S M I T H S O N I A N C O N T R I B U T I O N S T O B O T A N Y 0 N U M B E R 7 6 The Morphology and Taxonomy of Aneilema R. Brown (Commelinaceae) Robert B. Faden SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1991 A B S T R A C T Faden, Robert B. The Morphology and Taxonomy of Aneilema R. Brown (Commelinaceae). Smithsonian Contributions to Botany, number 76, 166 pages, 57 figures, 17 maps, 7 plates, 1991.-Vegetative and reproductive morphology and their variation within Aneilema, one of the largest genera of Commelinaceae, are described in detail. The taxonomic usefulness of the various characters is considered, and floral attributes are found to be particularly important. Within the genus, great diversity is found in habit, pedicel curvature, petal color, stamen curvature, floral hair types and distribution, and seed shape and testa patterns. Anatomy, cytology, phytochemistry, and reproductive biology of Aneilema are treated briefly. Aneilema is divided into seven sections, with sections Rendlei, Somaliensia, Brevibarbata, and Pedunculosa described as new. Keys to the sections and to all 62 species rwognized are provided. The 22 species comprising sections Rendlei, Somaliensia, and Lamprodithyros are treated in detail, with nine new species and five new subspecies described. One naturally occurring hybrid is also newly recognized. Two species originally described in Aneilema but of uncertain generic position, A. calandrinoides and A. brasiliense, are discussed in detail. A list of all specific names that have ever been included in Aneilema and their current status (including new combinations and new synonymy, when appropriate) is appended. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. Library of Congress Cataloging-in-Publication Data Faden, Robert. B. The morphology and taxonomy of Aneilema R Brown (Cmmelinawae) /Robert B. Fadem. p. cm. -(Smiths& contributions to botany : no. 76) Includes bibliographic references. SupL of Docs. no.: SI 1.2976 1. Aneilema-Morphology. 2. Aneilema-Classification I. Title. 11. Series. QKlS2741 no. 76 [QK495.C73] [584'.38] 581 s-dc20 90-9629 8 The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials 239.48-1984. Contents Page Preface and Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Morphology of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Introduction to Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Vegetative Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Shoots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Internodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Reproductive Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Inflorescences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Flowers in General. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Androecium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Staminodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Gynoecium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Capsules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Anatomy of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Introduction to Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Raphides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Stomata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Cytology of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Phytochemistry of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Reproductive Biology of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Taxonomy of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Introduction to Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Taxonomic Units . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Distribution of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 History and Typification of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . 55 Generic Relationships of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . 55 Citations of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Synonymy of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Description of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Section 1: Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Section 2: Amelina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 Division of Aneilema into Sections . . . . . . . . . . . . . . . . . . . . . . . . . 58 Key to the Sections of Aneilema . . . . . . . . . . . . . . . . . . . . . . . . . 59 Key to the Species of Section Aneilema . . . . . . . . . . . . . . . . . . . . 60 iv SMITHSONIAN CONTRIBUTIONS TO BOTANY Key to the Species of Section Amelina . . . . . . . . . . . . . . . . . . . . 61 Section 3: Rendlei. new section . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Key to the Species of Section Rendlei . . . . . . . . . . . . . . . . . . . . 64 1 . Aneilema taylorii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 2 . Aneilema usambarenre. new species . . . . . . . . . . . . . . . . . . . 69 3 . Aneilema rendlei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 4 . Aneilema brenanianum. new species . . . . . . . . . . . . . . . . . . 73 Section 4: Somaliensia. new section . . . . . . . . . . . . . . . . . . . . . . . 77 Key to the Species of Section Somaliensia . . . . . . . . . . . . . . . . . . 77 1 . Aneilema somaliense . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 2 . Aneilema granalibracteolatum. new species . . . . . . . . . . . . . . . 82 3 . Aneilema obbiadense . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 4 . Aneilema longicapsa. new species . . . . . . . . . . . . . . . . . . . . 85 5 . Aneilema pusillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Key to the Subspecies of Aneilema pusillum . . . . . . . . . . . . . . . . 5a . Aneilema pusillum subsp . pusillum . . . . . . . . . . . . . . . . . . 88 88 5b . Aneilema pusillum subsp . variabile. new subspecies . . . . . . . . . 88 5c . Aneilema pusillum subsp . gypsophilum. new subspecies . . . . . . . 88 5d . Aneilema pusillurn subsp . thulinii. new subspecies . . . . . . . . . . 89 90 1 . Aneilema petersii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 95 95 lb . Aneilema petersii subsp . pallidif2orum. new subspecies . . . . . . . 97 2 . Aneilema indehiscens . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 Key to the Subspecies of Aneilema indehiscens . . . . . . . . . . . . . . 102 2a . Aneilema indehiscens subsp . indehiscens . . . . . . . . . . . . . . . 102 2b . Aneilema indehiscens subsp . lilacinum . . . . . . . . . . . . . . . . 104 3 . Aneilema recurvatum. new species . . . . . . . . . . . . . . . . . . . 106 4 . Aneilema sebitense. new species . . . . . . . . . . . . . . . . . . . . . 110 5 . Aneilema forskalii . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 6 . Aneilema woodii. new species . . . . . . . . . . . . . . . . . . . . . . 118 6a . Aneilema woodii x forskalii . . . . . . . . . . . . . . . . . . . . . 120 7 . Aneilema benadirense . . . . . . . . . . . . . . . . . . . . . . . . . . 122 8 . Aneilema tanaense . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 9 . Aneilema calceolus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128 10 . Aneilema clarkei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 11 . Aneilema lamuense. new species . . . . . . . . . . . . . . . . . . . . . 133 12 . Aneilema succulentum. new species . . . . . . . . . . . . . . . . . . . 136 13 . Aneilema zebrinum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 Section 6: Brevibarbata. new section . . . . . . . . . . . . . . . . . . . . . . . 143 Key to the Species of Section Brevibarbata . . . . . . . . . . . . . . . . . . 144 Section 7: Pedunculosa. new section . . . . . . . . . . . . . . . . . . . . . . . 146 Key to the Species of Section Pedunculosa . . . . . . . . . . . . . . . . . . 146 Species of Uncertain Generic Position . . . . . . . . . . . . . . . . . . . . . . 147 Aneilema calandrinoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 Aneilema brmiliense . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 Appendix I: Rejected Generic Synonyms of Aneilema . . . . . . . . . . . . . . . . 151 Appendix 11: Index to Specific Epithets in Aneilema . . . . . . . . . . . . . . . . . 152 Appendix 111: Specimens Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159 Plates 1-7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . following page Section 5: Lamprodithyros . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 Key to the Species of Section Lamprodithyros . . . . . . . . . . . . . . . . Key to the Subspecies of Aneilema petersii . . . . . . . . . . . . . . . . . l a . Aneilema petersii subsp . petersii . . . . . . . . . . . . . . . . . . . 2c . Aneilema indehiscens subsp . kniense. new subspecies . . . . . . . . 104 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 74 Preface and Acknowledgments This monograph is based on a doctoral dissertation (Faden, 1975). It differs from the thesis in several significant ways: three major portions of the original have been only briefly summarized in the present work: a chapter on cytology, which will be published as a separate paper; a section on the history and typification of Aneilema, which has already been published (Faden, 1978b); and a chapter on the generic relationships of Aneilema, which will also be published separately. As a result of new collections, three species and four subspecies, all described as new, have been added to the 1975 treatment of the three monographed sections. Two of these three species, A. sebitense and A. woodii, were treated as inadequately known taxa in the dissertation under A. forskalii and A. petersii, respectively. The third, A. longicapsa, and the new subspecies A. indehiscens subsp. keniense and A. pusillum subsp. variabile, gypsophilum, and thulinii, were not recognized in the thesis. Among the other differences from the dissertation, the species descriptions and discussions have been shortened, and all new collections seen through 1986 have been added. For the sections that are not monographed, distributions of the species and a discussion about the naturalness of the section and taxonomic problems that remain have been added. Two of the species that were described as new in the dissertation, A. indehiscens and A. tanaense, were formally published subsequently (Faden, 1984) because the names were needed for the treatment of the family in the Flora of Southern Africa (Obermeyer and Faden, 1985) and other publications. Changes in the key to the species of section Brevibarbata and in the circumscription of A. schlechteri are the result of a study of that species and its relatives (Faden, 1984). The ncw species of Aneilema described and new combinations in other gencra made in the disscrtation were erroneously listed in Index Kewensis, supplement 17 (Davies, 1987) as having bccn validly published in 1975, bascd on an on-dcmand copy of the dissertation that was obtained by the library of the Royal Botanic Gardcns, Kew. That this listing was in error was acknowledged in the following (R.K. Brummitt, in litt., 3 March 1988): We are very surprised that your Commelinaceae names were taken up in I[ndex] K[ewensis]. It is certainly our policy now not to treat one-off copies produced from microfilm as effectively pubhshed, and I thought this policy had been applied for a long time. We consider that it was a mistake on our part that the names were included in Supplement 17, and we offer our apologies.. . . If you should re-publish any of h e m we would hst h e m again from the new publication with a note about the previous enuy. The first and hitherto only valid publication of any Aneilema species described in Faden (1975) was in Faden (1984). The rest are published hcrein. The names first cffectively published in Index Kewensis (Davies, 1987) as nomina nuda are sections Brevibarbata (as ?Brevibarbatum?), Pedunculosa (as ?Pedunculosum?), Rendlei and Somaliensia (as ?Somaliense?), and A. brenanianum, grandibracteolatum, lamuense, petersii subsp. pallidiflorum, recurvatum, succulentum, and usambarense. In order to avoid furthcr clutlcring up the litcralure, the Index Kewensis citations have bcen omitted from the text. With regard to combinations made in othcr gcnera in the disscrlation and mislakcnly listcd in Index Kewensis, thc situation is more complicatcd. The combinations listcd for Rhopalephora and attributed to Faden (1975) were first validly published in Fadcn (1977), except for R. monadelpha, which was abandoned in favor of R. micrantha (Vahl) Faden in the lattcr papcr and thus was nevcr validly publishcd. None of the combinations made in Dictyospermum in Faden (1975) and listed in Index Kewensis (Davies, 1987) has been validly published, nor will they ever be by me, for my generic concepts havc changed. V vi SMITHSONIAN CONTRIBUTIONS TO BOTANY ACKNOWLEDGMENTS I am greatly indebted to the many persons who have assisted in various aspects of this research. The following are gratefully acknowledged for having supplied me with living material of Aneilema or related genera: Dr. P. Andrews, Dr. B.G. Briggs, Dr. G. Davidse, Mr. D.B. Fanshawe, Dr. I. Friis, Dr. A. Gentry, Mr. M.G. Gilbert, Mr. J.B. Gillett, the late Mr. J.B. Hall, Mr. K. Inoue, Mr. A.B. Katende, Mr. 3. Lavranos, Dr. J.M. Lock, Mr. K.A. Lye, Dr. Gordon McPherson, Mrs. G.E. Organ, Mr. C.F. Paget-Wilkes, Mrs. J. Pawek, Dr. T P&s, Mr. M.G. Price, Dr. C.K. Rao, Dr. C.J. Saldanha, Mrs. J. Stewart, Mr. R.G. Strey, Dr. M. Thulin, the late Mrs. E.M. Tweedie, and Mr. J.R.I. Wood. Living materials were also supplied by the Royal Botanic Gardens, Kew, and Royal Botanic Gardens, Sydney. I wish to thank the directors of the following institutions for the loan of specimens (abbreviations after Holmgren, Keuken, and Schofield, 1981): B, BM, BOL, BR, BRI, C, CANB, COI, EA, F, FHI, FHO, FT, G, GC, GH, K, LISC, LMA, LUA, M, NDO, NH, NSW, NU, NY, P, PAV, PRE, PUC, QRS, RO, S , SRGH, UPS, US, VEN, WAG, Z. The curators of BM, EA, K, L, MAL, MPR, PRE, and Tsavo Research Centre Herbarium allowed me to study additional specimens at their institutions, The following are also thanked for their invaluable assistance: Dr. B.B. Chimphamba, Mr. D.B. Fanshawe, Dr. and Mrs. W.L. Handlos, Miss C.H.S. Kabuye, Dr. and Mrs. J.M. Lock, and Mr. and Mrs. A. Pooley, for extending hospitality and helping with the 1974 field work in Africa: Dr. W.H. Lewis, Dr. J. Dwyer, Dr. R. Andrews, Dr. A. Covich, and Dr. D.H. Nicolson, for reading part or all of the original thesis: the late Prof. J.P.M. Brenan, for his stimulating correspondence about Aneilema and other Commelinaceae and for his continuing interest in my research: Dr. Y. Suda, with whom the cytological studies were carried out jointly, for helping with all aspects of that work; the late Mr. P.R.O. Bally, Mr. P. Bamps, Mr. J.B. Gillett, Mr. J. Lavranos, and Dr. R.M. Polhill, for supplying geographic coordinates for collection localities; Dr. J. Dwyer, for assisting with some of the Latin descriptions: Dr. J. Nowicke, for preparing the scanning electron micrographs: the late Dr. J. Ridgway, for making available his Wild Dissecting Microscope: Mr. J. Lavranos, for supplying detailed ecological data for one of his Aneilema collections; Dr. P.H. Raven, for making all of the facilities of the Missouri Botanical Garden freely available to me: the government of Kenya, for permission to do research in that country. Reviewcrs Dr. W.C. Burgcr, Dr. D.R. Hunt, and an anonymous reviewer provided useful suggestions. Some of the line drawings were made by Mrs. Audrey J. Faden, Miss Tana Acton, Mr. Stanley Dolasinski, and Miss Alice Tangerini. Miss Susan Wiser and Mrs. Marie Uehling Kopp helped prepare the word processing copy. The insects were dctcrmined by Dr. C.D. Michener (bees) and the Commonwealth Institute of Entomology (all but the Hymenoptera). Financial support for field work in 1974 was provided by the National Science Foundation Doctoral Dissertation Improvement Grant No. GB-40817. I am particularly grateful to Dr. Walter H. Lewis for guiding this research, and to Mr. J.B. Gillett and Miss C.H.S. Kabuye, Botanists-in-Charge of the East African Herbarium during the period when I was employed there as a Peace Corps Volunteer (1969-1971), for allowing me ample time for field work. Finally, I must thank my wife Audrey J. Faden, without whose help, patience, and encouragement the original dissertation could not have been completed. The Morphology and Taxonomy of Aneilema R. Brown (Commelinaceae) Robert B. Faden Introduction The Commelinaceae are familiar to botanists of temperate North America largely through the native genera Commelina and Tradescantia and through a variety of tropical species commonly cultivated as houseplants and greenhouse ornamen- tals. Some Commelinaceae are used in introductory biology or botany laboratories to demonstrate cytoplasmic streaming and plasmolysis. Species with large chromosomes have been employed in research on chromosome structure and karyotype evolution. Certain Tradescantia clones have been used to test the mutagenicity of airborne substances (see Schairer et al., 1982); but overall, the members of this family are little utilized by man. Aneilema is an important genus in the Commelinaceae for several reasons. It is the second or third largest genus (after Commelina and Tradescantia, depending upon the latter?s circumscription), containing some 62 species. It is one of only six genera (out of 40) occurring in both the Old and New Worlds. It has been much confused taxonomically, having been circumscribed in many different ways by various authors (see Faden, 1978b,c). The purposes of this investigation were the (1) delimitation of Aneilema, (2) division of the genus into sections, and (3) monographic treatment of one or more sections. The circum- scription of Aneilema used here was arrived at through an intensive study of the morphology of all species in the genus. This was paralleled by in-depth investigations of related genera, which were then compared with Aneilema. The morphological studies of Aneilema led to its division into seven sections. Three sections comprising 22 species have been monographed. The large size of the genus prevented an exhaustive survey of the variation in many kinds of characters. Anatomy was looked into briefly and found not to be very promising for providing specific or sectional characters, with the exception of floral hair types and distributions. Cytology, on the other hand, proved Robert B. Faden Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. much more significant and was investigated more thoroughly. Those results will be reported in a separate publication. Materials and Methods Herbarium specimens of all species have been studied (see list of herbaria from which specimens were borrowed under ?Acknowledgments?). Living plants and preserved materials, including flowers, of Aneilema species were available as follows-the remaining species having been studied only from dried specimens: 1. Preserved material only: plagiocapsa (section Amelina); silvaticum (section Brevibarbata); richardsiae (section Pedun- culosa). 2. Preserved material and living plants only from cultiva- tion: acuminatum, biflorum, neocaledonicum (section Anei- lema); gillettii (section Amelina); longicapsa, pusillum subsp. gypsophilum, pusillum, and thulinii (section Somaliensia); forskalii, woodii, woodii x forskalii (section Lamprodithyros); lanceolatum subsp. lanceolatum, pomeridianum (section Bre- vibarbata). 