantha var. vernicosa, 854 Platyestes, 1265 Platystachys, 1107 Platystachys bulbosa, 1114 Platystachys ehrenbergii, 1157, 1158 Platystachys erythraea, 1114 Platystachys heptantha, 844 Platystachys inanis, 1116 Platystachys kunthiana, 1138 Platystachys latifolia, 1138 Platystachys pruinosa, 1149 Platystachys tortilis, 1158 Pleomele, 1337 Pleomele aloifolia, 1367 Pleomele hookeriana, 1338 Poaceae, 1307 Podaechmea, 851 Poellneria, 825 Poellnitzia, 827 Poellnitzia rubriflora, 827 Poellnitzia rubriflora var. jacobseniana, 828 Polea telegraphica, 250 Polemannia, 1247 Polemannia hyacinthoides, 1248 Polianthes, 12, 14, 21

1450 Polianthes alboaustralis, 43 Polianthes americana, 86 Polianthes bicolor, 68 Polianthes blissii, 68 Polianthes brachystachys, 228 Polianthes brunnea, 73 Polianthes bundrantii, 76 Polianthes cernua, 180 Polianthes debilis, 96 Polianthes densiflora, 91 Polianthes durangensis, 55 Polianthes elongata, 123, 216, 217 Polianthes ensifolia, 40 Polianthes geminiflora, 86 Polianthes geminiflora ssp. clivicola, 87 Polianthes geminiflora ssp. pueblensis, 88 Polianthes geminiflora var. clivicola, 87 Polianthes geminiflora var. graminifolia, 124 Polianthes geminiflora var. pueblensis, 88 Polianthes gracilis, 48 Polianthes graminifolia, 124 Polianthes guttata, 128 Polianthes howardii, 136 Polianthes jaliscana, 144 Polianthes lata, 271 Polianthes longiflora, 103 Polianthes maculata, 244 Polianthes maculosa, 161 Polianthes mexicana, 75 Polianthes michoacana, 103 Polianthes montana, 221 Polianthes multicolor, 176 Polianthes nelsonii, 180, 181 Polianthes oaxacana, 184 Polianthes oliveriana, 228 Polianthes palustris, 193, 194 Polianthes planifolia, 209, 210 Polianthes platyphylla, 210 Polianthes potosina, 213 Polianthes pringlei, 55 Polianthes purpurea, 40 Polianthes pygmaea, 40 Polianthes quilae, 218, 219 Polianthes revoluta, 220 Polianthes rosei, 123 Polianthes rubescens, 215 Polianthes runyonii, 155 Polianthes sessiliflora, 55 Polianthes singuliflora, 239, 240 Polianthes tuberosa, 48 Polianthes tuberosa fa. plena, 48 Polianthes tuberosa var. gracilis, 48 Polianthes tubulata, 86 Polianthes variegata, 264 Polianthes venustuliflora, 265 Polianthes virginica, 271 Polianthes virginica fa. tigrina, 271 Polianthes zapopanensis, 281 Polifreda, 22

Index Polygonataceae, 1315 Polynome, 1201 Polyxena, 1265 Portea legrelliana, 855 Portea leptantha, 841 Portea tillandsioides, 855 Portemea, 852 Pothaceae, 457 Pothos, 21 Pothuava, 851 Pothuava nudicaulis, 854 Pothuava weberbaueri, 856 Pourretia, 1073 Pourretia africana, 844 Pourretia alpestris, 1075 Pourretia argentea, 1042 Pourretia auruacensis, 1074 Pourretia coarctata, 1079 Pourretia coerulea, 1081 Pourretia ferruginea, 1083 Pourretia flexilis, 844 Pourretia gigantea, 1093 Pourretia hanisiana, 1114 Pourretia joinvillei, 986 Pourretia lanata, 1088 Pourretia lepidota, 1074 Pourretia mexicana, 844 Pourretia nivosa, 1156 Pourretia rubricaulis, 1081 Pourretia speciosa, 1074 Pourretia stricta, 1135 Pourretia violacea, 1081 Pourretia yuccoides, 1074 Prantleia, 1051 Prionophyllum, 893 Prionophyllum maritimum, 930 Prionophyllum selloum, 950 Prochnyanthes, 12, 22 Prochnyanthes bulliana, 75 Prochnyanthes mexicana, 75 Prochnyanthes viridescens, 75 Protasparagus, 467 Protasparagus crassicladus, 469 Protasparagus densiflorus, 469 Protasparagus mucronatus, 470 Protasparagus oxyacanthus, 470 Protasparagus striatus, 471 Protoyucca sadishii, 365 Pseudoanamea, 852 Pseudobravoa, 22 Pseudobravoa densiflora, 91 Pseudogaltonia, 1247–1248 Ptyas, 486 Puckia, 1069 Pucohnia, 1071 Puya, 1073 Puya acris, 1074 Puya adscendens, 1074 Puya aequatorialis, 1075

Index Puya aequatorialis fa. albiflora, 1075 Puya aequatorialis var. aequatorialis, 1075 Puya aequatorialis var. albiflora, 1075 Puya alpestris, 1075 Puya alpestris ssp. alpestris, 1076 Puya alpestris ssp. zoellneri, 1076 Puya asplundii, 1076 Puya berteroniana, 1076 Puya boliviensis, 1077 Puya bravoi, 1077 Puya butcheriana, 1087 Puya cardenasii, 1077 Puya castellanosii, 1078 Puya cerrateana, 1078 Puya chilensis, 1079 Puya chilensis var. gigantea, 1079 Puya claudiae, 1080 Puya coarctata, 1079 Puya cochabambensis, 1080 Puya coerulea, 1080 Puya coerulea var. coerulea, 1081 Puya coerulea var. intermedia, 1081 Puya coerulea var. monteroana, 1081 Puya coerulea var. violacea, 1081 Puya copiapina, 1077 Puya coquimbensis, 1097 Puya densiflora, 1081 Puya dyckioides, 1082 Puya echinotricha, 1083 Puya edulis, 895 Puya eryngioides, 1082 Puya ferruginea, 1082 Puya flava, 884 Puya flora, 1098 Puya gaudichaudii, 1097 Puya gigantea, 1079 Puya gilmartiniae, 1083 Puya glabrata, 1081 Puya glomerifera, 1084 Puya grafii, 1084 Puya grandiflora, 1083 Puya gummifera, 1096 Puya gutteana, 1085 Puya harmsii, 1085 Puya harry-lutheri, 1086 Puya hauthalii, 1086 Puya hofstenii, 1086 Puya hoxeyi, 1087 Puya hromadnikii, 1097 Puya humilis, 1087 Puya ibischii, 1087 Puya lanata, 1088 Puya laxa, 1069, 1071, 1088 Puya lopezii, 1089 Puya macrura, 1089 Puya medica, 1090 Puya mirabilis, 1090 Puya mitis, 1091 Puya navarroana, 1091