3. Preserved material and living plants from the field and often also from cultivation: somaliense (section Somaliensia); brenanianum, rendlei, taylorii (section Rendlei); aequinoc- tiale, ephemerum, hockii, johnstonii, longirrhizum, nyasense (section Amelina); calceolus, clarkei, indehiscens (all subspe- cies), lamuense, petersii (both subspecies), recurvatum, sebi- tense, succulentum, tanaense, zebrinum (section Lamprodithy- ros); arenicola, beniniense, brunneospermum, dispermum, dregeanum, macrorrhirum, umbrosum, welwitschii (section Brevibarbata); chrysopogon, hirtum, leiocaule, minutijlorum, nicholsonii, pedunculosum, spekei, termitarium (section Pedun culosa) . The taxa for which living material was obtained only after the completion of Faden (1975) are forskalii, indehiscens subsp. keniense, lanceolatum subsp. lanceolatum, longicapsa, pomeridianum, pusillum subsp. gypsophilum, subsp. pusillurn and subsp. thulinii, sebitense, woodii, woodii x forskalii. To determine pollen viability, pollen grains from anthers of 1 2 SMITHSONIM CONTRIBUTIONS TO BOTANY mature buds, preserved in three parts absolute ethanol and one part glacial acetic acid, were stained with cotton blue. Pollen grains for scanning electron microscopic study were acetolyzed according to the procedure of Erdtman (1966), vacuum coated with gold, and examined and photographed with a Cambridge S tereoscan Mark IIa Scanning Electron Microscope. Capsule walls and seeds for SEM study were coated and examined as above. Seeds for light microscopic study were photographed with a Wild Dissecting Microscope. Stem and leaf transverse sections were made freehand from living and preserved material and stained with 0.01% meth- ylene blue for 30 to 60 minutes. They were then irrigated with 10% glycerine and stored as semi-permanent mounts. The Aneilema species investigated were as follows: acuminatum, neocaledonicum (section Aneilema); taylorii (section Rendlei); somaliense (section Somaliensia); aequinoctiale (section Ame- lina); petersii subsp. pallidiflorum, succulentum, zebrinum (section Lamprodithyros); spekei (section Pedunculosa). Paired petal color was determined by means of an R.H.S. Colour Chart (1966). Comparisons with the color patches of the chart were made in daylight in bright shade, almost always by two observers. All paired petal colors noted in the population were recorded. Floral hairs were studied from preserved mature buds or, occasionally, preserved or fresh flowers. The buds or flowers were dissected into separate organs, which were mounted whole or slightly macerated, when necessary. These were stained with 0.01% methylene blue and irrigated with 10% glycerine as above. Glandular cells stained darkly in less than 60 seconds. Other cells stained weakly or not at all, even after 60 minutes. The species examined are listed in Tables 3,4, and 5. Buds for meiotic chromosome counts were collected in the field or from cultivated plants. Root tips for mitotic counts were collected from greenhouse grown plants. The procedures followed were those of Fadcn and Suda (1980). Species descriptions were drawn up using a variety of materials, including herbarium specimens, and (when avail- able) living plants, preserved flowers and inflorescences, separately collected capsules and seeds, detailed descriptions of morphology made from living plants, and color transparencies. Of the 22 species studied in detail, 14 were seen in the field and were latcr cultivated, another four were cultivated from material obtained from other collectors, and only four (A. usambarense, A. grandibracteolatum, A. obbiadense, A. bena- dirense) had to be studied exclusively from herbarium spccimcns. Among the 40 species belonging to sections that are not monographed, 28 were obscrved in the field or under cultivation, and the rest had to be studied from only dried or preserved specimens. One hundred and twenty-one populations representing 32 Aneilema species plus four infraspecific taxa were observed in the field in Africa (Ghana, Kenya, South Africa, Zambia, Malawi, and Tanzania) from January to August 1974. Specimens were collected and ecological notes made for every population. When possible, detailed notes were made of vegetative and reproductive morphology, including paired petal color, on specially designed data sheets. Further notes were made on the reproductive condition of the plants, e.g., whether mature capsules were present, flowering times, and the presence of pollinators, which were collected whenever possible. Color transparencies were taken of the flowers and sometimes of the inflorescence and habit using a Miranda DR single-lens reflex camera, Kodachrome I1 35 mm film, a Honeywell Strobonar 100 electronic flash and extension tubes for close-ups. In addition to dried specimens, materials frequently collected included. flowers and vegetative organs preserved in FAA, separate capsules and seeds placed in seed packets, buds fixed for cytological study, and cuttings or whole plants for later cultivation. Plants were cultivated at the Missouri Botanical Garden until 1976, at the University of Chicago 1976-1980, and at the Smithsonian Institution Department of Botany greenhouse from 1980 until the present. Most plants studied in cultivatim were derived from the 1974 field collections. Other materials for cultivation came from earlier field collections made by the writer, from the writer?s 1977 Kenyan collections, from plants or seeds collected by various persons at the writer?s request (see ?Acknowledgments?), and from materials in cultivation at botanical gardens. These plants were described and photo- graphed as above. Flowers and other materials were collected from the cultivated plants for later study. Preserved flowers of some species were studied at the Royal Botanic Gardens, Kew, or obtained on loan from that institution. Determinations of habit types have been based largely on field notes and living plants. Descriptions of vegetative and inflorescence morphology have utilized dry specimens for the most part, being supplemented by data from living plants, when available. In contrast, whenever possible, floral descriptions have been derived almost entirely from living and preserved materials, with data added from field notes and color transparencies. When such material was lacking, pressed flowers (or their remains) and buds, soaked in a solution of Aerosol OT and usually dissected in situ, provided the basis for the descriptions. In order to avoid ambiguity, the manner of measuring certain vegetative and floral parts has been described under the relevant structures. All specimens seen through 1986 of the species in sections Rendlei, Somaliensia, and Lamprodithyros have been cited under the species (also see Appendix 111). The countries are listed from west to east and north to south. For political subdivisions, listed alphabetically within the countries, the Flora of Tropical East Africa (1952-) has been used for Uganda, Kenya, and Tanzania, Flora Zambesiaca (1960-) for Mozambique, and the Times Atlas of the World (1971) for other countries. For a few countries, political subdivisions were unclear and have been omitted, The system proposed by NUMBER 76 3 Edwards and Leistner (1971) has been used for citing South African collections. Only specimens seen have been mapped. All lectotypes are those of the writer unless otherwise stated. Morphology of Aneilema I may here mention, for the encouragement of parties who may have an opportunity of collecting specimens, that I have learned in the course of their investigation, that much more can be done with dried specimens than I previously supposed possible, and would therefore urge their collection, as I feel quite convinced that the Commelynaceae is much richer in species than the latest publications would lead one to suppose. (Wight, 1853:28.) INTRODUCI'ION TO MORPHOLOGY The Commelinaceae is one of the most awkward families to study using dried specimens because the deliquescent flowers are normally poorly, if at all, preserved. The difficulty in making use of such material is no doubt a contributing factor to the paucity of modern monographic studies in the family. In the present investigation, living material has been utilized when- ever possible. However, the large number of taxa surveyed has necessitated an examination of numerous herbarium speci- mens. For a number of species this has been the only kind of material available. Like Wight (1 853), I have been pleasantly surprised as to how much could be learned from dried specimens, particularly from those with mature flower buds. The genera of Commclinaceae are defined largely on the basis of morphological characters. The total morphological varialion in Aneilema has never been studied. The most useful contributions have been those of Bruckncr (1926, 1930), Brenan (1952), and Morton (1966). These studies have either been limited to a few characters (Bruckner, 1926. 1930; Brenan, 1952) or have been geographically biased and the generalizations made not always applicable to the genus as a whole (Morton, 1966). In the present account, an attempt has been made to examine the morphology of all species in the genus. The extent to which each species has been studied depended upon the type of material available and whether the species belonged to one of the sections selected for detailed taxonomic investigation. All aspects of morphology have been examined, but greatest emphasis has been placed on floral and seed morphology. VEGETATIVE MORPHOLOGY HABIT.-Aneilema shows great diversity in habit. I recog- nize 17 habit types (Figures 1-3), but the boundaries between adjacent types are not always clean-cut. The basic type (IA) is a decumbent perennial in which lhe main shoot is overtaken by lateral shoots that in turn are overtaken by other lateral shoots. The decumbent perennial has given rise to a variety of other habit types: less decumbent to ascending or erect perennial or annual (IB and IC); mat-forming perennial or annual (IIA1 and IIB); rhizomatous perennial with fibrous roots (IIA2); long- trailing, sometimes looping or scrambling perennial (IIA3); ascending perennial that produces occasional long-trailing shoots that give rise to new plants (IIA4). The erect to ascending perennials and annuals (IC) have themselves produced two major lines of habit types. One is a series of reduced annuals (IIIA, IIIB, and IIIC) that secondarily may become decumbent again (IIIB). The other line consists of tuberous-rooted perennials. Among these the basic type (IVA) has sessile root tubers, a single or sometimes two subterranean growing points that persist only one or two growing seasons, and a moderately well-branched, ascending shoot system. From this type have arisen forms with stalked, distal tubers (IVBl), more persistent shoot bases that become shortly rhizomatous (IVB2), or reduced shoot systems (IVB3). In the type with stalked tubers, the shoots may sometimes root at the lower nodes (IVC1). The stalked tuber type has also given rise to forms with dimorphic vegetative and reproductive shoots (IVC2). The reproductive shoots are sometimes produced before the vegetative shoots during the growing season. Varying amounts of dieback occur in the perennial species at the end of the growing season. In the decumbent to erect, fibrous-rooted species, the lower portions of the shoots normally survive and produce new growth from axillary buds during the next growing season. In species with subterranean storage organs, the entire aerial shoot system or the greater part of it (depending upon the species) dies during the dry season. The shoots may gradually die back like the fibrous-rooted perennials or they may abscise at the base. It is sometimes difficult to determine whether a plant is an annual or a perennial. As used here, annuals are monocarpic plants that, under natural conditions, live less than a year. Perennials live more than one year and have the potential to flower and fruit more than once. Two examples will serve to illustrate the problem in applying these definitions. Aneilema petersii subsp. pallidiflorum, new subspecies, may be distinctly annual. However, when the plants produce shortly decumbent shoots, which they often do, they may appear perennial, even in the field. From prolonged observations of populations in the Tsavo area of Kenya, it was learned that even these perennial-appearing plants are monocarpic and die after fruiting. Fitting these plants, however, into the above definition of an annual is complicated by the fact that they normally require two growing seasons, at least in this area, to complete their life cycle. The seeds germinate during the short rains (November to December) but, typically, the plants do not flower then. Despite the absence of clearly adapted water storage organs, many plants survive a three-month dry season, flower and fruit during the long rains (April to May), and then die. Since the entire life cycle does occur within 12 months (although over two calendar years), and there is only one flowering period, the plants are considered annuals. Plants of Aneilema sebitense, new species (Gilbert & Thulin 266) and A. forskalii (Gilbert & Thulin 969), when grown from seed, developed identically, rapidly producing numerous 4 SMITHSONIAN CONTRIBUTIONS TO BOTANY ep H I:, ,,> \ ,,,*p cp w ... . . . . . . . I:.:. i:: U w U H NUMBER 76 5 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY m m W NUMBER 76 7 shoots with terminal and axillary inflorescences, a characteris- tic of annuals; but whereas the plants of A. forskalii, proved to be annual by continuing this pattern of growth and eventually losing vigor and dying, those of A. sebitense began to produce strong vegetative shoots and have been cultivated continuously since 1976. From this evidence and later cultivation of plants from two additional populations of A. forskalii (Wood 1997 and 2895) and one of A. sebitense (Faden & Faden 771803 and 77/803A), it was concluded that A. forskalii is always annual. On the other hand, although A. sebitense is perennial, young plants appear capable of functioning as annuals during very short rainy seasons. Plants of this species may thus be considered perennials or facultative annuals. The perennial habit of this species would be entirely unpredictable, based solely on the morphology of the young plants. Perennials and annuals are known to occur within five species of Aneilema. In A. petersii and A. paludosum, the pcrennials are considered sufficiently differentiated to be trcatcd as separate subspecies. In A. taylorii, A. tanaense, and A. calceolus, they are not distinct enough to merit taxonomic recognition. In A. taylorii the perennials occur at higher clcvations and probably in moister situations than the annuals. In the other two spccics the pcrennials occur as peripheral populations in otherwise annual spccics. Most other species show rclatively little variation in habit whcn only mature plants are considered. In the course of dcvclopmcnt they may exhibit other habit types transitionally. For example, many decumbent perennials are erect to ascend- ing in their early stages. Other variation, such as the degree of branching exhibited by annuals, may be caused by edaphic factors or the length of a particular growing season. An exception is A. hockii in which fibrous-rooted decum- bent, fibrous-rooted rhizomatous, and tuberous-rooted, ascend- ing pcrcnnials occur. A great deal of variation in the amount of branching is also prescnt within some of these types. Further study is rcquired to dctcrmine how to treat these forms of A. hockii taxonomically. For convcnience, the 17 habit types recognized can be rcduccd to three gcncral categories: pcrcnnials lacking special- ized, Subterranean storage organs, pcrcnnials with such organs, and annuals. The Aneilema spccics with each type are as follows: Pcrcnnials lacking spccializcd, subterranean storage organs: acuminatum, biflorum, neocaledonicum (section Aneilema); aequinoctiale, hockii, nyasense (section Amelina); rendlei, taylorii, usambarense (section RendleQ; calceolus, clarkei, indchiscens, petersii subsp. petersii, recurvatum, new species, sebitense, succulentum, ncw species, tanaense, woodii, new spccics, rebrinum (section Lamprodithyros); beniniense, dis- permum, dregeanum, silvaticum, umbrosum (section Brevibar- bata); leiocaule (scction Pedunculosa). Pcrcnnials with spccialized, Subterranean storage organs: sclerocarpum, siliculosum (scc Lion Aneilema); gillettii, hockii, johnstonii, longirrhizum, plagiocapsa (scction Amelina); bre- nanianum, new species (section Rendlei); longicapsa, new species, obbiadense, pusillum, somaliense (section Somalien- sia, new section); homblei, lanceolatum, macrorrhirum, palu- dosum subsp. pseudolanceolatum, pomeridianum, setiferum, welwitschii (section Brevibarbata, new section). Annuals: aparine? (section Aneilema); ephemerum (section Amelina); calceolus, forskalii, lamuense, new species, petersii subsp. pallidiflorum, tanaense (section Lamprodithyros); an- golense, arenicola, brunneospermum, mortonii, paludosum subsp. paludosum and subsp. pauciflorum, schlechteri (section Brevibarbata); chrysopogon, hirtum, minutiflorum, nichol- sonii, peduncul osum, richardsiae, spekei, termitarium (section Pedunculosa). Species of unknown habit: grandibracteolatum, new species (section Somaliensia); benadirense (section Lamprodithyros). ROOTS.-ROO~S in mature plants are all adventitious. Seedlings of A. umbrosum have been observed to develop such roots when only 24 hours old, These adventitious roots rapidly overtake the radicle. In older plants adventitious roots are produced from nodes-often several from a node-that are close to or in contact with the substrate. In some species they are produced only from subterranean shoots. Roots may be fibrous or variously thickened. The latter are collectively termed tuberous in this account. Tuberous roots may be of uniform thickness for their entire length, e.g., A. longirrhizum, but, more commonly, they taper from their bases or at both ends. The fusiform tubers are sessile in some species, e.g., A. pusillum, and shortly to longly stipitate in others, e.g., A. lanceolatum. Stipitate, tuberous roots have been found to reach a length of 60 cm (including the stipe) in A . somaliense. Aneilema hockii and A. paludosum are the only species in the genus that include both fibrous-rooted and tuberous-rooted plants. In A. hockii the type of root system has been found to be under genetic control and is maintained in cultivation. Cuttings from each typc produce the same kind of roots as the plant from which they were taken. Tuberous root systems have also been reproduced by shoot cuttings of A. brenanianum, A. gillettii, and A. welwitschii, all tubcrous-rooted species. Aerial roots have bcen observed on a cultivated plant of A. somaliense. Short lateral shoots produced such roots from their bases. The thus formed planllets were easily detachable from the main shoots and could function as independent plants. This peculiar vivipary has not been observed in herbarium speci- mens of A. somaliense nor in any other Aneilema species. It must therefore bc considcred an artifact of cultivation. SIIO0TS.