1451 Puya nigrescens, 1091 Puya pachyphylla, 1097 Puya paniculata, 1081 Puya pectinata, 1094 Puya pichinchae, 1091 Puya pitcairnioides, 1092 Puya prosanae, 1092 Puya pumila, 1076 Puya quillotana, 1079 Puya raimondii, 1093 Puya retrorsa, 1093 Puya robinfosteri, 1094 Puya roezlii, 1094 Puya roldanii, 1094 Puya roseana, 1095 Puya rubricaulis, 1081 Puya saxatilis, 978 Puya sehuencasensis, 1095 Puya serranoensis, 1097 Puya serranoensis var. brevispica, 1097 Puya sodiroana, 1096 Puya spathacea var. harmsii, 1085 Puya suberosa, 1079 Puya tillii, 1096 Puya tuberosa, 1097 Puya vallo-grandensis, 1097 Puya vallo-grandensis var. simplex, 1097 Puya venusta, 1097 Puya vestita, 1097 Puya violacea, 1081 Puya violacea var. intermedia, 1081 Puya violacea var. monteroana, 1081 Puya weberi, 884 Puya weberiana, 1098 Puya weddelliana, 1098 Puya werdermannii, 1087 Puya whytei, 1076 Puya yakespala, 1099 Pycnanthaceae, 1299 Q Quesmea, 852 Quesnelia, 841, 859, 860 Quesnelia chacoensis, 844 Quesnelia distichantha, 853 Quesnelia strobilispica, 841 R Rajania, 1201 Rauhia, 445 Rauhia decora, 445 Rauhia megistophylla, 445 Rauhia multiflora, 445 Rauhia occidentalis, 445 Rauhia peruviana, 445 Rauhia staminosa, 446 Rectanthera, 1173

1452 Rectanthera fragrans, 1174 Renealmia, 1107 Resnova, 1269 Resnova megaphylla, 1271 Rhadamanthopsis, 1250 Rhadamanthopsis karooicus, 1258 Rhadamanthopsis namibensis, 1261 Rhadamanthus, 1249 Rhadamanthus albiflorus, 1251 Rhadamanthus arenicola, 1252 Rhadamanthus convallarioides, 1253 Rhadamanthus cyanelloides, 1254 Rhadamanthus fasciatus, 1255 Rhadamanthus karooicus, 1258 Rhadamanthus montanus, 1253 Rhadamanthus namibensis, 1261 Rhadamanthus platyphyllus, 1262 Rhadamanthus rotundus, 1262 Rhadamanthus secundus, 1262 Rhipidodendron, 486 Rhipidodendrum, 485 Rhipidodendrum dichotomum, 546 Rhipidodendrum distichum, 636 Rhipidodendrum plicatile, 636 Rhizemys, 1201 Rhodocodon, 1250 Rhodocodon campanulatus, 1262 Rhodocodon cryptopodus, 1254 Rhodocodon rotundatus, 1262 Rhodocodon rotundus, 1262 Rhodostachys, 1041 Rhodostachys albobracteata, 986 Rhodostachys andina, 1042 Rhodostachys bicolor, 986 Rhodostachys carnea, 1042 Rhodostachys chamissonis, 1042 Rhodostachys grandiflora, 1042 Rhodostachys joinvillei, 986 Rhodostachys leiboldiana, 1042 Rhodostachys litoralis, 1043 Rhodostachys micrantha, 986 Rhodostachys pitcairniifolia, 1043 Rhodostachys pitcairniifolia var. kirchhoffiana, 986 Rhodostachys urbaniana, 875 Rhoeo, 1183 Rhoeo discolor, 1186 Rhoeo spathacea, 1186 Rhopalephora, 1171 Robynsia, 21 Robynsia geminiflora, 86 Roeslia, 323 Roezlia, 323 Roezlia bulbifera, 337 Roezlia regia, 337 Roezlia regina, 337 Rokautskyia, 865 Rokautskyia microglazioui, 870 Roulinia gracilis, 1330 Roulinia humboldtiana, 1350

Index Roulinia karwinskiana, 1350 Roulinia serratifolia, 1332 Rowleyara, 823 Ruckia, 1041 Ruckia ellemetii, 1042 Runyonia, 22 Runyonia longiflora, 155 Runyonia tenuiflora, 155 Runyonia tubiflora, 155 Ruscaceae, 1315 S Sagittanthera, 1250 Sagittanthera cyanelloides, 1254 Sagittanthera mzimvubuensis, 1261 Salmia ebracteata, 1362 Salmia guineensis, 1381 Salmia spicata, 1367 Samuela, 363 Samuela carnerosana, 380 Samuela faxoniana, 385 Sanseverinia, 1353 Sanseverinia rorida, 1377 Sanseverinia thyrsiflora, 1367 Sansevieria, 1353 Sansevieria abyssinica, 1364 Sansevieria abyssinica var. angustior, 1364 Sansevieria abyssinica var. sublaevigata, 1363 Sansevieria aethiopica, 1355 Sansevieria aethiopica ssp. itumei, 1367 Sansevieria andradae, 1379 Sansevieria angolensis, 1360 Sansevieria angustiflora, 1367 Sansevieria angustifolia, 1367 Sansevieria arborescens, 1356 Sansevieria ascendens, 1356 Sansevieria aubryana, 1368 Sansevieria aubrytiana, 1368 Sansevieria bacularis, 1357 Sansevieria bagamoyensis, 1357 Sansevieria ballyi, 1357 Sansevieria ballyi var. ballyi, 1357 Sansevieria ballyi var. robertsoniae, 1357 Sansevieria bella, 1357–1358 Sansevieria bequaertii, 1372 Sansevieria bhitalae, 1358 Sansevieria bracteata, 1356 Sansevieria braunii, 1358 Sansevieria burdettii, 1358 Sansevieria burmanica, 1358 Sansevieria caespitosa, 1355 Sansevieria canaliculata, 1359 Sansevieria caulescens, 1359 Sansevieria chahinianii, 1359 Sansevieria chinensis, 1369 Sansevieria concinna, 1360 Sansevieria conspicua, 1360 Sansevieria cornui, 1378

Index Sansevieria cylindrica, 1355, 1357, 1360, 1363 Sansevieria cylindrica var. cylindrica, 1360 Sansevieria cylindrica var. patula, 1361 Sansevieria dambilonensis, 1372 Sansevieria dawei, 1361 Sansevieria deserti, 1373 Sansevieria dhofarica, 1361 Sansevieria dooneri, 1361 Sansevieria downsii, 1361 Sansevieria dumetescens, 1362 Sansevieria ebracteata, 1362 Sansevieria ehrenbergii, 1355, 1362, 1382 Sansevieria eilensis, 1362–1363 Sansevieria elliptica, 1363 Sansevieria ensifolia, 1366 Sansevieria erythraeae, 1363 Sansevieria fasciata, 1363 Sansevieria fischeri, 1364 Sansevieria formosa, 1364 Sansevieria forskaoliana, 1365 Sansevieria francisii, 1365 Sansevieria frequens, 1365 Sansevieria fulvocincta, 1355 Sansevieria gentilis, 1369 Sansevieria glauca, 1355 Sansevieria gracilis, 1365 Sansevieria gracilis var. humbertiana, 1381 Sansevieria gracilis var. somaliensis, 1365 Sansevieria gracillima, 1365 Sansevieria grandicuspis, 1366 Sansevieria grandis, 1367 Sansevieria grandis var. zuluensis, 1367 Sansevieria guineensis, 1360, 1364, 1367, 1371, 1381 Sansevieria guineensis var. β, 1367, 1371 Sansevieria guineensis var. γ, 1355 Sansevieria guineensis var. angustior, 1364 Sansevieria guineensis var. latifolia, 1367 Sansevieria hallii, 1366 Sansevieria hargeisana, 1366 Sansevieria humbertiana, 1381 Sansevieria humiflora, 1366–1367 Sansevieria hyacinthoides, 1367 Sansevieria indica, 1382 Sansevieria intermedia, 1381 Sansevieria itumei, 1367 Sansevieria jacquinii, 1381 Sansevieria kirkii, 1368 Sansevieria kirkii var. kirkii, 1368 Sansevieria kirkii var. pulchra, 1368 Sansevieria laetevirens, 1355 Sansevieria laevifolia, 1369 Sansevieria lancifolia, 1367 Sansevieria lanuginosa, 1362 Sansevieria lasciata, 1364 Sansevieria latifolia, 1367 Sansevieria laurentii, 1381 Sansevieria lavranii, 1369 Sansevieria liberica, 1369 Sansevieria liberiensis, 1378