-In mature plants of most Aneilema species there is neither a distinct main shoot nor a single, clear, sympodial axis, but rather a diffuse branching, which is here termed a diffuse shoot system. Such branching systems were considered sympodial by Bruckner (1926), who used the term for any shoot system that was not distinctly monopodid. Aneilema beniniense and A. aequinoctiale are examples of species with diffuse shoot systems. Sympodial shoots do occur in Aneilema, but they are 8 SMITHSONIAN CONTRIBUTIONS TO BOTANY uncommon. They are largely confined to rhizomatous species such as A. brenanianum and A. obbiadense, in which the rhizomes are sympodial, being composed of the bases of successive lateral shoots. This type of branching never results in leaf-opposed inflorescences, as in Commelina and a few other genera of Commelinaceae with sympodial shoots, because the sympodial growth occurs at the base of the flowering shoot, rather than at the apex. Distinctly monopodial shoots systems are also infrequent in Aneilema. They occur only in annuals and tuberous-rooted perennials with reduced shoot systems. Even in these species, the main shoot tends to flower and die before the lateral shoots and is, in that sense, overtaken by them. Monopodial shoot systems are often the result of poor growth conditions: plants of the same species growing in more favorable situations may produce diffuse branching systems. Taxa in which monopodial shoot systems have been noted include A. hockii, A. petersii subsp. pallidijlo- rum, A. nicholsonii, and A. termitarium. The shoots of Aneilema species vary from monomorphic to strongly dimorphic. In the monomorphic types there is no differentiation into vegetative and reproductive shoots. In other species, e.g., A. neocaledonicum and A. calceolus, the flowering shoots differ from the vegetative shoots only in being erect or ascending while the latter are repent. In A. clarkei, which has a similar habit to the above named species, the leaves on the vegetative shoots are uniform in size and shape; those on the flowering shoots are very diverse (see p. 133). Strongly dimorphic shoots are produced in only a few species. The flowcring shoots in A. lanceolatum subsp. subnudum have very reduced leaves, while the vegetative shoots, which are often produced later in the season (Morton, 1966), have distinctly longer leaves. In A. pomeridianum the flowcring shoots are elongate and have relatively small leaves; the vegctative shoots form basal rosettes of longer leaves (see habit type IVC2, Figure 3). Lateral shoots arise from axillary buds within sheaths. Such shoots may emerge from the tops of the sheaths or they may perforate them. Emergent or non-perforating shoots sometimes cause the sheaths to split more or less irregularly along the fuscd cdge. Pcrforating shoots are more common than non-pcrforating oncs and characterize species of scctions Aneilema and Lamprodilhyros. Non-perforating shoots are typical of scctions Brevibarbata and Pedunculosa. Within plants of many spccics normally having one shoot type, a few shoots of the othcr type may be present. A latcral shoot always has a bladeless sheath or prophyll at its base. In some Aneilema species an axillary bud within the prophyll may give rise to another shoot that in turn has a prophyll at its base. Successive shoots may arise in this fashion and produce a cluster of up to seven shoots at a node. Shoots arising from prophyllar buds have been noted in 11 Aneilema spccics (arenicola, brunneospermum, calceolus, hirtum, inde- hiscens, minutiflorum, pedunculosum, spekei, tanaense, termi- tarium, zebrinum), nine of which are annuals. It is likely to be found in most annual species of the genus. Such branching has previously been reported in the Commelinaceae only in Tinantia (Tomlinson, 1969). Leaves are produced on the lateral shoots above the basal prophyll in a distinctive sequence of shapes and sizes. The first leaf always has a very small and broad lamina. Successive leaves show a gradual increase in total length and width and in the ratio of length to width. In most species a petiole is also developed by about the third or fourth leaf above the prophyll. Successive leaves have longer petioles. Towards the inflorescence there is another sequence of leaf shapes and sizes. This transition may be abrupt or gradual and always includes some reduction in leaf length and elimination of the petiole (when present in the lower leaves). The uppermost leaf below the inflorescence is sometimes highly modified and is termed the inflorescence bract in this account. It is discussed in more detail under the inflorescence. The middle internodes of the lateral shoots are generally the longest. The upper are often very strongly reduced. A sequence of leaves and internode lengths on a lateral shoot of Aneilema umbrosum subsp, ovato-oblongum is shown in Figure 4. In most Aneilema species there is a region of the lateral shoots, between the basal and apical transition zones, in which leaf size and shape and internode length are relatively uniform. This region may be short or long, according to the species. The sequence of leaf shapes and sizes and internode lengths in the lateral shoots is very similar to the sequence that occurs in the main shoot. The prophyll of the lateral shoot is analogous (or homologous?) to the cotyledonary sheath at the base of the main shoot in the seedling. Leaves in Aneilema are always alternate and are either spirally or distichously arranged on the shoots. The spiral arrangement is much more common and is the sole type present in sections Aneilema, Rendlei, Brevibarbata, and Pedunculosa. In some species of section Aneilema, e.g., A. neocaledonicum, the leaves of the repent vegetative shoots are sometimes two-ranked, but they are clearly spirally attached to the stems. Both distichous and spiral arrangements occur in sections Amelina, Somaliensia, and Lamprodithyros. The species in which distichous leaves have been noted are A. aequinoctiale, A. hockii, A. longirrhizum, and A. nyasense (section Amelina); A. pusillum (section Somaliensia); and A. calceolus, A. clarkei, A. lamuense, A. petersii, A. succulentum, A. tanaense, and A. zebrinum (section Lamprodithyros). The constancy within a species of the distichous arrangement is variable. In A. aequinoctiale, A. nyasense, A. calceolus, A. clarkei, A. lamuense, A. succulentum, and A. zebrinum, all lateral shoots always have distichously arranged leaves. This is also the case for most shoots of A. tanaense. In some shoots of this species, however, the arrangement changes to spiral towards the inflorescences. Plants of A. petersii with distichous leaves on the lateral shoots have been noted in only two populations; it is unknown whether all plants of these populations have this leaf arrangement. In A. hockii relatively NUMBER 76 9 5 P FIGURE 4 . S e q u e n c e of leaf shape and size and internode length in a lateral shoot of Aneilema umbroswn (Vahl) Kunth subsp. ovato-oblongum (P. de Beauvois) J.K Morton (p = prophyll). From Lye & Katende 6235 in cultivation. (Bar = 1 cm.) few populations have distichous leaves, and none has exclu- sively this arrangement. In plants of these populations many individual shoots have a mixture of spiral and distichous arrangements. The last situation also obtains to the single population of A. longirrhizum seen in the field. The distichous leaf arrangement is commonly associated with the mat-forming habit in such species as A. succulentum and A. zebrinum. However, A. aequinoctiale and the popula- tions of A. hockii with this leaf arrangement have erect to decumbent shoots that do not form mats. Furthermore, other mat-forming species, e.g., A. neocaledonicum and A. bijlorum, have spirally arranged leaves. Seedlings of seven of the 10 species that typically or occasionally have a distichous leaf arrangement on the lateral shoots have been observed to have, as the normal condition, a spiral leaf arrangement on the primary shoot. In A. lamuense, A. petersii, and A. tanaense, the spiral arrangement is maintained right up to the inflorescence. In A. calceolus the mainshoot could not be identified with certainty in the mature plant. This probably is indicative that the leaf arrangement of the main shoot changes to distichous when the shoot becomes repent. The seedlings of A. aequinoc- tiale, A . succulentum, and A. zebrinum have not been followed to maturity. The presence of only spiral leaf arrangements in the primary shoot of all 23 Aneilema species in which seedlings have been observed (acuminatum, aequinoctiale, arenicola, beniniense, brunneospermum, calceolus, ephemerum, hirtum, indehiscens, johnstonii, lamuense, minutiflorum, pedunculosum, petersii, rendlei, spekei, succulentum, tanaense, taylorii, term'tarium, umbrosum, welwitschii, zebrinum), and the subsequent devel- opment of a distichous arrangement in the lateral shoots of some of these species support the hypothesis that in the Commelinaceae the spiral arrangement (radial symmetry) is the primitive condition and the distichous arrangement (dorsiven- tral symmetry) the derived one (Clark, 1904). The data for Aneilema do not support the hypothesis of Tomlinson (1966) that the distichous arrangement is the juvenile state and the spiral arrangement the adult condition. INTERNODES.-Internodes are generally terete, succulent, and green. In A. succulentum they are sometimes laterally compressed. In all species of dry habitats, the internodes of persistent aerial shoots become shriveled during the dry season. The fleshiness of the internodes also varies according to the species. The Australo-Oceanic species have less succulent internodes than the African species. Internodes in some Aneilema taxa may be striped or suffused with purple, the color sometimes becoming more pronounced during the dry season. LEAVES.-Aneilema leaves are typical of the family and are always simple with unlobed, entire-margined, parallel-veined laminae. The sheaths are closed except occasionally in the uppermost leaves of the flowering shoot where they may be only slightly fused basally or even completely free. The formation of a lateral shoot sometimes causes the sheath to split. Commonly the sheaths persist after the laminae have died. In a few species, e.g., A. brenanianum, the sheaths are very long and completely cover the lower portions of the flowering shoots, protecting them from desiccation during the dry season. In most species the sheaths are ciliate at the apex. Laminae are generally herbaceous and pubescent on both surfaces and on the margins. In a few species the laminae are somewhat succulent, but except in A. pusillum, they are not as fleshy as certain species of other genera of Commelinaceae, such as Cyanotis, Murdannia, and Tradescantia. The laminae in Aneilema have symmetric bases (except A. succulentum), 10 S M I T H S O W CONTRIBUTIONS TO BOTANY which in many species are narrowed into petioles. Although these are not true petioles in the sense used for dicotyledons, the term ?petiolate? is employed here to conform with common usage. Generally the laminae are entirely green, but in some species they may become suffused or mottled with purple or maroon, particularly in the dry season. In many species the laminae are discolorous, the abaxial surface being paler or sometimes gray-green. Stomata are present predominantly or exclusively on the abaxial surface. The great variation in lamina shape and size present in every individual plant of Aneilema as a result of the developmental sequences in the shoots would tend to obscure specific characters and interspecific differences if the full range of variation were included in the species descriptions. Therefore, descriptions of the leaves are based entirely on the larger laminae present on the shoots. Lamina length always includes the pctiole, when present. Mature leaves in all but two Aneilema species that have been seen alive have involute vernation. The exceptions are A. pomeridianum, which consistently had convolute (including supervolute) vernation, and A. pusillum, the very succulent leaves of which were slightly conduplicate to nearly flat. In many species, the first leaf of the seedling has a convolute vernation. This has been observed in single populations of A. calceolus (four seedlings), A. ephemerum (four), A. johnstonii (seven), A. lamuense (one), A. pedunculosum (eight), A. spekei (two), A. tanaense (34), and A. termitarium (four), and in two populations of A. hirtum (total of four seedlings). Solely involute vernation in the first seedling leaf has been recorded in single populations of A. acuminatum, A. aequinoctiale, A. beniniense (-10 seedlings), A. petersii subsp. pallidiforum (four), A. umbrosum subsp. ovato-oblongum, and A. wel- witschii (two). Both types of vernation have been found in two populations of A. welwitschii (one seedling with each type in both populations) and in one of A. minutiforum (three seedlings with involute vernation, one with convolute). In all of the above species, except A. wefwitschii, the second and every subsequent leaf of the seedling have the involute vernation of mature plants. In a few seedlings of A. welwitschii the second seedling leaves had a convolute vernation, but succcssive leaves all had the adult type. The significance of the above observations is unclear. The two types of vernation prcscnt in the seedlings do not separate species along sectional lines. Involute vernation is clearly the usual type in mature plants of nearly all Aneilema species, so the convolute vernation observed in the first leaves of some sccdlings may be considered a developmental stage. Perhaps it results from the very narrow lamina in the seedlings. The only species in which convolute vernation has been noted in mature plants, A. pomeridianum, also has a very narrow lamina, but so does A. brenanianum, which consistently has involute vema- tion. REPRODUCTWE MORPHOLOGY INFLORESCENCES.-The basic type of inflorescence in Aneilema is a terminal thyrse consisting of several to many cincinni attached to a central axis, the inflorescence axis (Figure 5; Plates lg, 2c-e,m,o, 4i,m,o). The designation of this raceme of cincinni, the Wickeltraube of Eichler (1875), as a thyrse follows the usage of Clarke (1881a), Pichon (1946), Brenan (1966). and Faden (1974). Most authors, however, have termed such inflorescences panicles (Bentham, 1849; Bentham and Mueller, 1878; Bentham and Hooker, 1883; Schumann, 1895; Clarke, 1897, 1901; Hutchinson, 1934, 1959; Morton, 1966). The differences are not due to varying interpretations of the inflorescence morphology but rather to diverse uses of ?panicle? and ?thyrse.? The definition of a thyrse (or thyrsus) as a ?panicle-like cluster with main axis indeterminate and the lateral axes determinate? (Lawrence, 1951) has been followed here. Aneilema inflorescences conform to this definition in that the cincinni flower in an acropetal sequence: therefore the inflorescence axis is indeterminate. The cincinni themselves are determinate, because every flower is morphologically terminal. Aneilema thyrses show great variation in size, density, number and arrangement of the cincinni, and in position on the plant. Thyrses are commonly 2-15 cm long, but they can reach a length of at least 25 cm in A. aparine and A. gillettii. The number of cincinni per inflorescence has a characteristic range for each species and can be as many as 55 (or perhaps more in some forms of A. beniniense). In a few species, e.g., A. clarkei, every thyrse is composed of a single cincinnus (Plates 3p, 4e,g). The density of the inflorescence is a function of the length of the inflorescence axis, the number of cincinni attached to it, and the length of the cincinni. The terms lax, moderately lax, moderately dense, and dense are used in this account and are illustrated in Figure 6. Lax or moderately lax thyrses characterize sections Amelina and Aneilema, while moderately lax to dense inflorescences are typical of the remaining sections, the densest thyrses occurring in sections Brevibarbata and Pedunculosa. The larger thyrses-those more than 10 cm long-are always lax. The cincinni have a basically alternate arrangement on the inflorescence axis. In some species, e.g., A. aequinoctiale and A. johnstonii, some or all of them may appear opposite or verticillate, but closer inspection and observations of their sequence of flowering reveal that they too are truly alternate. Therefore, such arrangements have been termed subopposite and subverticillate here. The three basic types of cincinnus arrangement are illustrated in Figure 7. Moderately lax to dense inflorescences usually have predominantly alternate cincinni; lax thyrses commonly have mainly subopposite or subverticil- late cincinni. Not infrequently all three arrangements may occur within a single inflorescence. Thyrses terminate the main shoots in nearly all Aneilema species, They are also terminal on lateral shoots of various lengths in most species. The lateral, inflorescence-terminated NUMBER 76 11 i. a. P. ? L- 1 c . b. br. c . a. c . p. FIGURE 5.-Diagram of the structure of the Aneilema inflorescence (br. = bracteole, c.a. = cincinnus axis, c.b. = cincinnus bract, c.p. = cincinnus peduncle, i.a. = inflorescence axis, i.b. = inflorescence bract, p = peduncle). shoots may be elongate and have normal foliage leaves, or abbreviated and bear only reduced leaves or bracts below the inflorescence. These shoots may perforate the sheaths or emerge from their mouths. Only A. zebrinum has strictly axillary inflorescences (Plate 4g). The abbreviated, inflorescence-terminated, lateral shoots that bear only bracts or very reduced leaves below the inflorescence are the ?cyme-branches? of Clarke (1901:73) and ?inflorescence-shoots? of Forman (1962). These differ from strictly lateral inflorescences, according to Forman (1962), by having a basal prophyll. By applying that criterion, there are no truly lateral inflorescences in Aneilema, even in A. zebrinum, only inflorescence-shoots. However, it is questionable whether a special term like ?inflorescence-shoot? is desirable for a condition that is merely one end of a continuous range of variation in size and development of these inflorescence- bearing shoots. Often the whole range may be found on a single plant. Furthermore, whether or not a basal prophyll is present-ne may ask if the prophyll can ever be lost by reduction-the inflorescence is still a shoot system and it is still lateral. From the above discussion it is obvious that the distinction among an inflorescence terminal on a lateral shoot, an inflorescence-shoot, and a lateral inflorescence derives partly from the length and degree of development of the shoot and partly as a matter of definition. In the present account, an ?inflorescence terminal on a lateral shoot? is applied only to the leafy shoots that are normally produced at least several nodes below the terminal inflorescence of the main shoot. The terms lateral inflorescence and inflorescence-shoot have been used interchangeably for the very reduced inflorescence-terminated shoots that may be produced from leaf a i l s anywhere along the stem, The term secondary inflorescence has sometimes been employed for an inflorescence-shoot arising from the inflores- cence bract. Although inflorescence-shoots may be produced from any node in some species, e.g., A. zebrinum and A. petersii subsp. pallidiflorum, more commonly they are restricted to the uppermost few nodes below the terminal thyrse. Under these circumstances they usually form loose associations with that inflorescence. In a few species, e.g., A. indehiscens and A. umbrosum, the association of the terminal thyrse and lateral inflorescence-shoots is so close that the whole group may be treated as a larger inflorescence unit (Brenan, 1966). This is the Synfloreszenz of Troll (1961). The uppermost leaf on the flowering shoot is often highly modified, always differing from the foliage leaf below it in size and sometimes in shape, texture, and pubescence. It varies from foliaceous and herbaceous to bract-like and membranous; its lamina is always sessile. In some species, e.g., A. pedunculo- sum and A. spekei, it completely encloses and protects the young, pre-flowering inflorescence. In A. clarkei, A. succulen- rum, and A . lamuense, it surrounds the inflorescence at all stages. In A. petersii subsp. pallidiflorum it is often erect and serves as a background for the white or whitish flowers (Plate 20). In these species and others the uppermost leaf is clearly more of a reproductive structure than a vegetative one. For this reason it is called the inflorescence bract in this account. For consistency the same terminology has been applied to all Aneilema species, including those in which the uppermost leaf is not highly modified. Because the uppermost leaf is considered a reproductive structure, the peduncle, which is almost always present, is treated as the sum of the lengths of the two uppermost internodes below the inflorescence, i.e., the distance between the node of the second leaf below the inflorescence and the lowermost cincinnus (Figure 5) . As in all Commelinaceae, the basic unit of the inflorescence in Aneilema is the cincinnus. The cincinnus has usually been considered a 12 SMITHSONIAh! CONTRIBUTIONS TO BOTANY t t a b C d FIGURE 6.