1453 Sansevieria lineata, 1369 Sansevieria livingstoniae, 1359 Sansevieria longiflora, 1368, 1370 Sansevieria longiflora var. fernandopoensis, 1370 Sansevieria longiflora var. longiflora, 1370 Sansevieria longistyla, 1370 Sansevieria lunatifolia, 1370 Sansevieria macrophylla, 1367 Sansevieria maduraiensis, 1359 Sansevieria malawiana, 1378 Sansevieria masoniana, 1370 Sansevieria massae, 1371 Sansevieria metallica, 1371 Sansevieria metallica var. longituba, 1371 Sansevieria metallica var. metallica, 1371 Sansevieria metallica var. nyasica, 1371 Sansevieria newtoniana, 1371 Sansevieria nilotica, 1371 Sansevieria nilotica var. nilotica, 1372 Sansevieria nilotica var. obscura, 1372 Sansevieria nitida, 1372 Sansevieria nobilis, 1367 Sansevieria paniculata, 1338 Sansevieria parva, 1372 Sansevieria patens, 1372–1373 Sansevieria patersonii, 1380 Sansevieria pearsonii, 1373 Sansevieria pedicellata, 1373 Sansevieria perrotii, 1373–1374 Sansevieria pfennigii, 1359 Sansevieria pfisteri, 1374 Sansevieria phillipsiae, 1374 Sansevieria pinguicula, 1374 Sansevieria pinguicula fa. disticha, 1374 Sansevieria pinguicula ssp. disticha, 1374 Sansevieria pinguicula ssp. nana, 1374 Sansevieria pinguicula ssp. pinguicula, 1375 Sansevieria pinguicula var. nana, 1374 Sansevieria polyphylla, 1355 Sansevieria polyrhitis, 1381 Sansevieria powellii, 1375 Sansevieria powysii, 1376 Sansevieria pumila, 1355, 1366 Sansevieria quarrei, 1381 Sansevieria raffillii, 1376 Sansevieria raffillii var. glauca, 1376 Sansevieria raffillii var. raffillii, 1376 Sansevieria rhodesiana, 1373 Sansevieria robusta, 1376–1377 Sansevieria rorida, 1377 Sansevieria rosulata, 1377 Sansevieria roxburghiana, 1359 Sansevieria rufocincta, 1367 Sansevieria sambiranensis, 1377–1378 Sansevieria scabrifolia, 1355 Sansevieria schimperi, 1359 Sansevieria schweinfurthii, 1363 Sansevieria scimitariformis, 1378 Sansevieria senegambica, 1378

1454 Sansevieria singularis, 1364 Sansevieria sinus-simiorum, 1378 Sansevieria sordida, 1379 Sansevieria specksii, 1379 Sansevieria spicata, 1367 Sansevieria splendens, 1380 Sansevieria stenophylla, 1355 Sansevieria striata, 1355 Sansevieria stuckyi, 1379 Sansevieria subspicata, 1380 Sansevieria subspicata var. concinna, 1360 Sansevieria subtilis, 1380 Sansevieria suffruticosa, 1380 Sansevieria suffruticosa var. longituba, 1380 Sansevieria suffruticosa var. suffruticosa, 1380 Sansevieria sulcata, 1359 Sansevieria thunbergii, 1355 Sansevieria thyrsiflora, 1367 Sansevieria transvaalensis, 1355 Sansevieria trifasciata, 1380 Sansevieria trifasciata var. laurentii, 1381 Sansevieria trifasciata var. trifasciata, 1381 Sansevieria tsavoensis, 1357 Sansevieria varians, 1381 Sansevieria venosa, 1355 Sansevieria volkensii, 1381 Sansevieria wanguiae, 1369 Sansevieria zanzibarica, 1382 Sansevieria zebrina, 1381 Sansevieria zeylanica, 1355, 1377 Sansevieriaceae, 1315 Sarcoyucca, 363 Sarcoyucca aloifolia, 371 Sarcoyucca australis, 385 Sarcoyucca baccata, 375 Sarcoyucca brevifolia, 378 Sarcoyucca decipiens, 382 Sarcoyucca elephantipes, 394 Sarcoyucca endlichiana, 385 Sarcoyucca macrocarpa, 385 Sarcoyucca mohavensis, 408 Sarcoyucca periculosa, 403 Sarcoyucca schottii, 408 Sarcoyucca treculiana, 410 Sarcoyucca valida, 412 Schizobasis, 1250 Schizobasis angolensis, 1257 Schizobasis bruce-bayeri, 1262 Schizobasis buchubergensis, 1257 Schizobasis cuscutoides, 1257 Schizobasis dinteri, 1257 Schizobasis gracilis, 1257 Schizobasis intricata, 1256 Schizobasis macowanii, 1257 Schizobasis schlechteri, 1257 Schizobasis sigmoidea, 1262 Schizobasis volubilis, 1244 Schizobasopsis, 1243 Schizobasopsis kilimandscharica, 1244

Index Schizobasopsis volubilis, 1244 Schizocarphus, 1269 Schoenolirion, 12, 13 Scilla, 1269 Scilla albomarginata, 1271 Scilla anthericoides, 1259 Scilla brevifolia, 1248 Scilla cicatricosa, 1271 Scilla climatocarpha, 1271 Scilla collina, 1271 Scilla concolor, 1270 Scilla dracomontana, 1277 Scilla ecklonii, 1270 Scilla elevans, 1271 Scilla ensifolia, 1270 Scilla fugax, 1259 Scilla guttata, 1271 Scilla kraussii, 1278 Scilla lancifolia, 1271 Scilla laxiflora, 1271 Scilla ludwigii, 1270 Scilla maritima, 1259 Scilla micrantha, 1251 Scilla natalensis, 1278 Scilla ovatifolia, 1271 Scilla paucifolia, 1271 Scilla plumbea, 1278, 1289 Scilla prasina, 1270 Scilla pusilla, 1270 Scilla socialis, 1271 Scilla undulata, 1259 Scilla violacea, 1271 Scillaceae, 1225 Scillopsis angustifolia, 1266 Scillopsis contaminata, 1266 Scillopsis hyacinthoides, 1266 Sekanama, 1250 Sekanama delagoensis, 1254 Separotheca, 1183 Serena, 441 Setcreasea, 1183 Setcreasea hirsuta, 1185 Setcreasea pallida, 1185 Setcreasea purpurea, 1185 Sincoraea, 866, 1101 Sincoraea albopicta, 1047, 1102–1103 Sincoraea burle-marxii, 1103 Sincoraea hatschbachii, 1103–1104 Sincoraea heleniceae, 1103, 1104 Sincoraea mucugensis, 1104 Sincoraea navioides, 1049, 1067, 1101–1104 Sismondaea, 1201 Smilacina forskaoliana, 1364 Smithara, 743 Spetaea, 1289 Spetaea lachenaliiflora, 1278, 1289 Spinifex squarrosus, 1308 Spironema, 1173 Spironema fragrans, 1174