-Density of the Aneilema inflorescence: a, lax; b, moderately lax; c, moderately dense; d, dense. a b C FIGURE 7.4incinnus arrangements in Aneilentu inflorescences: a, alternate; 6, subopposite; c, subverticdlate. scorpioid cyme (e.g., Brenan, 1966), but Lawrence (1951) has cincinnus bract. Cincinnus bracts commonly have subapical termed it a helicoid cyme, noting that such cymes are often glands that appear to be functional only in the very young, mistakenly called scorpioid. It is beyond the scope of this pre-flowering inflorescence. In some species of section discussion to determine whether the monochasial cymes in the Pedunculosu each cincinnus bract has a linear apex terminated Commclinaceae are helicoid or scorpioid: they will simply be by a clavate gland. In a few Aneilemu species, e.g., A. forskalii, called cincinni, as they have been by most authors. glands are sometimes present along the margins of the Every cincinnus is subtended by a persistent bract, the cincinnus bracts. NUMBER 76 13 The cincinnus consists of a sympodial axis along which two ranks of bracteoles are arranged (Figure 8). The most basal internode of the cincinnus, that between the inflorescence axis and the first bracteole, is commonly much longer than the others and is called the cincinnus peduncle. (The term stipe has been used for this structure by Hunt (1986), but I prefer cincinnus peduncle because it is less ambiguous.) The remaining internodes collectively constitute the cincinnus axis. Both the cincinnus peduncle and axis are usually elongate, but in some species, the latter or both may be much abbreviated or suppressed. No species has very reduced cincinnus peduncles with elongate cincinnus axes. The number of flowers that will be produced by a cincinnus is essentially indeterminate, but it may regularly be few or even one in some species. Persistent, pedicel-opposed bracteoles are present in all Aneilema species. They are usually amplexicaul and commonly cup-shaped and perfoliate. When amplexicaul they enclose and protect the cincinnus apex and young buds. In nearly all species every bracteole has a subapical gland. In a few taxa, e.g., A. forskalii, smaller glands are also present along the bracteole margin. Those species of section Pedunculosa that have cincinnus bracts terminating in clavate glands also have similarly shaped bracteoles. In the present account, bracteole length is considercd the length of the bracteole midrib. For those species with perfoliate bracteoles, a second measurement, bractcole height, is the length of the fused edge. The morphology of nearly all Aneilema inflorescences can f* --r( ped. b. c. a. 1 ~ (2, I \ I I/ c. b. --ti%-- i. a. FIGURE &-Diagram of an Aneilema cincinnus. (The broken lines represent the positions of previous flowers; b. = bud, br. = bracteole, c.a. = cincinnus axis, c.b. = cincinnus bract, c.p. = cincinnus peduncle, f = position of flower, i.a. = inflorescence axis, ped. = pedicel.) be understood from the above discussion. The most highly reduced, specialized types require further comment. As noted above, all inflorescences of A. clarkei consist o f solitary cincinni (Figure 9; Plate 3p; see discussion under A. clarkei, p. 133). These cincinni lack a peduncle and have a very contracted axis. Each cincinnus is subtended by and partially enclosed in a pair of generally leafy bracts, the outer, larger one the inflorescence bract, the inner the cincinnus bract. The some- what spathe-like inflorescence bract is analogous to but not homologous with the spathe of Commelina, which is a cincinnus bract. Inflorescences of the type of A. clarkei also occur in A. succulentum and A. lamuense (Plate 4e). The perforating inflorescence-shoots of A. zebrinum are similar to those of A. clarkei except that the inflorescence and cincinnus bracts are very reduced and the cincinnus is more elongate. Further variation in A. rebrinum is discussed under that species. ROWERS IN GENERAL.-The flower in Aneilema shows greater interspecific variation than any other organ. This variation is both biologically important, reflecting differences / i.b. 1 c.b. i .b . c.b. f.1. f.l. T f . 1 . FIGURE 9.-Diagram of the distal end of a flowering shoot of Aneilema clarkei Rendle showing the position and structure of the inflorescences: I1 is the terminal inflorescence of the shoot; 12,13, and I4 are teminal on short axillary shoots (c = cincinnus. c.b. = cincinnus bract, f.1. = foliage leaf, i.b. = inflorescence bract). 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY in breeding systems, including pollination mechanisms, and taxonomically useful, providing approximately as many char- acters as all the other organs combined. Floral characters also include the majority of attributes found useful for phylogenetic deductions. In previous studies of Aneilema, e.g., Clarke (1881a, 1901), Morton (1966), floral characters have not been given the emphasis they deserve, partly because they are difficult to interpret or often lacking in dried specimens and partly because their great variability within the genus as a whole has not been recognized. Flowers of 34 species of Aneilema are illustrated in Plates 1-4. Aneilema flowers are borne in two ranks along the cincinni. They are pedicellate, pentacyclic, trimerous, and zygomorphic. Their basic structure consists of three free, sepaloid sepals, three free, petaloid petals, three staminodes (one antesepalous, two antepetalous) borne on one side of the flower, three stamens (two antesepalous, one antepetalous) borne on the other side, and a fundamentally trilocular ovary, in which one locule is frequently abortive (Figure 10). The flowers are gcnerally unscented and remain open for a few daylight hours at the end of which they fade. Fading includes deliquescence of the corolla. Perfect, pistillate, and staminate flowers occur, the type or types present usually being species specific. Flowers vary in size from four to 40 mm wide. The zygomorphic symmetry of Aneilema flowers is due primarily to the one-sidcd arrangement of the stamens and staminodes. More striking in most species is the differentiation of the corolla into two larger, often more brightly colored, clawed petals and a smaller, duller, sessile one. A less obvious zygomorphy is also present in the calyx: the medial sepal is always morphologically distinct from the lateral sepals. The , , Of symmetry Of the flower bisects One member FIGURE 10.-Floral diagram of Aneilema. (The solid circle represents the of each whorl: the medial sepal, medial staminode, dorsal locule of the ovary, medial stamen, and medial petal (Figure cinchnus =is, the dotted fine the of Symmetry.) 10). The flowers of most Aneilema species are regularly held horizontally, i.e., like a pea flower. In these species one can unambiguously refcr to upper and lower sepals and petals. In A. neocaledonicum and A. biflorum, however, the flowers are primarily vertical, and the calyx and corolla members are best tcrmed medial (or outer) and latcral (or paircd or inner). For consistency, the tcrminology required for the vertical flowers has been applied to all spccies. The horizontal flowers characteristically have the medial sepal, paired petals, and staminodcs on the upper side and the paired sepals, medial petal, and stamens on the lower side. The tcrms outer and inner can be used for the sepals without confusion because, in bud, one sepal (the medial) always overlaps, i.e., is outside of, the other two sepals. Similarly, the medial petal in bud consistently overlaps the other two petals. The tcrms adaxial, abaxial, dorsal, vcntral, anterior, and posterior, which are all defined with reference to an axis, have not been used for the scpals and pctals, because they are ambiguous when applied to a flower that is morphologically terminal, and because they can readily be avoided. The primary orientation of the flowers appears to be to gravity, not to other inflorescence structures. It has been observed in cultivation that when flowering shoots fall over, the flowers that are produced subsequently are still horizontal despite the change in position of the inflorescence. A reorientation of the buds occurs. This has been noted to happen in as few as 12 hours in A. brenanianum, but no experiments have been conducted. Aneilema species may have only perfect, perfect, and staminate, or pistillate, perfect, and staminate flowers. When flowers of more than one type are present, they occur in the same inflorescences, and the plants are andromonoecious or polygamomonoecious. In andromonoecious and polyga- momonoecious species the different types of flowers are produced in a characteristic sequence in new inflorescences: perfect then staminate flowers in the former; pistillate, then perfect, then staminate flowers in the latter (Figures 11-14). These sequences occur during the first few days of flowering in NUMBER 76 a 15 FIGURE 11.-Idealized flowering sequence in an inflorescence of an I andromonoecious Aneilema species: diagram of the inflorescence showing the distribution of perfect and staminate flowers over the first nine days of flowering. The numbers are the days in the flowering sequence on which particular flowers opened. FIGURE 13.-Idealized flowering sequence in an inflorescence of a polyga- momonoecious Aneilema species: diagram of the inflorescence showing the distribution of pistillate, perfect, and staminate flowers over the first 10 days of flowering. The numbers are the days in the flowering sequence on which particular flowers opened. EZl Perfect flowers Staminate flowers f 5 =Pistillate flowers Perfect flowers I s t a m i n a t e flowers Day in Flowering Sequence FIGURE 12.-Idealized flowering sequence in an inflorescence of an andromonoecious Aneilema species: graph of the numbers of perfect and staminate flowers produced on each day of the flowering sequence. The numbers of flowers are based on Figure 11. 1 2 3 4 5 6 7 8 9 10 Day in Flowering Sequence FIGURE lrl.-Idealized flowering sequence in an inflorescence of a polyga- momonoecious Aneilema species: numbers of pistillate, perfect, and staminate flowers produced on each day of the flowering sequence. Ihe numbers of flowers are based on Figure 13. 16 SMITHSONIAN CONTRIBUTIONS TO BOTANY an inflorescence. Flowers somewhat intermediate between two successive types, particularly pistillate and perfect, are some- times produced. In older inflorescences the sequence is unpredictable, except that pistillate flowers almost never occur. Few Aneilema species produce either solely perfect or all three types of flowers. Only perfect flowers are normally present in A. arenicola, A. bijlorum, A. neocaledonicum, and A. zebrinum and in some populations of A. beniniense. Pistillate, perfect, and staminate flowers are regularly produced in populations of A. hockii, A. somaliense, and A. welwitschii. They have also been observed in populations of A. leiocaule and A. macrorrhizum, but the consistency of their occurrence in the last two species has not been established. The majority of Aneilema species are andromonoecious. Staminate flowers in addition to perfect ones were first recorded in Aneilema by Kunth (1843) in A. dregeanum and have since been mentioned by Bruckner (1930:163), Vogel (1955), Druten (1959), Brenan (1961). and Morton (1966). No author has indicated how widespread the Occurrence of staminate flowers is in the genus. Bruckner (1930:163) mentioned Aneilema merely as an example of a genus of Commelinaceae with staminate flowers; the other authors reported such flowers only in the context of their being present in certain Aneilema taxa. Perfect and staminate (but not pistillate) flowers have been noted by the writer in the following Aneilema species: acuminatum, aequinoctiale, beniniense (some populations), brenanianum, brunneospermum, calceolus, chrysopogon, clarkei, dispermum, dregeanum, ephemerum, forskalii, gillettii, hirtum, hockii (some populations), indehiscens (all subspecies), johnstonii, lamuense, lanceolatum subsp. lanceolatum, Iongi- capsa, longirrhizum, minutijlorum, nicholsonii, nyasense var. brevisepalum, obbiadense, pedunculosum, petersii (both sub- species), pomeridianum, pusillum (all subspecies but gypsophi- lum), recurvatum, rendlei, schlechteri, sebitense, spekei, succulentum, tanaense, taylorii, termitarium, umbrosum, usambarense, woodii. Bccausc first flowers of inflorescences have not becn sccn in every case, a few of the above might ultimatcly prove to be polygamomonoecious rather than andromonoecious. Morton (1966) has rcportcd perfect and staminate flowers in the additional taxa, A. lanceolatum subsp. subnudum and A. setiferum. The distinction bctwecn andromonoecious and polyga- momonoccious species in Aneilema is not always clean-cut. This is due to the first flowers in inflorcsccnces of normally andromonoccious populations often having reduced anthers. It appears that ccrtain environmental factors can cause the complctc supprcssion of pollcn production in these anthers. Undcr such circumstances the flowers are pistillate and the plant polygamomonoccious. Great variation in the proportion of pistillate flowers has bccn notcd in a population of A. hockii in Kenya (Faden & Faden 74/227), possibly in response to water availability. and 82% of the flowers were pistillate. During a normal rainy season no bud abortion was noted, and only 5% of the flowers were pistillate. Prolonged observations of A. hockii plants in cultivation suggest that the ratio of flowers of different types is also mediated by temperature extremes, but no experimental studies have been conducted. The morphological differences between pistillate and perfect flowers on the one hand and between staminate and perfect flowers on the other often extend beyond the mere suppression of the stamens or the gynoecium. In A. somaliense and A. welwitschii, the pistillate flowers are smaller than the perfect flowers that follow them (Plate 2f). All three anthers are reduced and lack pollen. The filaments of the lateral stamens are very short. The pistillate flowers of A. hockii (Plate 11) also show a reduction in flower size and lateral stamen length. However, the medial stamen is fully developed and its anther pollenifemus. These flowers are functionally pistillate because the pollen produced by the medial anther does not stain with cotton blue and is apparently sterile (Figure 15; Table 1, p. 32). These pistillate flowers of A. hockii further differ from the perfect ones in having shorter styles, larger stigmas, and, in at least one population, more ovules. Bruckner (1926:32) rcported dimorphism in the staminodes of A. beniniense, but no such differences have been found in that or any other Aneilema species in the present study. Bruckner?s cultivated plants should therefore be considered exceptional. In many andromonoecious or polygamomonoecious species, the lateral stamens are dimorphic in the staminate and perfect flowers, Stamen dimorphism has been observed by the writer in the following Aneilema species: aequinoctiale, beniniense FIGURE 15.-Pollen grains of Aneiletnu hockii De Wildeman stained and mounted in cotton blue: a, medial anther pollen (x-300); b, lateral antherpollen Undcr conditionsbf watcr strcss thcre was much bud abortion, (x-300). Both from Faden et al. 72/234: NUMBER 76 17 (some populations), brenanianum, gillettii, hockii, lamuense, longicapsa, rendlei, succulentum, taylorii (some populations), usambarense. It should be expected to be found in other taxa, because no monoecious species has yet been proven not to have dimorphic stamens. However, detailed stamen measurements have not been made for any species that lacked conspicuous stamen dimorphism. Dimorphic lateral stamens in staminate and perfect flowers always differ in filament length, the filaments being longer in the staminate than perfect flowers (Figure 16; Plates lg, 2b,d). Differences only in length have been noted in A. aequinoctiale, A. gillettii, A. hockii, and A. longicapsa. Further differences may occur in filament curvature, divergence from the floral midplane, and in anther orientation. Filament divergence differences are usually manifested in the stamens of the perfect flowers being more divergent than those of the staminate flowers. This has been noted in A. beniniense, A. brenanianum, A. rendlei, and A. taylorii. Conversely, in A. lamuense and A. succulentum, the stamens of the staminate flowers are more divergent (Figure 28c,d). Dimorphism in stamen curvature occurs in A. brenanianum and A. rendlei and that in anther orientation only in the latter species (Plate 2b,d). The stamens of perfect and staminate flowers in A. rendlei, the most dimorphic stamens yet observed in any Aneilema species, are illustrated in Figure 17. The extent of the stamen dimorphism in A. usambarense has not been determined because no living or preserved flowers have been seen. Another type of floral dimorphism reported in Aneilema is heterostyly (Vogel, 1955, 1978). Vogel's record of long- and short-styled flowers in A. aequinoctiale was undoubtedly based on a misinterpretation of flowers that were morphologically intermediate between perfect and staminate flowers. Such flowers have been observed occasionally in this species by the writer. They have well-developed styles that are shorter than those of the perfect flowers. They are functionally staminate, however, because their ovaries, although large, are not fully developed. This is not a case of true heterostyly, which is unknown in Aneilema. The suggestion by Omduff (1974) that Vogel (1955) might have been referring to an androdioecious condition rather than to true heterostyly is also inaccurate; androdioecism does not occur in Aneilema. As noted above, A. aequinoctiale is andromonoccious. The styles of many species may curve out of the floral midplane, either to the right or to the left (Plates 2n,p, 3a,b,d,g,k,l,n, 4d, and see below). This does not constitute true cnantiostyly because there are no differences in stamen curvature in right-styled or Icft-styled flowers. A distinct but weak fragrance has been noted in the flowers of A. brenanianum, A. brunneospermum, A. hockii, A. johnstonii (some populations), A. longirrhizum, A. nicholsonii, A. rendlei, and A. somaliense. Among these species, the strongest scents are in A. hockii, A. longirrhizum, A. rendlei, and A. somaliense. A possible scent was recorded in one population of A. dregeanum subsp. dregeanum. The fragrances 20 51 12 13 14 perfect a flowers staminate 0 flowers 1 I 15 16 17 18 Lateral stamen filament length (mm) 10 I- 5 6 7 8 9 10 11 12 13 b Lateral stamen filament length (mm) FIGURE 16.--lateral stamen filament lengths in perfect and staminate. flowers of two Aneilema species: u, A. aequinoctiale (F'. de Beauvois) G. Don. collective data from several days (from Faden ef al. 711922); b, A. rendlei C.B. Clarke, data from one day (from Kabuye & Evans 163). The data are from single plants in cultivation. have not been characterized chemically. No odors could be detected in approximately 25 species. The corolla and calyx in Aneilema buds have an imbricate aestivation. As a result, the sepals and petals can be numbered following Eichler (1875): 1, outer, overlapping the other two; 2, middle, overlapping one, overlapped by the other; 3, inner, overlapped by the other two. Sepal 1 and petal 1 are the medial sepal and petal from the standpoint of floral symmetry. Two types of aestivation of the inner petals have been found in Aneilema buds (Figure 18). In Type I (Figure 18a), petal 2 overlaps petal 3 on the side of the flower towards the outer sepal. Petal 3 overlaps petal 2 on the opposite side of the flower. In the Type I1 arrangement (Figure 18b), petal 2 completely envelops petal 3. Both types of arrangement have been found in approximately equal numbers in inflorescences of A. hockii and A. taylorii. Only Type I was found in 10 buds 18 SMITHSOMAN CONTRIBUTIONS TO BOTANY I FIoIJRE 17.-Perfect and staminate flowers in Aneilema rendlei C.B. Clarke: a, perfect flower, side view; b, staminate flower, side view (from Kabuye & Evans 163). (Bar = 1 mm.) 1 1 a 0 b 0 FIGURE 18.-Lateral petal aestivation types in Aneilema: a, Type I: petal 2 and petal 3 mutually overlapping; b, Type 11: petal 2 enveloping petal 3 (the solid circles represent the cincinnus axis). Both from A. hockii De Wildeman (Faden & Evans 711502). NUMBER 76 1 19 1 1 a b 0 0 C 0 FIGURE 19.--lateral petal aestivation development in Aneilema: a, petal 2 overlapping petal 3 dorsally, no overlap ventrally; b, petals 2 and 3 meeting in a valvate fashion ventrally; c, petal 3 overlapping petal 2, resulting in a Type I arrangement. The solid circles represent the cincinnus axis. All from A. hockii De Wildeman (Faden &Evans 711502). of A. umbrosum subsp. umbrosum. The taxonomic significance of these differences, if any, is unknown. The distribution of the two petal aestivation types within the inflorescence has been investigated in A. hockii. An inflores- cence of 12 cincinni was found to have a Type I arrangement in the largest buds of six cincinni and a Type I1 arrangement in the equivalent buds of the other six cincinni. In five of the cincinni, it was possible to check two successive buds for petal arrangement. In three of these cincinni, successive buds had the same type of petal aestivation; in the other two, sequential buds had different petal aestivation types. No pattern to the distribution of the petal aestivation types could be found within the inflorescence. The characters considered were bud size, position on the cincinnus, and cincinnus position in the inflorescence. The random distribution of the two petal arrangements in buds of A . hockii could be understood when buds of different sizes were examined (Figure 19). It was found that the overlap of petal 3 by petal 2 on the side towards sepal 1 was completely predictable and develops long before any overlap occurs on the opposite side of the flower (Figure 19u). At a later stage the as yet non-overlapping petal margins meet lengthwise in a valvate fashion (Figure 19b). At this point it appears to be a matter of chance as to which petal grows over the other. The same pattern of development may be expected in A. tuylorii, which is more difficult to work with than A. hockii, not only because of its smaller bud size, but also because the inner petals tend to spring apart when the outer one is rolled back. Furthermore, it appears that in many of the largest buds, the inner petals do not overlap at all on the lower ventral side (the reason that they spring apart?). In A. umbrosum subsp. umbrosum, it may be predicted that the lower edge of petal 3 turns inwards before this petal meets petal 2. Therefore, the edge of petal 2 always passes over that of petal 3 and a Qpe I1 arrangement regularly occurs. A further variation is that petal 2 may be either on the right or left side, viewing the bud from the front. The position of petal 2 depends upon in which of the two floral ranks of the cincinnus the bud is located. When the bud is in the right rank (viewing the cincinnus from the apex) petal 2 is on the right and petal 3 on the left. A bud in the left rank of the cincinnus has its petal 2 on the left and petal 3 on the right. Thus petal 2 is always oriented away from the cincinnus axis and petal 3 towards it. Conversely, sepal 2 is always inclined towards the cincinnus axis and sepal 3 away from it (Figure 20). Because buds are produced alternately in the two ranks of the cincinnus, successive flowers always have mirror image symmetry with regard to the position of petals 2 and 3 and sepals 2 and 3. The first flower of the cincinnus may be produced in either rank. Although the cincinni have a basically alternate arrange- ment on the inflorescence axis, no regular alternation of the rank of the first flower in successive cincinni has been found. Thus a cincinnus following one that had had its first flower in the left rank is just as likely to begin flowering in the same rank as in the opposite one. No wholly satisfactory mcthod of measuring flower size has been found. Flower width, the distance between the outer margins of the paired peul limbs, has been used here because of its ease of measurement. However, this distance is a function not only of the size of the paired petals but also of the angle at which they are held. Flowers in which the paired petals overlap each other will measure smaller than flowers that have non-overlapping petals of the same size. Furthermore, the angle at which the petals are held can vary depending upon whether 20 1 SMITHSONIAN CONTRIBUTIONS TO BOTANY 1 b 0 a 0 FIGURE 20.-Mirrorimage symmetry in Aneilema buds: a, bud from the right floral rank on the cincinnus; b, bud from the left floral rank on the cincinnus. The cincinnus is viewed from the apex. The solid dots represent the cincinnus axis. Both buds have a Type I aestivation. Both from A. hockii De Wildeman (Fuden & Evans 71/502). the flower is just opening, fully open, or beginning to fade. For standardization, measurements of flower width have been made from fully open flowers, whenever possible. Within a species, flower size and hence flower width can be a function of the stage in the life cycle of the plant when measurements are made. The first flowers of inflorescences are usually smaller than later ones, and the last flowers produced by a plant, particularly an annual, are often diminutive. In the species descriptions the total range of flower widths measured in the field and in cultivation have been included. PEDICELS.-Pedicels in Aneilema species vary in length, thickness, color, pubescence, curvature, and persistence. All except thickness and to a large extent color are useful taxonomically. Pedicel length usually increases measurably in fruit. The length by which the longest fruiting pedicel exceeds the longest flowering pedicel has been indicated in the species descriptions. Where no such difference has been noted, e.g., in A. tanaense, it is assumed that additional material would include longer fruiting pedicels than have yet been found. Flowering pedicels are generally aligned with the axes of the cincinni, i.e., horizontal to ascending. In the event of pollination and fruit set-and often even if they do not occur-the pedicel orientation changes, in order that there may be room for elongation of the cincinnus for the production of successive flowers. The pedicel may curve upward and backward (recurved), downward (decurved) or laterally. Curva- ture may occur along the whole length of the pedicel or chiefly at the base and/or apex. The various types of fruiting pedicel curvature in Aneilema are shown in Figure 21. The overwhelming majority of Aneilema species have recurvcd fruiting pcdicels. Most commonly the capsules are held erect or further inclined towards the inflorescence axis. Further recurvature occurs in most species of section Lampro- dithyros and in a few taxa belonging to other sections. The method used to designate the angle of recurvature is shown in Figure 22. Other fruiting pedicel curvatures are uncommon in the genus. Decurved pedicels, which always result in the capsules pointing downward, occur only in A. bijlorum, A. neocaledoni- cum, A . umbrosum (both subspecies), and A. taylorii (fre- quently). Laterally spreading pedicels have been observed in A. acuminatum, A . nyasense, and A. taylorii. The pedicels of flowers that do not set fruit abscise at their bases within a few days after flowering. However, fruiting pedicels in almost all Aneilema species persist even after the capsules have dehisced and the seeds have been shed. The only definite exception is A. umbrosum (both subspecies), in which the pedicels often abscise at the base about the time of capsule dehiscence. The fruiting pedicels of A. bijlorum are also easily detached at the base when the seeds are mature: some of the pedicels and their attached capsules may therefore be naturally deciduous like those of A. umbrosum. CALYX.-The calyx in Aneilema consists of three free, persistent, herbaceous, convexo-concave, sepaloid sepals. They are almost always distinctly glandular near the cucullate apex. The medial sepal always has bilateral symmetry; the lateral sepals are individually asymmetric, but they have mirror image symmetry. In addition to the difference in symmetry, the medial sepal commonly differs from the lateral sepals in being more deeply convexo-concave with a more strongly hooded apex, more prominently glandular and narrower. Differences in venation and distribution of pubescence are also frequent. All three sepals usually become appressed to the young capsule; they generally do not become reflexed. The sepal characters NUMBER 76 21 a C d C . S . P. b. c . a f i e s 1 i FIGURE 21.-Fruiting pedicel curvature types in Aneilema: 04, g, recurved;f, erect; h, spreading laterally; i j , decurved. The uncinnus apex is to the right. Examples of each type. are u, A. petersii (Hasskarl) C.B. Clarke; b, A. longirrhizwn Faden; c, A. welwitschii C.B. Clarke; d, A. clarkeiRendle; e, A. tanoense Faden$ A. rendlei C.B. Clarke; g, A. succulenturn Faden; h, A. nyasense C.B. Clarke; i, A. wnbroswn (Vahl) Kunth; j, A. neocoledonicwn Schlechter. (Not to scale; b = bracteole, c = capsule, c.a. = cincinnus axis, p = pedicel, s = sepal.) 22 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 22.-Method of designating the angle of curvature in recurved Aneilema fruiting pedicels. The flowering pedicel is considered horizontal (except in A. biflorum R. Brown and A. neocaledonicwn Schlechter, in which it is vertical), so that in u, an erect fruiting pedicel is recurved 90"; in b, a downward pointing pedicel is recurved 270". that are of taxonomic use are size, shape, color, prominence and distribution of glands, pubescence, and probably venation. The convexity of the sepals makes their actual or curvilinear dimensions very difficult to measure. As a result linear dimensions have been used instead. The sepals have been measured as if they were planar figures. The sepals are always partly or wholly green. Commonly the margins are transparent and may be tinged with various shades of purple or violet. In several species, e.g., A. hockii, A. rendiei, and A. somaliense (Plates lh,o, 2d,f,g), the colored portions are more extensive and may form distinctive patterns with the green parts, e.g., alternating lengthwise stripes. In these species the sepals appear to constitute a significant part of the visual floral attraction to pollinators. The presence or absence of non-green areas on the sepals is sufficiently constant in most species to be useful taxonomically. The major function of the sepals is protection of the inner floral whorls in bud. Protection against desiccation is provided by the secretion of the subapical sepaline glands. These vary from inconspicuous and unlobed to very prominent and distinctly bilobed (e.g., Plates 3e,f,k,l, 4j,k,n-p); rarely they appear to be completely lacking in some or all of the sepals, e.g., in A. succulentum and A. zebrinum. In many species the gland of the medial sepal is bilobed while that of the lateral sepals is unlobed. In A. benadirense and A. tunaenre, small, marginal glands are present on the medial sepal in addition to the larger, subterminal gland. Sepal pubescence probably also serves a protective function, Glandular microhairs, which are present on the adaxial surface of the sepals of all 27 Aneilema taxa in which this character has been checked, together with the sepaline glands, keep the inner floral parts and young fruit from desiccating. Other hair types present abaxially are two-celled hook-hairs, prickle hairs, and uniseriate, acicular hairs (Figure 40). Their distribution in 19 Aneilema species representing all seven sections of the genus is shown in Table 3 (p. 47). Such hairs may discourage feeding by herbivorous insects. The sepals have three to about seven parallel veins, the larger of which converge at the subapical glands. None reaches the margin. A small number of cross veins are usually present. The venation pattern of the sepals may prove useful taxonomically, but no detailed investigations have been conducted. COROLLA.-The corolla in Aneilemu is composed of three free, deliquescent, petaloid petals: two equal, clawed, lateral petals, usually oriented upwards, and a morphologically distinct medial petal generally inclined downwards. The medial petal generally differs from the lateral petals in size, shape, color and lack of a claw. In sections Aneilema and Lampro- dithyros, it is subequal to and usually concolorous with the paired petals (Plates la-c, 2m-p, 3lth,j-l,n-p, 4c-h). In other sections it is always greatly reduced and generally dull colored relative to the paired petals. However, in A. gillettii (Plate l e , f ) the three petals are concolorous; in A. hirtum and A. chrysopogon (Plate 4n,o), the medial petal, although reduced and largely dull, has a conspicuous, subapical maroon spot; in A. minutiflorum all three petals are very reduced and thus secondarily subequal. The degree of differentiation between the medial and lateral petals as well as details of size, shape, and color of both are taxonomically useful. The paired petals are usually planar, although the limbs may be reflexed at anthesis. In a few species, e.g., A. johnstonii and A. longirrhizum (Plate ld,k), the apex of the limb is slightly hooded. The limb varies from elliptic or suborbicular to subdeltate, but most commonly it is ovate or broadly ovate. The limb apex is normally rounded to obtuse, but it may frequently or occasionally be acute or truncate to slightly emarginate in some species. The limb base varies from more or less symmetric to strongly asymmetric and from cuneate to cordate. In the asymmetric typcs, the medial side (side towards the floral midplane) is more narrowly cuneate than the lateral side, or NUMBER 76 23 cuneate versus truncate, or truncate versus cordate to subcor- date. The limb margins are entire to crenulate. The limbs range in color from white to lilac, lavender, or rarely blue (but never the clear, sky blue of Commelina species), or very rarely red or from yellow to orange. The white to blue color range is the more common, occurring in all sections of the genus. A single population of A. pusillum subsp. pusillum (Plate 2i , j ) has been noted with red flowers. Yellow to orange petals are infrequent in Aneilema, as they are in all other gcnera of Commelinaceae in which they occur: Co- leotrype, Commelina, Floscopa, and Murdannia. In Aneilema they are present only in some species of sections Amelina (aequinoctiale, ephemerum, johnstonii, nyasense) and Bre- vibarbata (angolense, homblei, macrorrhizum, welwitschii) (Plates ld,i,j, 4j). Gcncrally the entire limb is uniform in color. In A. rendlei, however, the veins are contrastingly dark compared to the background color. In some populations of A. hockii, the base of the limb is white, contrasting with the rest which is lavender. Petal lustcr is causcd by reflectance from the facets of pyramidal epidcrmal cells on the adaxial surface of the limbs. The claws are cithcr concolorous wilh the limbs or paler than thcm. Like the limbs, they are usually glabrous, but in A. aequinoctiale (Plate l i ) and A. ephemerum, they are incon- spicuously beardcd with a mixture of glandular hairs and hook-hairs (Figure 40; Table 4, p. 48). Elsewhere in the Commclinaceae, pctaline hairs have been reported only in Cochliostema and Geogenanthus, in both of which they are marginal. They have also been observed by the writer on the petal bases of Murdannia simplex in Ghana, Kenya, and Zambia. The medial petal may be planar, but it is generally slightly to strongly convcxo-concave. In section Lamprodithyros, it is large and cup-, boat- or slipper-shaped (Figures 23, 24; Plates 2m9, 3a-h,j-1,n-p, 4c-h). In most species of this section, it rctains the latcral stamens for some time after the flower opcns (Plate 3c,d,k,l). The stamcns are hcld by the involute margins of this pctal and are rclcascd only whcn thcse margins uncurl, not through the action of pollinators. The stamcns may be caught for a pcriod of only a few minutcs or up to nearly thrce hours, the mcan lcngth of time for which thcy are hcld bcing spccics spccific (Figure 25). The stamcn-rctcntion mechanism is part of a pollinalion system, which is discusscd in Fadcn (1983b) and bclow. The reduced mcdial petals of the other scctions are often grecn-tinged, a color not found in the paired petals. The mcdial pctal is glabrous in all species examined except A. ephemerum in which hairs, identical to those of the paired pctal claws, are present at the base. All thrce petals have three traces at the base. The medial trace usually remains unbranched for most of its length, ultimately dividing a fcw times ncar the pctal apex. In some flowers of A. pelersii, it may branch much more. The lateral traces rcmain more or lcss parallcl to the mcdial trace for most of thcir lcngth, giving off numcrous branchcs unilatcrally. Thcse side vcinlcts fork repeatedly as they approach the margin (see Brenan, 1952:191, fig. 5). Anastomoses occur occasionally. None of the veinlets reaches the margin. This venation pattern shows little variation between species and is therefore not useful taxonomi- cally. Paired petal dimensions have been measured curvilinearly, except that the length of the hooded apex, in those species having one, has been excluded. Like the sepals, the medial petal has been measured linearly when it is not planar and cannot be flattened. An additional measurement, depth, has been used for the species in section Lamprodithyros. The width of the medial petals of these species is measured after the margins have fully unrolled. Medial petal shape, the linear outline when viewed from the front, is also recorded when the margins are fully extended in section Lamprodithyros species. ANDROEC1UM.-The androecium consists of three stami- nodes opposite the lateral petals and medial sepal (the antesepalous one occasionally lacking) and three stamens opposite the lateral sepals and medial petal. Thus the outer, antesepalous androecial whorl contains two stamens and a staminode, and the inner, antepetalous whorl two staminodes and a stamen (Figure 10). The usually horizontal orientation of the flowers results in the shorter staminodes being held above the longer stamens. The staminode and medial stamen filamants are always glabrous: the lateral stamen filaments are glabrous or variously bearded, according to the species. All of the filaments are usually free, except in section Lamprodithy- ros, in which those of the three stamens are always fused basally and frequently also shortly adnate to the filaments of the lateral staminodes (Figures 26-28: Plate 46). A further short adnation to the medial petal base is also general in these species. In the remainder of the genus, fused stamen filaments have been observed only in A. hockii (section Amelina). Staminodes: The medial staminode is usually differenti- ated from the lateral staminodes in size and form. Its filament is generally shorter and its antherode either larger or smaller and frequently with differently shaped lobes. The medial staminode is complctcly lacking or vestigial in the following Aneilema species: benadirense, beniniense (some populations), calceolus, clarkei (occasional flowers in some populations), minutiflorum, setiferum (sometimes lacking, fide Morton, 1966), tanaense, termitarium, and umbrosum (some popula- tions). All three staminodes are equal or subequal in species of section Aneilema and in A. johnstonii (section Amelina) (Plate la,c,d). Although sometimes distinctly thickened basally, the stami- node filaments are always more slender than those of the stamens. Except in section Rendlei and in some populations of A. hockii, the staminode filaments are little curved, being straight or gently arcuate for all or most of their length (e.g., Plates lc,z',j, 3c,d,o,p, 4b,c,f) . The lateral staminode filaments are sometimes sharply dcflcxcd near the apex. The strongly curved latcral staminode filaments of A. rendlei, A. brenania- 24 SMITHSONIAN CONTRIBUTIONS TO BOTANY a d I 0 m $: b 6 e :i h k n 4. C f I I 0 FIGURE 23.-Medial petals of five species of Aneilema section Lamprodithyros: a-c, A. petersii (Hasskarl) C.B. Clarke subsp. petersii (from Andrews s.n.); d$ A. indehiscens Faden subsp. indehiscens (from Faden & Faden 72/72); g i , A . recurvatum Faden (from Faden 6911300); j-l, A. tanaenre Faden (from Gillett 19973); m-o, A. calceolrcs Brenan (from Organ in EA14970). All are after stamens have been released (where appropriate). (a,d,gj,m, front views; b,e,h,k,n, lateral views; c, $i,l,o, apical views). (Bar = 1 mm.) NUMBER 76 25 a d Q i 3 b e -4 h b k C f i U I FIGURE 24.