Index Spironema robbinsii, 1176 Spironema warszewicziana, 1176 Squilla, 1249 Squilla anthericoides, 1259 Squilla insularis, 1259 Squilla littoralis, 1259 Squilla maritima, 1259 Squilla numidica, 1259 Squilla pancration, 1259 Squilla sphaeroidea, 1259 Stellarioides, 1231 Stellarioides canaliculata, 1233 Stellarioides lebaensis, 1238 Stellarioides longibracteata, 1233 Stellarioides spiralifolia, 1283 Stenomeridaceae, 1199 Strepsia, 1107 Strepsiphylla, 1249 Streptomea, 852 Strophis, 1201 Stylochaeton, 458 Succosaria, 486 Sugillaria, 1269 Synandrospadix, 458 Syncodium, 1281 Sypharissa, 1249 Sypharissa multifolia, 1260 T Tacca, 1199 Taccaceae, 1199 Taeniola, 1231 Taetsia, 1295 Tamaceae, 1199 Tamnaceae, 1199 Tamus, 1199, 1201 Tamus elephantipes, 1203 Tamus sylvestris, 1207 Tenicroa, 1249 Tenicroa multifolia, 1260 Testudinaria, 1201 Testudinaria cocolmeca, 1205 Testudinaria elephantipes, 1203 Testudinaria elephantipes fa. montana, 1204 Testudinaria elephantipes var. montana, 1203 Testudinaria glauca, 1205 Testudinaria macrostachya, 1205 Testudinaria montana, 1203, 1206 Testudinaria montana var. paniculata, 1207 Testudinaria multiflora, 1206 Testudinaria paniculata, 1207 Testudinaria paniculata var. brevipes, 1206 Testudinaria rehmannii, 1207 Testudinaria sylvatica, 1206 Testudinaria sylvatica var. brevipes, 1206 Testudinaria sylvatica var. lydenbergensis, 1206 Testudinaria sylvatica var. multiflora, 1206 Testudinaria sylvatica var. paniculata, 1207

1455 Testudinaria sylvatica var. rehmannii, 1207 Testudinaria sylvestris, 1206 Thaumaza, 1219 Thaumaza paradoxa, 1220 Themidaceae, 11 Thuranthos, 1250 Ti kouka, 1296 Tillandsia, 1107, 1108, 1302 Tillandsia acutifolia, 844 Tillandsia aizoides, 1109–1110 Tillandsia albertiana, 1110 Tillandsia andicola, 1110–1111, 1130 Tillandsia angulosa, 1111 Tillandsia araujei, 1111–1112 Tillandsia araujei var. minima, 1111–1112 Tillandsia argentea, 1156 Tillandsia argentina, 1112–1113 Tillandsia arhiza, 1113 Tillandsia arhiza var. rupestris, 1113 Tillandsia ariza-juliae, 1113 Tillandsia armadae, 844 Tillandsia atrichoides, 1140 Tillandsia baileyi, 1114 Tillandsia boryana, 844 Tillandsia bracteata, 853 Tillandsia brealitoensis, 1114 Tillandsia breviscapa, 1149 Tillandsia breweri, 1113 Tillandsia bulbosa, 1114–1115 Tillandsia bulbosa var. brasiliensis, 1114 Tillandsia bulbosa var. picta, 1114 Tillandsia burle-marxii, 1115 Tillandsia butzii, 1115–1116 Tillandsia caerulea, 1154 Tillandsia caespitosa, 1153 Tillandsia calamifolia, 844 Tillandsia caliginosa, 1116 Tillandsia capillaris, 1116–1117 Tillandsia capillaris fa. cordobensis, 1159 Tillandsia capillaris fa. hieronymi, 1117 Tillandsia capillaris fa. incana, 1117 Tillandsia capillaris fa. typica, 1117 Tillandsia capillaris fa. virescens, 1159 Tillandsia capillaris var. incana, 1117 Tillandsia capillaris var. lanuginosa, 1117 Tillandsia caput-medusae, 1117–1118 Tillandsia castellanii, 1118 Tillandsia chiapensis, 1118–1119 Tillandsia chilensis, 1144 Tillandsia chlorantha, 881 Tillandsia circinalis, 1125 Tillandsia circinnatoides, 1119–1120 Tillandsia coarctata, 1146 Tillandsia coarctata var. pedicellata, 1146 Tillandsia colganii, 1120 Tillandsia comata, 853 Tillandsia compressa, 1130 Tillandsia confusa, 1125 Tillandsia confusa var. minor, 1125

1456 Tillandsia confusa var. saxatilis, 1125 Tillandsia copanensis, 1120–1121 Tillandsia cordobensis, 1159 Tillandsia cotagaitensis, 1121 Tillandsia crocata, 1121–1122 Tillandsia crocata fa. major, 1116 Tillandsia crocata var. tristis, 1116 Tillandsia curvifolia, 1122–1123 Tillandsia decomposita, 1125 Tillandsia dependens, 1159 Tillandsia dependens fa. percordobensis, 1159 Tillandsia dependens fa. perusneoides, 1159 Tillandsia dependens var. percordobensis, 1159 Tillandsia dependens var. perusneoides, 1159 Tillandsia dependens var. sanzinii, 1159 Tillandsia diaguitensis, 1123–1124 Tillandsia dorisdaltoniae, 1124 Tillandsia dorotheae, 1124–1125 Tillandsia duratii, 1125, 1125–1126 Tillandsia duratii ssp. reichenbachii, 1152 Tillandsia duratii var. saxatilis, 1125 Tillandsia eggersii, 1154 Tillandsia ehlersiana, 1126 Tillandsia ehrenbergiana, 1158 Tillandsia ehrenbergii, 1158 Tillandsia ensifolia, 844 Tillandsia erecta, 1127 Tillandsia erecta var. rigida, 1127 Tillandsia erici, 1127–1128 Tillandsia erubescens, 1134 Tillandsia erythraea, 1114 Tillandsia euosma, 1152 Tillandsia favillosa, 1144 Tillandsia ferrisiana, 1128 Tillandsia flavescens, 1154 Tillandsia floribunda, 1125 Tillandsia fresnilloensis, 1128–1129 Tillandsia funebris, 1129 Tillandsia fusca, 1144 Tillandsia gigantea, 1125 Tillandsia gilliesii, 1129–1130 Tillandsia glabrior, 1130–1131 Tillandsia grisea, 1138 Tillandsia grisebachii, 1154 Tillandsia guerreroensis, 1131 Tillandsia hasei, 1131–1132 Tillandsia hegeri, 1132–1133 Tillandsia helmutii, 1133–1134 Tillandsia heptantha, 844 Tillandsia herzogii, 1152 Tillandsia hieronymi, 1117 Tillandsia hieronymi var. lichenoides, 1117 Tillandsia hirta, 1134 Tillandsia imbricata, 844 Tillandsia inanis, 1116 Tillandsia incana, 1116 Tillandsia intermedia, 1134 Tillandsia ionantha, 1134–1135 Tillandsia ionantha var. vanhyningii, 1135

Index Tillandsia ionantha var. zebrina, 1135 Tillandsia juncea, 116 Tillandsia juruana, 1145 Tillandsia kirschnekii, 1135–1136 Tillandsia klausii, 1136–1137 Tillandsia kuehhasii, 1137 Tillandsia kunthiana, 1138 Tillandsia lanata, 1138 Tillandsia langlassei, 1117 Tillandsia latifolia, 1137–1138 Tillandsia latispatha, 844 Tillandsia lepidosepala, 1139 Tillandsia lichenoides, 1117 Tillandsia lineata, 844 Tillandsia lithophila, 1139 Tillandsia loliacea, 1140 Tillandsia lydiae, 1140–1141 Tillandsia macrocnemis, 1160 Tillandsia mandonii, 1143 Tillandsia mauryana, 1141–1142 Tillandsia mitlaensis, 1142 Tillandsia mollis, 1142–1143 Tillandsia murorum, 1138 Tillandsia myosura, 1116, 1143 Tillandsia myosura var. saxicola, 1157 Tillandsia nappii, 1143 Tillandsia nappii var. darwinii, 1153 Tillandsia neglecta, 1143–1144 Tillandsia obscura, 844 Tillandsia osyana, 860 Tillandsia oxysepala, 1138 Tillandsia paleacea, 1144–1145 Tillandsia paraensis, 1145–1146 Tillandsia paucifolia, 1146 Tillandsia paucifolia var. schubertii, 1134 Tillandsia pedicellata, 1146–1147 Tillandsia peiranoi, 1147 Tillandsia permutata, 1117 Tillandsia platystachia, 844 Tillandsia polystachia, 853 Tillandsia porongoensis, 1147–1148 Tillandsia praschekii, 1148–1149 Tillandsia propinqua, 1117, 1159 Tillandsia propinqua var. rectangula, 1151 Tillandsia propinqua var. saxicola, 1117 Tillandsia pruinosa, 1149–1150 Tillandsia pseudobaileyi, 1150 Tillandsia pueblensis, 1150–1151 Tillandsia pueblensis var. glabrior, 1130 Tillandsia pumila, 1114 Tillandsia pusilla, 1117 Tillandsia quadriflora, 1140 Tillandsia quesneliana, 1135 Tillandsia ramealis, 844 Tillandsia rectangula, 1151–1152 Tillandsia recurvata, 1159 Tillandsia reducta, 1152 Tillandsia reichenbachii, 1152–1153 Tillandsia retorta, 1153

Index Tillandsia revoluta, 1125 Tillandsia rigida, 844, 1127 Tillandsia rosea, 855 Tillandsia rubentifolia, 1135 Tillandsia rupestris, 1113 Tillandsia rupestris var. pendens, 1113 Tillandsia sanctae-crucis, 1145 Tillandsia saxatilis, 854 Tillandsia saxicola, 1156 Tillandsia scalarifolia, 1144 Tillandsia schatzlii, 1153–1154 Tillandsia schenckiana, 1144 Tillandsia schiedeana, 1154 Tillandsia schiedeana ssp. glabrior, 1130 Tillandsia scopus, 1135 Tillandsia seleriana, 1154–1155 Tillandsia sericea, 1160 Tillandsia stolpii, 1159 Tillandsia suaveolens, 1160 Tillandsia subulifera, 1155 Tillandsia tectorum, 1156 Tillandsia tenebra, 1156–1157 Tillandsia tetrasticha, 853 Tillandsia tomasii, 1159 Tillandsia tomentosa, 1125 Tillandsia tortilis, 1157–1158, 1158 Tillandsia tortilis ssp. curvifolia, 1122 Tillandsia tortuosa, 844 Tillandsia tuberosa, 956 Tillandsia tucumanensis, 1152 Tillandsia unca, 1112–1113, 1160 Tillandsia undulata, 1140 Tillandsia variabilis, 844 Tillandsia variegata, 1116 Tillandsia vestita, 1154 Tillandsia violaceiflora, 1158 Tillandsia violata, 844 Tillandsia virescens, 1158–1159 Tillandsia virescens var. sanzinii, 1159 Tillandsia weberi, 1159–1160 Tillandsia weddellii, 1125 Tillandsia williamsii, 1159 Tillandsia xiphioides, 1160–1161 Tillandsia xiphioides var. minor, 1160 Tillandsia yucatana, 1146 Tillandsia yuncharaensis, 1161–1162 Tillandsia zecheri, 1162–1163 Tillandsiaceae, 835 Timmia obliqua, 438 Tomoxis, 1231 Tonningia, 1179 Tonningia nodiflora, 1181 Tonningia speciosa, 1180 Trachyandra, 829 Trachyandra adamsonii, 830 Trachyandra aridimontana, 830 Trachyandra blepharophora, 830 Trachyandra bracteosa, 830 Trachyandra canaliculata, 830

1457 Trachyandra ciliata, 830 Trachyandra falcata, 831 Trachyandra involucrata, 831 Trachyandra longifolia, 830 Trachyandra tabularis, 831 Trachyandra tortilis, 832 Trachyandra vespertina, 830 Tradescantia, 1183 Tradescantia albiflora, 1185 Tradescantia albiflora cv. Albovittata, 1185 Tradescantia anagallidea, 1185 Tradescantia blossfeldiana, 1184 Tradescantia brachyphylla, 1175 Tradescantia callisia, 1176 Tradescantia cerinthoides, 1184 Tradescantia crassipes, 1184 Tradescantia crassula, 1184 Tradescantia crassula var. gaudichaudii, 1184 Tradescantia cumanensis, 1191 Tradescantia cymbispatha var. villosissima, 1184 Tradescantia discolor, 1186 Tradescantia fluminensis, 1185 Tradescantia fluminensis cv. Argenteo-variegata, 1185 Tradescantia formosa, 1181 Tradescantia glandulosa, 1190 Tradescantia hirta, 1185 Tradescantia ionantha, 1190 Tradescantia koernickeana, 1184 Tradescantia martensiana, 1175 Tradescantia micrantha, 1174 Tradescantia multiflora, 1190, 1191 Tradescantia multiflora var. parviflora, 1191 Tradescantia multiflora var. tobagensis, 1191 Tradescantia mundula, 1185 Tradescantia navicularis, 1175 Tradescantia nodiflora, 1181 Tradescantia pallida, 1185–1186 Tradescantia pallida cv. Purple Heart, 1186 Tradescantia pallida cv. Purpurea, 1186 Tradescantia parviflora, 1191 Tradescantia pendula, 1187 Tradescantia pexata, 1186 Tradescantia pflanzii, 1190 Tradescantia procumbens, 1191 Tradescantia radiata, 1190 Tradescantia schwirkowskiana, 1184 Tradescantia sillamontana, 1186 Tradescantia spathacea, 1184, 1186 Tradescantia spathacea cv. Concolor, 1186 Tradescantia spathacea cv. Variegata, 1186 Tradescantia spathacea cv. Vittata, 1186 Tradescantia speciosa, 1180 Tradescantia subscaposa, 1176 Tradescantia virginiana, 1184 Tradescantia warszewicziana, 1176 Tradescantia zebrina, 1184, 1187 Tradescantiaceae, 1167 Treleasea, 1183 Tricharis, 1247