-Medial petals of four spedes of Aneilema secticm Lomprcdithyros: a-c, A . clarkei Rendle (from Evans & Maikweki 51); d$ A. lamuense Faden (from Faden & Faden 7411083); g-i, A . succdentum Faden (from Faden ef al. 701937); j l , A. zebrinum Chiovenda (from Faden et al. 74/330). All are after stamens have been released (where. appropriate). (a ,d ,gj , front views; b,e,h,k, lateral views; c, fii,l, apical views.) (Bar = 1 mm.) 26 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 25-Flowering times and lateral stamen retention in five species of Anei l em section Lamprodiihyras in Kenya. The bars represent the interval between the opening of the first flower and onset of fading. num, and A. taylorii are shown in Figures 17,47c,e and Plates ln,p, 2 b 4 , respectively. The sterile anthers or antherodes of the staminodes are typically bilobed (Figure 43). In A. neocaledonicum (Plate lc) , however, they are primitively scarcely lobed, while occasion- ally in other species they may be unlobed or hardly lobed through reduction. The lobes show much variation in shape according to the species. They may be sessile or stalked, the length of the stalk and angle of its attachment to the filament being characteristic in some m a . The antherodes of all staminodes are generally yellow except in A. acuminatum (Plate l a b ) , in which they may be white or violet; in A. rendlei and A. taylorii (Plate lm,o), in which the lobes of at least the medial staminode are maroon-spotted adaxially; and in A. lamuense (Plate 4b,c), in which the lobes have pinkish purple bases. The staminode lobes vary further in being either thin and flat or thick and tumid. Their surface is generally more or less smooth, but in A. welwitschii, it is verrucose. The connective is sometimes slightly elongate between the staminode lobes, appearing as an extension of the filament to which the lobes are seemingly subterminally attached. All characters of the staminodes are useful taxonomically. The most important are relative degree of development of medial and lateral staminodes, particularly of their antherodes, presence or absence of a strong curvature in the filaments, and shape and size of the antherode lobes. Staminode characters must be used with caution because, although relatively constant in some taxa, e.g., A. rendlei and A. petersii, they are variable in others, e.g., A. hockii and A. indehiscens subsp. lilacinum. Although the staminodes completely lack pollen, their function appears to be the attraction of pollinators. The usual yellow color of the antherodes may make them appear polleniferous to some insects. The staminodes may further attract pollinators by producing a scent in some taxa. Several flowers of A. somaliense were dissected into separate floral organs: sepals, petals, staminodes, stamens, and gynoecium. The staminodes appeared to be one source of the scent produced by these flowers, but the results were inconclusive. The possibility that the antherode lobes in Aneilema may be sources of liquid for pollinators, as has been suggested for Commelina staminodes by Faegri and Pijl (1971:71), has not been ruled out, but field observations of numerous flower visitors have thus far failed to support this hypothesis. Stamens: In all Aneilema species-except A. neocaledoni- cum (Plate lc), in which the three stamens are more or less equal-the lateral stamens always have longer filaments and further differ in filament curvature and anther form from the medial stamen. One aspect of lateral filament curvature that varies among species is the relationship of the filaments to each other, i.e., whether they are divergent, parallel, or convergent. This can be determined most satisfactorily from a top view of the flower. The filaments may be closely parallel for their entire length, as in A. aequinoctiale (Plate l i ) , but more commonly, they are divergent for at least part of their length. In some species different parts of the filament regularly have different degrees of convergence or divergence (Figures 264 2 8 ~ ) . The lateral filaments cross in A. petersii (Figure 266; Plate 2m-p), the only species in which this character is typical. Top views of NUMBER 76 21 a b C d FIGURE 26.-Top views of the androecia of four species of Aneilema section Lumprodithyros, medial staminode anitted, gynoecium removed: (I, A. indehiscens Faden subsp. indehiscens (from Faden & Faden 72172); b, A . petersii (Hasskarl) C.B. Clarke subsp. pefersii (from Andrews s.n.); c, A. forskolii Kunlh (from Gilberf & Thdin 969); d, A . recurvafum Faden (from Faden 69/1300). (Bar = 1 mm.) 28 SMITHSONIAN CONTRIBUTIONS TO BOTANY a I b C FIGURE 27.-Top views of the androecia of three species of Aneilemu section Larnprcxiithyros, medial staminode absent or omitted, gynoecium removed: a , A. fanaense Faden (from Gillelt 19973); b, A. calceolus Brenan (from Faden & Faden 7411069); c, A. zebrinwn Chiovenda (from Faden & Faden 741482). (Bar = 1 mm.) the filaments of 12 species of section Lamprodithyros are shown in Figures 26-28, SOg, 51g. The aspect of lateral stamen filament curvature that is most important taxonomically is the shape of the filament in lateral view. Four basic types are recognizable: straight, undulate, S-shaped or sigmoid, and J-shaped (Figure 29). The U-shaped curvature mentioned in section Rendlei, e.g., Plate lp , is considered a variant of the J-shape, having been derived by reduction of the longer arm. Variation in the angle at which the filaments are held relative to the horizontal can result in the final position of the anthers being quite different in relation to other floral parts, even between species with the same basic curvature type (Figures 2964,Jg). The four main curvature types have been useful in defining the sections in Aneilema. The lateral stamen filaments are straight or nearly so in section Aneilema (Plate la-c), straight or undulate in section Amelina (Plate Id-k), sigmoid in sections Somaliensia, Lamprodithyros, Brevibarbata, and Pedunculosa plates 2e9, 3, 4), and principally J- or U-shaped in section Rendlei (Plates Im-p, 2a-d). The curvature of the filaments in bud is indicative of that found in the flower (Figure 30). Thus the stamen curvature of species known only from dried NUMBER 76 29 W a b d FIGURE 28.-Top views of the androecia of three species of Aneilema section Lamprodithryos, medial staminode omitted, gyncecium, when present, removed: a, A. clarkei Rendle (from Faden & Faden 7411215); b, A . succulenturn Faden (from Faden & Faden 7411152); c,d, A. lamuense Faden: c, perfect flower. d, staminate flower (from Faden & Faden 7411083). (Bar = 1 mm.) 30 SMITHSONIAN CONTRIBUTIONS TO BOTANY a h b i A d e f J' FIGURE 29.--Lateral stamen filament curvature types in Aneilema: a , straight; b - d f , g , S-shaped or sigmoid; e, undulate; h, J-shaped. (Not to scale.) specimens can often be deduced through careful dissection of mature buds. The lateral stamen filaments are either glabrous or variously bearded. The presence or absence of hairs on the filaments is a specific character in Aneilema. It does not vary within a species, unlike the apparent situation in some Murdannia taxa (see Clarke, 1881a:207, under Aneilema spiratum = Murdannia spirata). Filament pubescence is also significant at the sectional level: sections Aneilema, Rendlei, Somaliensia, and Lamprodithyros are characterized by glabrous filaments, and sections Brevibarbata and Pedunculosa by bearded filaments. Only section Amelina includes some species with glabrous and others with bearded filaments. Considerable variation in the types of hairs and in their distribution on the lateral stamen filaments is present in the genus as a whole, but on closer FIGURE 30.-bteral stamen and lateral staminode fdament curvatures in buds of Aneilema species: a, A. neocaledonicwn Schlechter (from Bierhorst in Kew Accession No. 00069.12046); b , A . acwninatwn R. Brown (from Hind 683); c , A. aequinoctiale (P. de Beauvois) G. Don (from Faden et al. 711922); d , A . hockii De Wildeman (from Faden et al. 721234); e , A. welwitschii C.B. Clarke, bud of staminate flower (from Faden et al. 7 4 1 77); f, A. recurvaturn Faden (from Faden el al. 6911066); g, A. somalienre C.B. Clarke (from Faden & Faden 74/923); h, A. rendlei C.B. Clarke (from Kabuye & Evans 163); i, A . petersii (Hasskarl) C.B. Clarke subsp. petersii (from Faden & Faden 74/291). All buds are from cultivated plants. In flower, a-c have lateral stamen filaments straight; d , undulate; e-g,i, S-shaped or sigmoid; h, J-shaped. (Bar = 1 mm.) inspection, all of the unusual types and distributions occur solely in section Amelina and are usually restricted to single species. The only generalizations that hold for all species are that the hairs are never moniliform and that they are attached predominantly or exclusively to the abaxial surface of the filament. The hairs are generally uniseriate and eglandular, with a rounded terminal cell, but branched hairs (A. nyasense), glandular hairs (A. gillettii), and hook-hairs (A. aequinoctiale) NUMBER 76 31 also occur (Figure 41; Table 4). Typically the hairs are present only distal to the middle of the filament, but in A. nyasense and A. uequinoctiale (Plate l i , j ) they are distributed along most of its length, and in A. gillettii (Plate l f ) they are found only below the middle. The hairs vary in length from about 40 to 1000 pm. They may be very short, whitish and easily overlooked, e.g., in A. pomeridianum (Plate 4k), or long, brightly colored, and very conspicuous, as in most species of section Pedunculosa (Plate 4m-p). The lateral filaments may be terete or strongly flattened dorsiventrally and ribbon-like. In section Pedunculosa, they are broadened in the region to which the hairs are attached. The character of lateral stamen dimorphism is discussed above. The filament of the medial stamen is much less variable than those of the lateral stamens. It is always glabrous, usually straight or gently arcuate, and often slightly to strongly recurved apically. Aneilema anthers are four-celled and longitudinally dehis- cent (Figure 3 1). The lateral and medial stamen anthers vary in relative degree of development, shape, size, form of the connective, color, attachment, orientation, and pollen color. The medial anther may be larger or smaller than the lateral anthers. There is a general inverse correlation between medial anther size and the amount of lateral filament pubescence. The species with glabrous filaments always have the medial anther larger than the lateral anthers. Those with short, inconspicuous hairs on the filaments usually have the three anthers subequal in size, although the medial differs in shape. The species with dcnsely beardcd filaments generally have the medial anther distinctly smaller than the laterals. Aneilema nyasense (Plate l j ) is an exccption, for its lateral filaments are minutcly bearded, but its medial anther is reduced. Aneilemu gilletfii (Plate l e , f ) is also unusual in having finely bearded filaments and an enlarged medial anther. The lateral anthers, except in A. bijlorum, have narrow connectives (Figure 31c,d). In contrast, the medial anther has a broad connective, with the exception of A. neocaledonicum (Figure 3la,b,e,f). The medial anther connective may be shorter than the anther sacs, the whole anther thereby bowtie- or dumbbell-shapcd, e.g., A. beniniense, A. nicholsonii, and A. welwitschii, or more or less equal in length to the anthcr sacs. In the latter case the anther may be shield-shaped, e.g., A. ephemerum and A. johnstonii, saddle-shapcd, e.g., A. hockii and A. petersii (Plates l hJ , 2n,p), or convex-ellipsoid, e.g., A. rendlei and A. taylorii (Plate lm-p). The conncctives of the larger medial anthers are commonly strikingly spottcd or transverscly striped with maroon (Plate lm-p). The anthcrs of A. neocaledonicum and A. bijlorum are basifixed. More generally, however, the anthers are slightly dorsifixed. The enlarged medial anthers are usually versatile; the lateral anthers are not distinctly versatile. The thecae of all three anthers in A. neocaledonicum and A. bijlorum are situated on opposite ends of the connective (Figure 31a.b). The anther dehiscence is latrorse. In all other Aneilemu a C b d e FIGURE 31.-Aneilema anthers. A . neocaledonicum Schlechter (a ,b): a , anther. b, transverse section of a (from Bierhorst in Kew Accession No. 000-69.12046). A . hockii De Wildeman ( c d ) : c, lateral anther; d, transverse section of c (from Faden et al. 721234). A . taylorii C.B. Clarke ( e , f ) : e, medial anther;f, transverse section of e (from Faden & Faden 74/37]) The brdten lines indicate the planes of the sections. (Bars = 0.5 mm.) species the anther sacs in at least the lateral anthers are both located on one side of the connective or, in the case of the large medial anther, usually face in more or less the same direction due to the outgrowth of the connective (Figure 3 1 ~ 9 ) . The varied orientation of the anthers, erect, horizontal, or pendent, usually results in an introrse dehiscence. However, the lateral anthers of A. petersii (Plate 2n,p) and of one population of A. hockii and occasionally the medial anther of A. dregeanum subsp. dregeanum have a more or less extrorse dehiscence. 32 The medial stamen has been considered staminodial in five West African Aneilema species (A. beniniense, A. lanceolatum, A. pomeridianum, A. setiferum, and A. umbrosum) by Morton (1966). West African populations of A. beniniense, A. umbrosum, and A. lanceolatum subsp. lanceolatum have been checked and the medial anthers found to produce viable pollen (Table 1). In view of these results and of the observation that even more reduced medial anthers of other Aneilema species usually produce some pollen, Morton?s description of the medial anthers of the other two West African species as staminodial requires confirmation. The pollen grains in Aneilema vary in color from white to yellow or orange. Commonly the pollen of the medial anther differs in hue from that of the lateral anthers. This color difference is sometimes taxonomically useful. For example, in A. calceolus (Plate 3n,o) the medial anther has yellow pollen and the lateral anthers white pollen, while in the closely related A. tunaense (Plate 31), all three anthers have yellow pollen. Similarly, A. clarkei has yellow and white pollen in the medial and lateral anthers, respectively. Its close relative A. lamuense has white pollen in all anthers, the only Aneilema species noted with this character. In A. taylorii the pollen color differentiation is less conspicuous than in A. calceolus or A. clarkei: the lateral anther pollen is usually orange-yellow and the medial anther pollen yellow when the flower first opens. During the course of flowering, the pollen grains of the medial anther turn white, and so when the flower begins to fade, usually only white pollen is present in this anther. No such color change occurs in the lateral anther pollen; hence pollen color differentiation increases during the flowering period. A further difference occurs in A. hockii pollen. The medial anther pollen differs not only in color but also in stainability with cotton blue, an indicator of viability. All of its pollen grains stain weakly and show shriveled contents; they are apparently inviable. In contrast, 96%-98% of the lateral anther pollen grains stain strongly and appear viable (Figure 15; Table 1). In one population both types of pollen grains were equal in size in the three flowers checked. In a single flower from another population, the medial anther pollen grains were distinctly longer than those of the lateral anthers. Such SMITHSONAN CONTRIBUTIONS TO BOTANY differences in pollen stainability and size have previously been reported in the Commelinaceae only in Tripogandra (Lee, 1961; Handlos, 1970, 1975; Mattsson, 1976; Poole and Hunt, 1980) and Palisota (Faden, 1983c, 1988). No variation in pollen size or staining was found in the anthers of five other Aneilema species (Table 1). Aneilema pollen grains are monosulcate and ellipsoid to bean-shaped or sometimes irregularly spherical. Scanning electron micrographs (Figures 32, 33) show that the exine is spinulose or spinulose-verrucose and punctitectate. The spin- ules or warts are blunt and randomly arranged. Lengths of 32, 33, 53, and 59 pm have been reported for the pollen grains of four unnamed species by Erdtman (19%) and of 63-91 (mean 76.9) pm for A. aequinoctiale by Zinderen Bakker (1953). Poole and Hunt (1980) report mean diameters for six species ranging from 32.1 pm to 68.5 pm. The lengths of five species measured in the present study ranged from 30.5 pm to 47.6 pm (Table 2). The results of Poole and Hunt (1980) suggest that the following pollen characters may be useful taxonomically at the species level: dimensions of the pollen grain; length of the spinules and distance between them: and thickness of the wall. Populations of additional species need to be investigated before the general usefulness of pollen characters in Aneilemu can be confirmed. All stamen characters are useful taxonomically. The most important are the presence or absence of filament fusion, dimorphism, and pubescence; type of filament curvature; relative size of lateral and medial anthers; form of the medial anther; and pollen color. Staminode and stamen filament lengths are measured curvilinearly and include the length of the fused basal portion, when present. Antherode and anther measurements are linear. Antherode lobe length is measured at right angles to the staminode filament, or, when the lobes are stipitate, then parallel to the stalk. Lobe length does not include the stalk. Lobe width is measured at right angles to the length. GYN0ECIUM.-The gynoecium consists of a basically trilocular ovary and a simple, terminal style and stigma. The ovary may be sessile or stipitate, but the distinction between these conditions is seldom clean-cut. Three longitudinal lines TABLE 1.-Pollen viability in Aneilema species (the fint 100 pollen grains encountered were tallied; * = well-formed pollen grains that stained darkly with cotton blue were considered viable). Taxon Aneilema hockii A. hockii A. rendlei A. clarkei A. lanceolatwn subsp. lanceolalum A. beniniense subsp. beniniense A. umbroswn subsp. umbrmwn Lateral Medial Number anther anlher of pollen pollen Collection Ploidy buds % viable* % viable* Faden et al . 721234 2x 3 96-98 0 Faden &Evans 711502 2x 1 91 0 Kabuye & Evam 163 2x 2 99 99-100 Faden et al. 711633 6x 2 98 99 Lock 84193 2x 3 87-94 86-9 1 Faden et al. 74171 4% 2 99-100 99-100 Faden et al. 7412 4x 2 5-26 29-50 NUMBER 76 33 FIGURE 32.Scanning electron micrographs of acetolyzed Aneilema pollen grains (polar view): a, A. pefersii (Hasskarl) C.B. Clarke subsp. pallidiflorum Faden. a 9 4 5 (from Faden & Faden 74/237): b, A. somalienre C.B. Clarke, ~3400 (from Faden & Faden 74/923); c, A . beniniense (P. de Beauvois) Kunth. ~ 2 8 9 0 (from Faden er al. 74/71). are always visible, a middorsal and two lateral sutures. The latter represent lines of dehiscence in the capsule. In nearly all Aneilema species, the middorsal suture is a vestige of a former line of dehiscence. Only in some plants of A. acuminatum does dehiscence regularly occur along this suture. The ovary is generally entirely green, but in some species distinctive violet patches, spots, or stripes are present apically. The ovary may be glabrous or variously puberulous with 34 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 33.4canning electron micrographs of pollen grains of Aneilemu somulieme C.B. Clarke: (I, equatorial view, longitudinal position, ~ 3 4 0 0 , b, polar view, ~ 3 4 0 0 , c, portion of exine. ~8500. (All from Fuden & Fuden 741923.) glandular and occasionally other hair types. The glandular hairs may be appressed microhairs or longer, appressed to patent, uniseriate hairs. The latter may have a globose to ovoid and capitate or a more or less cylindrical, non-capitate terminal cell. Other hair types observed on ovaries are hook-hairs and prickle hairs (Figure 42; Table 5 , p. 49). Some sections of the genus are characterized by the distribution and types of hairs present on the ovary. For example, the species of section Rendlei have NUMBER 76 35 TABLE 2.-Pollen grain lengths in some Aneilemo species. Taxon I Aneilemo hockii A . rendlei A . somalieme A. pelersii subsp. pollidiflorum A. beniniense subsp. beniniense A. beniniense subsu. beniniense Number of Mean pollen grains length Collection Treatment measured OLm) Foden et 01. 721234 cotton blue' 15 47.6 Kobuye & Evans 163 cotton blue' 15 30.5 Foden & Faden 741923 acetolyzedZ 15 32 Foden & Faden 741237 acetolyzed2 15 38.9 Faden el 01. 74/71 cotton blue' 15 31.6 Foden el 01. 