1458 Trichopodaceae, 1199 Trimelopter, 1231 Tripogandra, 1189 Tripogandra cumanensis, 1191 Tripogandra floribunda, 1191 Tripogandra glandulosa, 1189 Tripogandra grandiflora, 1190 Tripogandra ionantha, 1190 Tripogandra multiflora, 1190 Tripogandra multiflora fa. parviflora, 1191 Tripogandra parviflora, 1191 Tripogandra pflanzii, 1190 Tripogandra radiata, 1190 Tripogandra warszewicziana, 1176 Tropitria, 1183 Tropitria crassula, 1184 Tuberosa, 21 Tuberosa amica, 48 Tulista, 769 Tulista aristata, 505 Tulista bicarinata, 709 Tulista bullulata, 702 Tulista congesta, 703 Tulista corrugata, 703 Tulista dinteri, 547 Tulista foliolosa, 704 Tulista herrei, 704 Tulista kingiana, 790 Tulista koelmaniorum, 791 Tulista koelmaniorum var. mcmurtryi, 791 Tulista margaritifera, 793 Tulista marginata, 794 Tulista marginata var. mortonii, 796 Tulista minima, 795 Tulista minima var. poellnitziana, 796 Tulista opalina, 796 Tulista opalina var. zenigata, 790 Tulista pumila, 793 Tulista pumila var. ohkuwae, 793 Tulista pumila var. sparsa, 793 Tulista pungens, 804 Tulista rubriflora, 828 Tulista sladeniana, 657 Tulista spiralis, 705 Tulista variegata, 674 Tulista viscosa, 813 Tulworthia, 769 Tulworthiopsis, 769 Tupistraceae, 1315 U Ubium, 1201 Urginavia, 1250 Urginavia altissima, 1251 Urginavia epigea, 1252 Urginavia micrantha, 1251 Urginavia viridula, 1252 Urginea, 1249

Index Urginea alooides, 498 Urginea altissima, 1251 Urginea anthericoides, 1259, 1260 Urginea aphylla, 1259 Urginea brevipes, 1251 Urginea delagoensis, 1254 Urginea epigea, 1252 Urginea eriospermoides, 1252 Urginea forsteri, 1251 Urginea fugax, 1259 Urginea gigantea, 1251 Urginea hesperia, 1259 Urginea insularis, 1259 Urginea kniphofioides, 1251 Urginea littoralis, 1259 Urginea lydenburgensis, 1255 Urginea maritima, 1259 Urginea maritima fa. angustifolia, 1260 Urginea maritima fa. latifolia, 1260 Urginea maritima var. angustifolia, 1260 Urginea maritima var. anthericoides, 1259 Urginea maritima var. maura, 1259 Urginea maritima var. numidica, 1259 Urginea maritima var. pancration, 1259 Urginea maritima var. sphaeroidea, 1259 Urginea maritima var. stenophylla, 1259 Urginea maura, 1259 Urginea micrantha, 1251 Urginea mouretii, 1234 Urginea multifolia, 1260 Urginea numidica, 1259 Urginea pancration, 1259 Urginea scilla, 1259 Urginea sphaeroidea, 1259 Urginea tazensis, 1260 Urginea ubomboensis, 1255 Urginea undulata, 1259, 1260 Urginea viridula, 1252 Urgineopsis, 1250 Urginia, 1249 Uropetalon, 1247 Uropetalon hyacinthoides, 1248 Uropetalum, 1247 Urophyllon, 1231 Ursumea, 853 V Vallota, 435 Vanilla, 1300 Vanillaceae, 1299 Vera-duthiea, 1250 Viridantha, 1107 Viridantha curvifolia, 1122 Viridantha longisepala, 1139 Viridantha mauryana, 1141 Viridantha tortilis, 1157 Vriesea, 841 Vriesea espinosae, 841

Index Vriesea glauca, 844 Vriesea glaucophylla, 1030 Vriesea imbricata, 844 Vriesea legrelliana, 855 Vriesea sanctae-crucis, 1145 W Wallisia, 1107 Whiteheadia, 1273 Whiteheadia bifolia, 1274 Whiteheadia etesionamibensis, 1275 Whiteheadia latifolia, 1274 Wittmackia, 851 Wolffiaceae, 457 X Xanthorrhoea, 1389, 1390 Xanthorrhoeaceae, 1389 Xeodolon, 1269 Y Yucca, 11, 14, 15, 363 Yucca acaulis, 325 Yucca acrotricha, 1330 Yucca acuminata, 392, 393 Yucca acutifolia, 392 Yucca agavoides, 410 Yucca albo-spica, 381 Yucca aletriformis, 1338 Yucca aloifolia, 371, 371–372 Yucca aloifolia fa. arcuata, 371 Yucca aloifolia fa. crenulata, 371 Yucca aloifolia fa. draconis, 371 Yucca aloifolia fa. marginata, 371 Yucca aloifolia fa. purpurea, 372 Yucca aloifolia fa. tenuifolia, 371 Yucca aloifolia fa. tricolor, 371 Yucca aloifolia var. arcuata, 371 Yucca aloifolia var. draconis, 371 Yucca aloifolia var. gigantea, 372 Yucca aloifolia var. marginata, 371 Yucca aloifolia var. menandii, 372 Yucca aloifolia var. purpurea, 372 Yucca aloifolia var. quadricolor-variegata, 372 Yucca aloifolia var. roseomarginata, 371 Yucca aloifolia var. serratifolia, 372 Yucca aloifolia var. stenophylla, 371 Yucca aloifolia var. tenuifolia, 371 Yucca aloifolia var. tricolor, 371 Yucca aloifolia var. variegata, 372 Yucca aloifolia var. versicolor, 372 Yucca aloifolia var. yucatana, 372 Yucca andreana, 371 Yucca angustifolia, 381, 383, 390, 393 Yucca angustifolia fa. genuina, 390 Yucca angustifolia var. mollis, 374

1459 Yucca angustifolia var. radiosa, 383 Yucca angustissima, 373 Yucca angustissima ssp. angustissima, 373 Yucca angustissima ssp. avia, 373 Yucca angustissima ssp. kanabensis, 373–374 Yucca angustissima ssp. toftiae, 374 Yucca angustissima var. avia, 373 Yucca angustissima var. kanabensis, 374 Yucca angustissima var. toftiae, 374 Yucca antwerpensis, 371 Yucca arborescens, 378 Yucca arcuata, 371 Yucca argospatha, 410 Yucca argyraea, 337 Yucca argyrophylla, 337 Yucca arizonica, 376 Yucca arkansana, 374–375 Yucca arkansana ssp. freemanii, 389 Yucca arkansana ssp. louisianensis, 389 Yucca arkansana var. paniculata, 374 Yucca armata, 371 Yucca aspera, 410 Yucca atkinis, 383 Yucca atkinsii, 371, 372 Yucca australis, 385, 388 Yucca australis var. valida, 385 Yucca baccata, 375, 385 Yucca baccata fa. circinata, 403 Yucca baccata fa. fragilifolia, 375 Yucca baccata fa. genuina, 376 Yucca baccata fa. parviflora, 376 Yucca baccata fa. periculosa, 403 Yucca baccata ssp. baccata, 375–376 Yucca baccata ssp. confinis, 376 Yucca baccata ssp. thornberi, 376 Yucca baccata ssp. vespertina, 376 Yucca baccata var. australis, 388 Yucca baccata var. brevifolia, 376 Yucca baccata var. circinata, 403 Yucca baccata var. filifera, 388 Yucca baccata var. hystrix, 376 Yucca baccata var. periculosa, 403 Yucca baccata var. scabrifolia, 375 Yucca baccata var. vespertina, 376 Yucca baileyi, 376–377 Yucca baileyi ssp. intermedia, 396 Yucca baileyi var. intermedia, 396 Yucca baileyi var. navajoa, 377 Yucca barrancasecca, 371 Yucca boerhaavii, 393 Yucca boscii, 116 Yucca brasiliensis, 371 Yucca brevifolia, 376, 377–378, 408 Yucca brevifolia fa. herbertii, 378 Yucca brevifolia fa. kernensis, 378 Yucca brevifolia ssp. herbertii, 378 Yucca brevifolia ssp. jaegeriana, 397 Yucca brevifolia var. herbertii, 378 Yucca brevifolia var. jaegeriana, 397