74/71 acetolyzed2 15 32.8 'Pollen grains removed from fresh mature buds and mounted in cotton blue, otherwise untreated. zPollen grains acetolyzed after Erdrman (1966) and mounted in glycerine jelly. only glandular microhairs, which are found chiefly on the lateral sutures and ventral surface. In the majority of Aneilema species, the dorsal locule is either empty or completely vestigial but identifiable in transverse section. In the rcmaining species one (rarely up to five) ovule is present in this locule. Species that regularly or occasionally have an ovule in the dorsal locule occur in all sections of the genus and are especially frequent in sections Aneilema, Somaliensia, Amelina, and Lamprodithyros. The ventral locules each contain one to six uniseriate, campylotro- pous ovules. The basal ovule in each locule characteristically has its micropylar end directed towards the dorsal surface of the ovary. Successive ovules alternately face ventrally, then dorsally, and so forth. When reduction in ovule number occurs, as in the later flowers of inflorescences in some A. hockii populations, it starts from the apical end of the ovary. The style is always slender. It is typically glabrous except for an occasional hair of the ovarian type on the base in some species. Four basic curvature types occur (in lateral view): straight, arcuate, undulate, and J-shaped (Figure 34). In the first two the style is sometimes recurved near the apex; in the latter two it is always recurved apically. Straight or arcuate styles that are not apically recurved are frequent in autogamous species. In most species the style lies in the floral midplane or nearly so. In section Lamprodithyros, the styles are usually strongly curved laterally out of the floral midplane (Plates 2ng, 3a,b,d,g,k,l,n, 4d). The distribution of left-handed and right- handed style curvatures within inflorescences has been studied in A. recurvatum and A. tanaense and found to be random. Either type is just as likely to occur in flowers of either cincinnus rank. The curvature of the stamen filaments is not affected by the style curvature. The stigma is terminal, usually capitate, and sometimes colorcd contrastingly to the style, e.g., in A. acuminatum (Plate la). The largest stigmas, e.g., in pistillate flowers of A. hockii (Plate ll), may be somewhat triangular in outline. In some species, particularly those of sections Rendlei and Somaliensia, the stigma is not at all enlarged, and is termed small in the species descriptions. All enlarged stigmas are called capitate. Owens and Kimmins (1981) surveyed stigma morphology in the Commelinaceae, including several species of Aneilema. Both dry and wet stigmas are reported in the genus, but which b C FIGURE 3 4 . 4 t y l e curvature types in Aneilemo: a, straight; b, arcuate- decurved; c, undulate; d , arcuate-decurved. recurved apically; e, J-shaped. (Not to scale.) 36 SMITHSONIAN CONTRIBUTIONS TO BOTANY species have which stigma type is not recorded. Aneilema is reported to be one of only four (out of the 17) genera investigated that lacks raised blisters on the stigma papillae. It also shares with only Commelina the character of the papillae covered by a distinctive folded cuticle. In a subsequent paper, Owens and Horsfield (1982) summarize the stigmas of Aneilema species as trifid and wet, but in the text they mention that exceptional stigmas may be circular in transverse section, and that both dry and wet stigmas were found in plants of A. aequinoctiale and A. hockii. In one accession of the former species, exclusively dry stigmas were found. No list of species was published in either Owens and Kimmins (1981) or Owens and Horsfield (1982), but Hunt (in litt., 23 February 1988) stated that the Aneilema species used were A. aequinoctiale (two accessions) and A. hockii, A. brenanianum, and an unnamed species (one accession each). Murdannia zeylanica (C.B. Clarke) Bruckner was also used as an Aneilema species. The characters of the gynoecium that are most useful taxonomically are ovarian hair types and distribution, dorsal locule development, ovule number in the ventral locules, and style curvature. Ovary shape is the linear outline viewing the ovary from either the dorsal or ventral surface. Ovary length and width are linear measurements. Length includes the stalk, when present. Style length is curvilinear. CAPSULES.-Aneilema capsules are sessile or stipitate, dry, usually dehiscent, bi- or trilocular, generally bivalved, usually lustrous, and gray to brown or stramineous. As in the case of the ovary, there is no clear distinction between sessile and stipitate capsules. The most plainly stipitate capsules are in the species group of A. grandibracteolatum, A. obbiadense, and A. longicapsa (section Somaliensia) and in several species of section Lamprodithyros, e.g., A. forskalii and A. petersii (Figure 35a,b). Aside from A. zebrinum and some populations of A. indehiscens, in which the capsules are regularly or frequently indehiscent or only partially dehiscent, all species have dehiscent capsules. In the dehiscent capsules the valves may spread only slightly (. .+'. A::. '."a . .,. 2;: ,:;.: 1 .>>.:: .. . :: . .v;: .f " ..L :9?, ...;a, . .a .:?/, r., C d e i i FIGURE 40.-Sepaline and petaline hairs in Aneilem. Sepaline hairs (a-g): (u-c, adaxial glandular micro- hairs): a, A. biflorwn R. Brown (from Tindale in NSW84495); b,c, A. beniniense (P. de Beauvois) Kunth (from Faden et al. 74171); (d,e, hook-hairs): A. hockii De Wildeman (from Fuden et al. 721234); f, acicular hair, A. hirtum A. Richard (from Paget-Wilkes B); g , prickle hair, A. zebrinwn Chiovenda (from Faden et al. 741330). Petaline hairs (k.3: h, uniseriate-glandular hair, A. aequinoctiule (P. de Beauvois) G. Don (from Faden et al. 711922); i, glandular microhair, A. ephemerwn Faden (from Faden & Faden 741794); j , hook-hair,A. ephemerwn Faden (from Fuden & Fuden 74794). (Bar = 0.1 mm for w,g- j ; 0.3 mm forf.) NUMBER 76 47 TABLE 3.--?Lpes and distribution of hairs on sepals in 19 species of Aneilema (figures refer to number of cells in the hairs; only presence (+) or absence (-) has been indicated for glandular microhairs because, by definition, they are always 3-celled). Aneilema sections and species Section Aneilema A . neocaledonicwn A . acwninatum A . biforum Section Amelina A . johnstonii A . gillettii A . hockii A . aequinoct iale A. ephemerum A. nyasense Section Rendlei A . taylorii A. rendlei Section Somaliensia A. somalieme Section Lamprodithyros A . clarkei A . recurvatum A. zebrinum A . beniniense A . brunneospermwn Section Peduncdosa A. hirtwn A . termilariwn Section Brevibarbata Abaxial Hook-hairs Adaxial Acicular glandular Glandular or prickle Uniform T~~ - - ___. . - microhairs microhairs hairs in size sizes + + + + + + + + + + + + + + + + + + + Chromosome numbers have been found to be very useful in delimiting the sections of Aneilema and in understanding some species and infraspecific m a . The basic numbers obtained for each section are included in the descriptions of the sections. Chromosome counts for species of sections Rendlei, Somalien- sia, and Lamprodithyros are listed in the accounts of these species. Because the full data will be presented elsewhere, only a brief summary will be included here. Five basic chromosome numbers were mentioned for Aneilema by Faden and Suda (1980); a sixth has been found subsequently among the more than 40 species of the genus that we have counted: x = 9, 10, and 13-16. The sections that appear to be natural on morphological grounds usually have a single basic number, except section Somaliensia: section Aneilema ( x = 16); section Rendlei (x = 16); section Somaliensia (x = 14, 16); section Lamprodithyros (x = 13); section Pedunculosa (x = 9). The remaining two sections are much more diverse in their morphology, and this is also reflected in their cytology: section Amelina (x = 13, 15, 16); section Brevibarbata (x = 10, 13, 15). The chromosomes of Aneilema are generally small and, in nearly all species, all or most chromosomes are metacentrics and submetacentrics. Phytochemistry of Aneilema Very little work has been done on the phytochemistry of Aneilema. Flavonoids are the only group that has been studied, as part of a general survey of these compounds in the Commelinaceae (Martinez and Swain, 1985). Twenty-four populations of 17 species, representing all seven sections of the genus, were examined. Flavone C-glycosides, the most common flavonoids in the family, were found in every population and were the only kinds of flavonoids present. Although the specific flavone C-glycosides found in each species are not listed, it would appear that flavone C- glycosides are of such wide distribution within the family and within Aneilema as to be worthless taxonomically. Reproductive Biology of Aneilema The only paper dealing extensively with the reproductive biology of Aneilema is that of Faden (1983b) in which the floral characters of five species from a single locality in Kenya are compared. Those data are in turn only a distillation of a great body of information that was accumulated from the field and from greenhouse observations and experiments in the course of 48 SMITHSONIAN CONTRIBUTIONS TO BOTANY TABLE 4.--Types and distribution of hairs on the petals and stamen filaments in 19 species of Aneilema (figures refer to number of cells in the hairs; only presence (+) or absence (-)has been indicated for glandular microhain because, by definition, they are always 3-celled). Aneilema sections and species Section Aneilema A . neocaledonicwn A. acuminatwn A. biflorum Section Amelina A . johnstonii A. gillettii A. hockii A . aequinoctiale A. ephemerum A. nyasense Section Rendlei A. taylorii A. rendlei Section Somalienria A . somalienre Section Lamprcdithyros A. clarkei A . recurvatwn A. zebrinwn A. beniniense A. brunneospermum Section Pedunculosa A. hirtum A. termitariwn Section Brevibarbata Petals Stamen filaments Glandular Hook- Uniseriate- Hook- Uniseriate- Uniseriate- microhairs hairs glandular hairs eglandular glandular Branched preparation of Faden (1975) and subsequently. Because a compilation of all these data is likely to be overlooked in a work that is largely taxonomic, only a summary will be included here, and the rest will be published elsewhere. Aneilema species differ from one another in a number of features of floral structure and biology that may be important for pollination and reproductive isolation. Among the biologi- cally significant morphological characters are types of flowers present, flower size, petal color, position of the stamens at anthesis, pubescence of the lateral stamen filaments, fertility of the pollen in the diffcrent anthers, and stigma position relative to that of the anthers. The variation in these characters within Aneilema is discusscd under ?Reproductive Morphology,? above. Floral reproductive attributes, other than strictly structural ones, that may also be important are presence or absence of a scent, daily flowering times, retention of the lateral stamens by the medial petal, and presence or absence of self-compatibility and autogamy. One hundred and eighty-four insects and two spiders were collected on or associated with 18 Aneilema species in six African countries (Ghana, Zambia, Malawi, South Africa, Kenya, and Tanzania) in 1974. The major insect orders collected were (1) Hymenoptera (79 individuals)- Megachilidae: two genera (at least four species); Halictidae: three genera (at least three species); Anthophoridae: seven genera (at least nine species); Apidae: two genera (at least two species); (2) Diptera (42 individualsjsciaridae: one genus (two species); Bombydiidae: two genera and species; Syrphi- dae: eight genera (at least eight species); Calliphoridae: one genus and species; (3) Coleoptera (53 individuals)- Scarabaeidae: two genera and species; Dermestidae: one genus and species; Melyridae: one genus and species; Meloidae: two genera (at least five species); Alleculidae: one genus and species; Chrysomelidae: five genera (at least nine species). Other insect orders collected were Hemiptera (four individu- als), Thysanoptera (one) and Lepidoptera (five larvae). Despite this large set of data, few generalizations can be made. To some degree, the numbers and diversity of insects collected from an Aneilema species Seem to be correlated with the number and length of the observation periods, the number of different populations of the species sampled, and the size of the population or populations. It also appears that a combina- tion of Aneilema species and, especially, the presence of a nectar source, such as species of Plectranthus (Lamiaceae), in NUMBER 76 49 TABLE 5.-Tms and distribution of hairs on the ovaxy in 19 species of Aneilema (figures refer to number of cells in the hairs; only presence (+) or absence (-)has been indicated for glandular microhairs because, by definition, they are always 3-celled). Aneilema sections and species Section Aneilema A. neocaledonicwn A. acuminatwn A . biJ?orwn Section Amelina A . johnstonii A. gillettii A. hockii A. aequinoctiale A . ephemerwn A . nyasense Section Rendlei A . iaylorii A . rendlei Section Somaliensia A . somaliense Section Lamprcdithyros A . c larb i A . recwvatwn A. rebrinwn A. beniniense A . brunneospermwn Section Pedunculosa A. hirtum A . termitarium Section Brevibarbata Uniseriate- Glandular Prickle Hook- glandular microhairs hairs hairs Capitate Non-capitate the vicinity may increase the number of insect species visiting particular Aneilema species. Among the 18 Aneilema species on which insects were caught, only A. petersii subsp. pallidijlorum and A. hockii, growing in the Tsavo area of Kenya, could be considered relatively well sampled. The largest numbers of insect species were collected on these taxa: A. petersii subsp. pallidiflorum (six Hymenoptera species, three Diptera, five Coleoptera, one Lepidopteran); A. hockii (six Hymenoptera, six Coleoptera). The single, large popula- tion of A. nicholsonii, which was visited only once, yielded two species of Hymenoptera, four Diptera, three Coleoptera, and one Thysanopteran. In contrast to these data, a month-long, intensive study of the reproductive biology of Commelina erecta L. in Texas in 1981 yielded 44 visiting insect species (Faden, unpublished). Because no Aneilema species was intensively studied for that length of time, one might expect that more insect visitors could have been obtained from further observations of even the best sampled Aneilema species. Two further generalizations may be made. Larger flowers, such as those of A. hockii, tend to attract larger bees than smaller flowers, e.g., A. succulentum. This is especially true if the stamens of the larger flower extend forwards to a considerable degree rather than bend downwards (Faden, 1983b). Finally, most Aneilema species seem to be visited (and presumably pollinated) chiefly by bees. Bees were collected on 16 Aneilema species and flies (mostly Syrphidae) on only eight species; both bees and flies were collected on seven species. It is noteworthy that equal or greater numbers of fly than bee species were obtained from only five Aneilema species. These species (aequinoctiale, welwitschii, nicholsonii, pedunculo- sum, and spekei) all have bearded filaments, and they include all three species of section Pedunculosa (nicholsonii, peduncu- losum, spekei) and the only species of section Amelina (aequinoctiale) with bearded filaments on which insects were collected. These four species all have longly bearded filaments, unlike the short-bearded A. welwitschii (section Brevibarbata). Only a single fly and no bees were collected on the last species. In contrast to the above, 10 of the 12 Aneilema species on which more bee than fly species were collected have glabrous filaments, and the other two have shortly bearded ones. The glabrous-filament species include all seven species of section Lamprodithyros (calceolus, clarkei, indehiscens, lamuense, petersii, succulentum, tanaense), both glabrous-filamented 50 SMITHSONIAN CONTRIBUTIONS TO BOTANY species of section Amelina (hockii, johnstonii), and the only species of section Rendlei (rendlei) that were sampled. The two bearded-filamented species (arenicola, umbrosum subsp. um- brosum) both belong to section Brevibarbata. On A. arenicola only a single bee and no flies were collected. From these data it may be concluded that flies, particularly Syrphidae, are attracted to and probably serve as the chief pollinators for Aneilema species with longly bearded stamen filaments; that bees are the main pollinators of the glabrous- filament species and possibly some short-bearded-filament species; and that the data for short-bearded-filament species is too inadequate to generalize about this group of species. The data for the three species of section Amelina are especially instructive: the two species with glabrous filaments are visited mainly by bees; the single species with long-bearded filaments is visited predominantly by flies. The occurrence of Coleoptera does not seem to follow any taxonomic or clear morphological pattern within Aneilem- perhaps the data are too incomplete-and it is probable that the beetles are merely feeding on the flowers, although some of them may occasionally effect pollination. Pollination of related, co-occurring species of Aneilema by different pollinators can lead to reproductive isolation. In a colony of five species, A. succulentum was found to not share any pollinators with the other species (Faden, 1983b). There are two pollination syndromes that require comment. In most species of section Lamprodithyros, the large medial petal retains the lateral stamens when the flowers opens, only releasing them when the margins of this petal uncurl (see ?Reproductive Morphology,? above, and Faden, 1983b). The stamen retention mechanism may promote outcrossing by the presentation, when a flower first opens, of only part of the flower?s total pollen to the pollinator, thereby forcing the insect to visit more flowers. The same mechanism may also prolong the period during which the flower can attract insects by the release of a second source of pollen-that of the lateral anthers-after the pollen of the medial stamen anther, which is always free, may have already been collected. Within section Lamprodithyros, there is much variation in the degree of development of the stamen retention mechanism. In species such as A. petersii and A. indehiscens, this syndrome appears to be primitively lacking; in A. tanaense, A. calceolus, and A. recurvatum, it is well developed; in A. succulentum it is poorly developed and perhaps disappearing; and in A. zebrinum it seems to have bcen lost entirely. These different degrees of development of this mechanism, perhaps in combination with diffcrences in flowering times, may result in reproductive isolation between related species (Faden, 1983b). The other syndrome of note is automimicry in A. rendlei and A. brenanianum. In those species the perfect and staminate flowers are the most dimorphic in the genus. The lateral stamens of the staminate flower are held close together in the same position as the style in the perfect flower, which they closely resemble in color, shape, and curvature, and so the two flower types strongly resemble one another (Figure 17a,b; Plate 2b,d). In A. rendlei it has been observed that the main insect visitors and presumed pollinators are workers of a very small, black bee (Trigona species). The bees have been observed to first land on the very prominent medial anther, collect pollen from it, and then, in the staminate flowers, to climb down the filaments of the lateral stamens in order to collect pollen from those anthers. In the field it has been observed that in populations of A. rendlei, staminate flowers are much more numerous than perfect flowers. Thus I believe that the bees are conditioned to the described behavior pattern, which they repeat when they visit perfect flowers and thereby transfer pollen to the stigma, with which they would otherwise not make contact because of the bees? diminutive size, All Aneilema species have characteristic daily flowering times (Figure 25). In cultivation it has been observed that these times may vary somewhat, according to the ambient tempera- ture, but the range of variation is normally quite small. I have noted in Faden (1983b) that flowering times in related, co-occurring species, e.g., A. petersii and A. succulentum, may be totally non-overlapping, thus preventing any gene exchange between the species. Aneilema flowers differ from those of some other genera of Commelinaceae, e.g., Commelina, in that the style does not roll up within the other floral parts when the flower fades. Thus, if the anthers are not in contact with or very close to the stigma when the flower is fully open, the flower will not self-pollinate, even in demonstrably self-compatible species such as A. rendlei. The species with autogamous flowers probably include all of the annuals in addition to many perennials such as A. umbrosum and A. zebrinum. Perennials, in addition to A. rendlei, that are not autogamous include A. hockii and A. johnstonii. Annual populations of A. tanaense, which have been observed in cultivation, regularly show autogamy. In contrast, a single perennial population of the same species, which has been cultivated for 10 years, has never set