1460 Yucca brevifolia var. wolfei, 378 Yucca californica, 358, 407 Yucca campestris, 378–379 Yucca canaliculata, 410 Yucca canaliculata var. filifera, 388 Yucca canaliculata var. pendula, 410 Yucca capensis, 379–380 Yucca carnerosana, 380 Yucca carrierei, 371 Yucca cernua, 380–381 Yucca circinata, 403 Yucca coahuilensis, 383 Yucca coloma, 395 Yucca concava, 388, 411 Yucca concava var. longifolia, 389 Yucca confinis, 376 Yucca conspicua, 371 Yucca constricta, 381, 383 Yucca contorta, 407 Yucca cornuta, 411 Yucca crassifila, 410 Yucca crenulata, 371 Yucca crinifera, 371 Yucca decipiens, 381–382 Yucca declinata, 382 Yucca desmetiana, 383 Yucca draco, 1339 Yucca draconis, 371 Yucca draconis var. arborescens, 378 Yucca ehrenbergii, 371 Yucca elata, 383 Yucca elata ssp. elata, 383–384 Yucca elata ssp. utahensis, 411 Yucca elata ssp. verdiensis, 384 Yucca elata var. magdalenae, 383 Yucca elata var. utahensis, 411 Yucca elata var. verdiensis, 384 Yucca eleana, 395 Yucca elephantipes, 394, 395 Yucca elephantipes var. ghiesbreghtii, 394 Yucca elephantipes var. gigantea, 390 Yucca ellacombei, 392 Yucca endlichiana, 384–385 Yucca engelmannii, 358 Yucca ensifera, 371 Yucca ensifolia, 371, 392 Yucca exigua, 389 Yucca eylesii, 393 Yucca falcata, 393 Yucca falcata var. semicylindrica, 393 Yucca faxoniana, 385–386 Yucca feeanoukiae, 386–387 Yucca filamentosa, 375, 387–388 Yucca filamentosa [?] foliis-aureovariegatis, 387 Yucca filamentosa [?] meldensis, 389 Yucca filamentosa fa. concava, 389 Yucca filamentosa fa. flaccida, 388 Yucca filamentosa fa. genuina, 387 Yucca filamentosa fa. glaucescens, 389 Yucca filamentosa fa. orchioides, 389 Yucca filamentosa fa. puberula, 389

Index Yucca filamentosa fa. variegata, 387 Yucca filamentosa ssp. concava, 389 Yucca filamentosa ssp. smalliana, 389 Yucca filamentosa var. angustifolia, 389 Yucca filamentosa var. antwerpensis, 389 Yucca filamentosa var. bicolor, 387 Yucca filamentosa var. bracteata, 387 Yucca filamentosa var. concava, 388 Yucca filamentosa var. elmensis, 387 Yucca filamentosa var. flaccida, 388 Yucca filamentosa var. glaucescens, 389 Yucca filamentosa var. grandiflora, 389 Yucca filamentosa var. laevigata, 387 Yucca filamentosa var. latifolia, 387 Yucca filamentosa var. maxima, 387, 389 Yucca filamentosa var. media, 387 Yucca filamentosa var. mexicana, 387 Yucca filamentosa var. nobilis, 387 Yucca filamentosa var. orchioides, 389 Yucca filamentosa var. patens, 387 Yucca filamentosa var. puberula, 389 Yucca filamentosa var. ramosa, 387 Yucca filamentosa var. recurvifolia, 387 Yucca filamentosa var. smalliana, 389 Yucca filamentosa var. variegata, 387 Yucca filifera, 376, 388 Yucca flaccida, 388–390 Yucca flaccida fa. integra, 389 Yucca flaccida fa. lineata, 389 Yucca flaccida fa. orchioides, 389 Yucca flaccida var. exigua, 389 Yucca flaccida var. glaucescens, 389 Yucca flaccida var. grandiflora, 389 Yucca flaccida var. major, 389 Yucca flaccida var. smalliana, 389 Yucca flexilis, 393 Yucca flexilis fa. boerhaavii, 393 Yucca flexilis fa. ensifolia, 392 Yucca flexilis fa. patens, 392 Yucca flexilis fa. peacockii, 393 Yucca flexilis fa. semicylindrica, 393 Yucca flexilis var. ensifolia, 392 Yucca flexilis var. falcata, 393 Yucca flexilis var. hildrethii, 393 Yucca flexilis var. nobilis, 393 Yucca flexilis var. semicylindrica, 393 Yucca flexilis var. tortulata, 392 Yucca fragilifolia, 375 Yucca freemanii, 389 Yucca fuauxiana, 371 Yucca funifera, 352 Yucca ghiesbreghtii, 394 Yucca gigantea, 390 Yucca gilbertiana, 395 Yucca glauca, 387, 390–391 Yucca glauca fa. montana, 391 Yucca glauca fa. stricta, 390 Yucca glauca ssp. albertana, 391 Yucca glauca ssp. gurneyi, 390 Yucca glauca ssp. stricta, 390 Yucca glauca var. arkansana, 390

Index Yucca glauca var. constricta, 381 Yucca glauca var. elata, 390 Yucca glauca var. gurneyi, 390 Yucca glauca var. mollis, 374 Yucca glauca var. radiosa, 390 Yucca glauca var. rosea, 390 Yucca glauca var. stricta, 390 Yucca glaucescens, 389 Yucca gloriosa, 391 Yucca gloriosa fa. acuminata, 392 Yucca gloriosa fa. genuina, 391 Yucca gloriosa fa. longifolia, 392 Yucca gloriosa fa. maculata, 392 Yucca gloriosa fa. mediostriata, 392 Yucca gloriosa fa. minor, 392 Yucca gloriosa fa. nobilis, 392 Yucca gloriosa fa. obliqua, 392 Yucca gloriosa fa. pendula, 393 Yucca gloriosa fa. planifolia, 371 Yucca gloriosa fa. pruinosa, 392 Yucca gloriosa fa. rufocincta, 393 Yucca gloriosa fa. superba, 392 Yucca gloriosa fa. tortulata, 392 Yucca gloriosa subvar. parviflora, 392 Yucca gloriosa var. acuminata, 392 Yucca gloriosa var. elegans, 393 Yucca gloriosa var. ellacombei, 392 Yucca gloriosa var. flexilis, 393 Yucca gloriosa var. glaucescens, 392 Yucca gloriosa var. gloriosa, 391–393 Yucca gloriosa var. longifolia, 392 Yucca gloriosa var. maculata, 392 Yucca gloriosa var. marginata, 393 Yucca gloriosa var. marginata-aurea, 393 Yucca gloriosa var. mediopicta, 392 Yucca gloriosa var. mediostriata, 392 Yucca gloriosa var. minor, 392 Yucca gloriosa var. mollis, 393 Yucca gloriosa var. nobilis, 392 Yucca gloriosa var. obliqua, 392 Yucca gloriosa var. planifolia, 393 Yucca gloriosa var. plicata, 392 Yucca gloriosa var. pruinosa, 392 Yucca gloriosa var. recurvata, 392 Yucca gloriosa var. recurvifolia, 393 Yucca gloriosa var. robusta, 392 Yucca gloriosa var. rufocincta, 393 Yucca gloriosa var. superba, 392 Yucca gloriosa var. tortulata, 392 Yucca gloriosa var. tristis, 393–394 Yucca gloriosa var. variegata, 393 Yucca gracilis, 371, 1330 Yucca graminifolia, 358, 1330 Yucca grandiflora, 394 Yucca grandis, 393 Yucca guatemalensis, 394–395 Yucca hanburyi, 376 Yucca harrimaniae, 395 Yucca harrimaniae ssp. gilbertiana, 395 Yucca harrimaniae ssp. harrimaniae, 395–396 Yucca harrimaniae ssp. neomexicana, 401

1461 Yucca harrimaniae ssp. sterilis, 396 Yucca harrimaniae var. gilbertiana, 395 Yucca harrimaniae var. neomexicana, 401 Yucca harrimaniae var. sterilis, 396 Yucca haruckeriana, 371 Yucca helkinsi, 383 Yucca horrida, 371 Yucca howard-smithii, 371 Yucca integerrima, 392 Yucca intermedia, 396–397 Yucca intermedia var. ramosa, 396 Yucca jaegeriana, 397 Yucca jaliscensis, 397–398 Yucca japonica, 393 Yucca juncea, 371 Yucca kanabensis, 374 Yucca karlsruhensis, 398 Yucca lacandonica, 398–399 Yucca laevigata, 393 Yucca linearifolia, 399–400 Yucca linearis, 399 Yucca longifolia, 393, 410, 411, 1350 Yucca louisianensis, 389 Yucca louisianensis var. paniculata, 374 Yucca luminosa, 405 Yucca lutescens, 407 Yucca macrocarpa, 385, 408 Yucca madrensis, 400 Yucca massiliensis, 371 Yucca mazelii, 395 Yucca meldensis, 389 Yucca mexicana, 371, 393 Yucca mixtecana, 400–401 Yucca mohavensis, 408 Yucca mooreana, 394 Yucca nana, 395 Yucca navajoa, 377 Yucca necopina, 401 Yucca neomexicana, 401–402 Yucca newberryi, 358 Yucca newberryi ssp. mckelveyana, 358 Yucca nitida, 359 Yucca nobilis, 393 Yucca obliqua, 392 Yucca orchioides, 389 Yucca orchioides var. major, 389 Yucca ortgiesiana, 358 Yucca pallida, 402 Yucca pallida var. edentata, 402 Yucca parmentieri, 337, 371 Yucca parviflora, 354 Yucca patens, 392 Yucca peacockii, 393 Yucca pendula, 393 Yucca pendula var. aurea, 393 Yucca pendula var. variegata, 393 Yucca peninsularis, 359 Yucca periculosa, 403 Yucca pitcairniifolia, 1007 Yucca plicata, 392 Yucca plicatilis, 392

1462 Yucca polyphylla, 381 Yucca potosina, 403–404 Yucca pringlei, 337 Yucca pruinosa, 392 Yucca puberula, 389, 408 Yucca purpurea, 372 Yucca quadricolor, 371, 372 Yucca quadricolor var. stokesii, 372 Yucca queretaroensis, 404 Yucca quinnarjenii, 371 Yucca radiosa, 383 Yucca recurva, 393 Yucca recurvata, 410 Yucca recurvifolia, 393 Yucca recurvifolia fa. elegans, 393 Yucca recurvifolia fa. rufocincta, 393 Yucca recurvifolia fa. variegata, 393 Yucca recurvifolia var. marginata, 393 Yucca recurvifolia var. rufocincta, 393 Yucca recurvifolia var. tristis, 393 Yucca reverchonii, 404–405 Yucca revoluta, 410 Yucca rigida, 371, 405–406 Yucca roezlii, 395 Yucca rostrata, 406–407 Yucca rostrata fa. integra, 410 Yucca rostrata var. linearis, 399 Yucca rubra, 371 Yucca rufocincta, 393 Yucca rupicola, 402, 407 Yucca rupicola fa. pallida, 402 Yucca rupicola var. edentata, 402 Yucca rupicola var. rigida, 405 Yucca rupicola var. tortifolia, 407 Yucca scabrifolia, 375 Yucca schidigera, 407–408 Yucca schottii, 408–409 Yucca schottii var. jaliscensis, 398 Yucca schottii var. valida, 412 Yucca semicylindrica, 393 Yucca serratifolia, 1332 Yucca serrulata, 371 Yucca serrulata var. angustifolia, 372 Yucca serrulata var. variegata, 372 Yucca smalliana, 389 Yucca smetiana, 383 Yucca spinosa, 410 Yucca standleyi, 377 Yucca stenophylla, 390, 393 Yucca sterilis, 396 Yucca striata, 371 Yucca striatula, 371 Yucca stricta, 390 Yucca stricta var. elatior, 390 Yucca stricta var. intermedia, 390 Yucca sulcata, 371

Index Yucca superba, 343, 392 Yucca tenuifolia, 371 Yucca tenuistyla, 409–410 Yucca thompsoniana, 410 Yucca thompsoniana fa. viridis, 410 Yucca thornberi, 376 Yucca toftiae, 374 Yucca toneliana, 337, 371 Yucca torreyi, 411 Yucca torreyi fa. parviflora, 411 Yucca tortifolia, 407 Yucca tortilis, 407 Yucca tortulata, 392 Yucca treculiana, 410–411 Yucca treculiana var. canaliculata, 410 Yucca treculiana var. glauca, 411 Yucca treculiana var. succulenta, 411 Yucca treleasei, 371, 376 Yucca tricolor, 372 Yucca undulata, 410 Yucca utahensis, 411–412 Yucca valida, 412 Yucca verdiensis, 384 Yucca vespertina, 376 Yucca vomerensis, 371 Yucca whipplei, 358, 359 Yucca whipplei fa. graminifolia, 358 Yucca whipplei ssp. caespitosa, 359 Yucca whipplei ssp. intermedia, 359 Yucca whipplei ssp. newberryi, 358 Yucca whipplei ssp. parishii, 359 Yucca whipplei ssp. percursa, 359 Yucca whipplei ssp. rigata, 359 Yucca whipplei ssp. typica, 359 Yucca whipplei var. caespitosa, 359 Yucca whipplei var. intermedia, 359 Yucca whipplei var. parishii, 359 Yucca whipplei var. percursa, 359 Yucca whipplei var. violacea, 358 Yucca yucatana, 372 Yuccaceae, 9 Yuccites, 365 Z Zahariadia, 1282 Zamioculcas, 461 Zamioculcas lanceolata, 462 Zamioculcas loddigesii, 462 Zamioculcas zamiifolia, 461–462 Zebrina, 1183 Zebrina pendula, 1187 Zebrina purpusii, 1187 Zephyranthaceae, 425 Zuccagnia, 1247