fruit without being intentionally pollinated. Self-compatibility at the level of the stigma and pollen grain in Aneilema has been reported on only by Owens (1981), who recorded all five species that he tested (A. aequinoctiale, A. brenanianum, A. sp. aff. spekei, A. zebrinum, A. sp.) as self-compatible. I have tried to deteremine compatibility by noting whether self-pollination leads to fruit set and and secd development. The results reported in Faden (1983b) (A. hockii, self-incompatible; A. petersii subsp. petersii, A. indehiscens subsp. indehiscens, and A. succulentum, possibly self- incompatible; and A. tanaense, self-compatible) were obtained in that way. Among those species, experimental pollinations were performed only on A. hockii and A. tanaense, hence the uncertainty about situation in the other taxa. There are no published data on seed dispersal in Aneilema. In general, the capsules and seeds do not show any clear adaptations for dispersal. Usually only one type of diaspore is produced, the individual ventral locule seeds, but in some species, e.g., A. pusillum and A. petersii (see ?Reproductive Morphology, Capsules,? above), a second type of disseminule, NUMBER 76 51 the deciduous, dorsal capsule valve containing the single, dorsal locule seed, is also formed. One may speculate that these two types of disseminules are dispersed by different means, particularly in species of section Lamprodithyros, where the dorsal and ventral locule seeds themselves may be strongly dimorphic. In A. tanaense a small number of hook-hairs is present on the dorsal valve, suggesting possible exozoochory (Faden, unpublished). It has also been found that the strongly dimorphic seeds in this species have different germination patterns (Faden and Teeri, unpublished). Taxonomy of Aneilema INTRODUCTION TO TAXONOMY The taxonomic treatment used here is based upon a study of a wide range of herbarium specimens (see ?Acknowledg- ments??) and, when available, living plants and preserved materials (see ?Materials and Methods?). All Aneilemu taxa have been investigated, and all types, except for the few destroyed during the Second World War, have been examined. This phase of the research led to an initial determination of species limits and synonymy. The list of species, thus established, has been used as a working hypothesis, being revised when necessary. It must still be considered tentative for those sections of the genus not thoroughly revised in the present study. Species recognized as distinct but undescribed were given specific names in Faden (1975), so they could be referred to readily. Some have been described subsequently (Faden, 1978a, 1984). Most of the others are described in this work; the remaining few will be formally described in future papers under the specific names used here. The large size of the genus prevented a study of all species in sufficient detail to understand their total variation. Therefore, once sectional limits were established, three sections were selected for more intensive investigation. Sections Rendlei and Lamprodithyros were chosen because large percentages of the taxa in each were known from living material, having been obscrvcd in the field and maintained in cultivation. Section Somaliensia was selected because its thitherto poorly known specics had to be thoroughly examined anyway, in order to dclimit the section. Keys to all species have been included. Those to the species in the four sections that have not been thoroughly revised should be considered provisional. In ordcr that no true Aneilemu species might be overlooked among the 260 names that have been included in the genus (see Appendix 11), types for most of these epithets have been examined. Particular emphasis has been placed on those taxa whose generic position was considered uncertain by Morton (1966) and whose geographic distributions and descriptions suggested possible valid inclusion in Aneilema, e.g., the Madagascan species of Perrier de la Bkhie (1936, 1938). The few specific epithets in Aneilemu for which the exact generic position has not bcen determined are very unlikely to belong to this genus, on the basis of their distributions and, in some cases, their descriptions. TAXONOMIC UNITS The species concept used here is fundamentally a taxonomic one (Davis and Heywood, 1964:91), the species being distinguished primarily on the basis of morphological disconti- nuities. Whenever possible an attempt has been made to refine the species limits, initially established on morphological criteria, through consideration of other kinds of data, such as distribution, ecology, phenology, breeding systems, karyo- types, and the result of crosses. The inclusion of such data has confirmed the preliminary taxonomic decisions in nearly all cases. Where the morphological data have not been supported by the biological data, the relative importance of each has had to be judged on a case-by-case basis. It is hoped that the use of a wide variety of characters in the final determination of specific limits will have made the resulting taxonomy as natural as possible within the bounds of the present state of our knowledge of these plants. Subspecies as used here may be defined as geographic segregates of a species that differ from one another in relatively few morphological characters. The morphological attributes that separate subspecies are presumed to not be in themselves of sufficient magnitude to prevent interbreeding. Subspecies may differ in biological characters, however, such as ploidy, that can be expected to have a more profound effect on potential gene exchange than their morphological differences. The ranges of subspecies may be discrete or overlapping. Varieties have not been used in the present treatment because the distinction between local variants, perhaps worthy of taxonomic rank, and odd specimens, not meriting formal recognition, is seldom clean-cut in Aneilemu. Minor variants and unusual specimens are dealt with in the discussion of each species. DISTRIBUTION OF Aneilemu The genus Aneilemu contains 62 species and occurs in three regions: eastern and northern Australia and the surrounding islands; tropical and subtropical Africa, Madagascar, and southern Arabian Peninsula; and tropical America (Map 1). Its absence from Asia except for two species occurring near the Red S e a is noteworthy. Aneilema is best represented in Africa with about 55 species, one of which extends to the Arabian Peninsula. One species is endemic to Madagascar and another to the Yemen Arabic Republic. Five species, all endemic, occur in the Australian region and only one in the neotropics. The latter, A. umbrosum, also occurs in Africa. Of the five Australo-Oceanic species, three are endemic to Australia, one (A. neocaledonicum) is confined to New Caledonia and adjacent islands, and the fifh (A. acuminatum, 52 SMITHSONIAN CONTRIBUTIONS TO BOTANY I I 3 I 0 9 0 N 0 0 3 0 c 3 3 5 2 3 0 > U 3 $ c 3 1 3 0 3 D NUMBER 76 53 0 400 MILES - 0 600 KILOMETERS Q 10 I I MAP 2.-Nmbers of Aneilema species in African countries. including A. papuanum) occurs throughout the region with the exception of New Caledonia. The number of Aneilema species in different parts of Africa shows considerable variation. The number of species per country is shown in Map 2. The greatest diversity of species is in the East African region extending from southern Ethiopia and central Somali Republic south to central Tanzania. Thirty-five species occur of which 18 are endemic to the region. A secondary concentration (15 species including five endem- ics) is located in southwestern Tanzania, southeastern Zaire, and northern Zambia. Another region of relatively high endemism (nine species, five endemic) occurs in southern Mozambique and southwestern Zimbabwe south to northern and eastern Transvaal and northeastern Cape Province, South Africa. The fourth such concentration is in the savannah region of West Africa (Sierra Leone and Senegal to Chad), where seven species (four endemic) occur. The species of this region include several very distinctive, infraspecific taxa, e.g., A. paluhsum A. Chevalier subsp. pseudolanceolatum J.K. Mor- ton, that may prove worthy of specific rank. It is significant that all four regions of relatively high species endemism (Map 3) are predominantly non-forested areas. The Congo Forest itself contains only five to six Aneilema species, most of which have wide ranges. All seven sections of Aneilema recognized herein occur in Africa or Madagascar. Section Aneilema, which includes all of the Australian species, is represented in the African region solely by the Madagascan endemic A. aparine. The remaining 54 S M I T H S O W CONTRTBUTIONS TO BOTANY h h ~ 3.4oncentrations of Aneilema species in Africa: 1 , East African center, 2, Zambian center. 3, Mozambique-South African center. 4, West African center. The shading is approximately proportional to the number of species in each area (see text). six sections are exclusively or predominantly African. All six have species in the East African center of diversity. Sections Rendlei and Somaliensia are wholly or largely confined to this region of endemism, while sections Amelina and Lamprodithy- ros include species endemic to it. The Zambian center contains species of sections Amelina, Brevibarbata, and Pedunculosa and endemics of the latter two. The Mozambique-South African center is comprised of species of sections Amelina, Brevibarbata, and Lamprodithyros including endemics of the first two. All of the species of the West African savannah region belong to section Brevibarbata. The distribution of Aneilema is unique in the Commelina- ceae. Only five other genera (Buforrestia, Commelina, Flo- scopa, Murdannia, Pollia) occur in both the Eastern and Western Hemispheres (Faden, 1978c, 1983a). Among them, Buforrestia is confined to Africa and South America. The other four occur in Australia, like Aneilema, but their ranges extend beyond that of the latter. Commelina is more or less cosmopolitan but best represented in Africa; Murdannia and Pollia are chiefly tropical and subtropical Asiatic, and Floscopa is primarily African but is also present in tropical Asia. It is significant that all six genera that occur in Africa and tropical America are better represented in the former, suggest- ing a dispersal from Africa to the neotropics (Raven and Axelrod, 1974). This is certainly correct for Aneilema. No genus is present in both Australia and Africa that does not also h?UMBER 76 55 occur in the neotropics. Among the 17 genera of Commelinaceae that occur in Africa and/or Madagascar, only Commelina has a distribution and concentration of species similar to that of Aneilema. Like that genus, Commelina contains relatively few forest species, the bulk of its taxa occurring in woodland, grassland, and bushland habitats. Although well represented in all four African regions in which Aneilema has concentrations of species, Commelina appears to have its greatest diversity in the Zambian region rather than in East Africa. Besides Aneilema and Commelina, only Floscopa among African Commelinaceae includes both forest and non-forest species. Unlike them, it occurs in primarily aquatic habitats. Its greatest concentration of species is in West Africa, with a secondary center in the Zambian region. In the East African and Mozambique-South African regions (as defined above) Flo- scopa is represented by a single widespread species, F. glomerata (J.A. & J.H. Schultes) Hasskarl. HISTORY Ah?) TYPIFICATION OF Aneilema The nomenclatural history of the genus Aneilema has been described in detail elsewhere (Faden, 1978b) and need only be summarized here. The genus was described by Robert Brown (1810) for 10 Australian species that are now recognized as belonging to three genera. The genus was considered synony- mous with Commelina by most workers until Kunth (1843) recognized it. Between 1843 and 1881 a number of generic segregates were described: Dictyospermum Wight and Dich- oespermum Wight (Wight, 1853), Lamprodithyros Hasskarl (Hasskarl, 1863), Rhopalephora Hasskarl (Hasskarl, 1864b), Piletocarpus Hasskarl and Prionostachys Hasskarl (Hasskarl, 1866), Bauschia Seubert (Seubert, 1872), and Amelina C.B. Clarke (Clarke, 1874). Clarke (1881a) returned all of these generic segregates to Aneilema in his monograph of the Commelinaceae. However, Clarke (1874) also narrowed the concept of Aneilema by removing from it two of Brown?s species, A. crispatum R. Brown and A. macrophyllum R. Brown, placing them in Polka, following Bentham and Mueller (1878) The concept of the genus remained unchanged until Briickner (1926) concluded that Clarke?s Aneilema included two disparate genera: Aneilema, which he typified by A. bijlorum, and Phaeneilema Briickner, which he described as new. Briickner (1930) subsequently recognized Phaeneilema as synonymous with Murdannia Royle and adopted the latter name. The fundamental distinction between Aneilema and Murdannia has been accepted by most authors with the notable cxccptions of Hutchinson (1934, 1959), who later did adopt it (Hutchinson, 1973), and Woodson (1942). Woodson?s Anei- lema was even broader than that of Clarke (1881a) and included, in addition to Murdannia, neotropical taxa that had never been placed in the genus and were totally unrelatcd to it. Unfortunately, Woodson?s nomenclature was adopted in a number of New World floristic works, and it has only slowly been abandoned. The most recent change in the generic concept of Aneilema was that of Morton (1966), who transferred all of the southeastern Asiatic species to Dictyospermum or Tricarpe- lema J.K. Morton. I agree with Morton?s circumscription of Aneilema, but I delimit the Asiatic genera that appear to be related to Aneilema differently (see ?Generic Relationships of Aneilema,? below). In reviewing the typification of Aneilema, I came to the conclusion that the retypification of the genus by either A. gramineum R. Brown or A. giganteum R. Brown-now both treated as species of Murdannia-which could be justified on historic grounds, should be rejected because of the severe nomenclatural dislocation that would ensue and, strictly speaking, it is not necessary (Faden, 1978b). I accept A. bijlorum as the lectotype of the genus, although it was selected in an arbitrary manner by Briickner (1926). The history of the species of Aneilema sensu strict0 is somewhat different from that of the genus. Except for a few that were described in genera such as Commelina, Bauschia, Amelina, Lamprodithyros, or Floscopa, nearly all species were first described in Aneilema. The first monograph of all of the known species was by Kunth (1843), who, as noted above, used a broad generic concept. Similarly, the next and last monograph to include all of the species was that of Clarke (1881a) whose subgenus Dicarpellaria is equivalent to Aneilema sensu stricto. All of the African species were monographed by Clarke (1901), who described many new ones. Since then many additional African species have been described, especially by Wildeman (1913, 1915), Chiovenda (1916, 1928, 1936), Brenan (1952, 1961, 1968), and Faden (1978a, 1984), and other species have been clarified, particularly by Morton (1966, 1967), Brenan (1952, 1961), and Faden (1984), but only regional floristic works have been published. A list of all specific names in Aneilema and their current status is given in Appendix 11. GENERIC RELATIONSHIPS OF Aneilema The generic relationships of Aneilema have been discussed in detail (Faden, 1975) and will be published separately. Therefore, only a summary will be included here. Aneilema appears to be most closely related to a group of three exclusively or mainly Asiatic genera that show the same one-sided stamen amangement, i.e., the three stamens (or three longer stamens) attached in front of the outer and usually lower petal and adjacent (inner) sepals. The three staminodes or smaller stamens, when present, are attached on the opposite side of the flower. Among these genera, Aneilema is undoubtedly more closely related to Rhopalephora Hasskarl than to Dictyospermum Wight or Tricarpelema J.K. Morton. Rhopalephora, a gcnus of about three ill-defined species that range from India and Sri Lanka throughout southeast Asia to New Guinea and the Fiji 56 SMITHSONIAN CONTRIBUTIONS TO BOTANY Islands plus one species in Madagascar (Faden, 1977), agrees with Aneilema (and differs from the other two genera) in having bivalved capsules that are bilocular or unequally trilocular (vs. trivalved and equally trilocular), perfoliate (vs. non-perfoliate) bracteoles, and clawed (vs. non-clawed) petals. In fact, Rhopalephora is so similar to Aneilema that it must be defined by a combination of characters, none of which is unique to it (except for the basic chromosome number x = 29). Dictyospermum (Dictyospermum section Dictyospermum sensu Faden, 1975) consists of four to five species that are distributed from southern India and Sri Lanka to New Guinea. In addition to the characters cited above, it differs from Aneilema by its equal petals, absence of staminodes, and strictly one-seeded capsule locules that contain coarsely reticulate seeds. Tricarpelema (Dictyospermum section Tricarpelema sensu Faden, 1975), the least understood of these genera, contains six species that range from the foothills of the Himalayas in India to the Philippines and Borneo, and possibly a seventh species in western Africa (Cameroon and Gabon). It differs from Aneilema by its shorter, upper androecium-members some- times polliniferous and in its lack of hook-hairs, plus the features mentioned above. Another species in this alliance, described as Floscopa yunnanensis Hong (Hong, 1974) from Yunnan Province, China, but clearly not belonging to that genus, is of uncertain generic position. It seems closer to Aneilema and Rhopale- phora than to Dictyospermum or Tricarpelema, but it cannot be comfortably accommodated in any of these genera. It is still inadequately known, having been collected only once. A detailed account of its morphology and possible relationships is given in Faden (1975). Two species that must either be placed in Aneilema or else segregated as monotypic, closely related genera are A. calandrinoides F. Mueller of Australia and A. brasiliense C.B. Clarke of Brazil and Venezuela. They are discussed at the end of the taxonomic treatment under ?Species of Uncertain Generic Position.? The genus Pollia may be the sister group for the entire alliance of genera and species discussed above. Although it includes species with the same one-sided arrangement of stamens and staminodes as in Aneilema and the other genera, Pollia also contains other species that have six equal, fertile stamens. The one-sided stamen and staminode arrangement has evolved within Pollia, and thus the genus is convergent rather than synapomorphic with the Aneilema alliance genera and spccics wih regard to this character. The genus Murdannia must be mentioned because of the historic placement of its species in Aneilema (see Faden, 1978a). Its alternating stamen and staminode arrangement, however, is clearly a separate, derived state from the androecial structure in Aneilema and related genera. Murdannia also exhibits further unusual and apparently derived characters, such as three-lobed anherodes (first noted by Brenan, 1952) (Figure 43) and a continuous adaxial hypodermis (Tomlinson, 1966), that are not present in the other genera and that indicate an early separation from Pollia and the taxa of the Aneilema alliance. Aneilema has also been considered closely related to a b e f n 9 C d h FIGURE 43.-Antherodes of Aneilema and Mwdannia. Aneilema (0-4: a, A. petersii (Hasskarl) C.B. Clarke subsp. jmllidiflorwn Faden, lateral antherode (from Faden et al. 6911068); b, A. umbrosurn (Vahl) Kunth subsp. ovato-oblongurn (P. de Beauvois) J.K. Morton, lateral antherode (from Lye & Katende 6235); c, A. rendlei C.B. Clarke, medial antherode (from Faden et al. 69/322); d, A . hockii De Wildeman, lateral antherode (from Faden et al. 72/234). Mwdannia (4): e, M. japonica frhunb.) Faden (after Clark, 1904, as Aneilema lineolatum (Blume) Kunth);f, M. clarkeana Brenan (from Faden & Evaru 69/304); g , M. simplex (Vahl) Brenan (from Faden et al. 74/64); h, M . zeylanica (C.B. Clarke) Briickner (from Davidse s.n.). (Not to scale.) NUMBER 76 57 Commelina by Rohweder (1969) on the basis of 13 shared characters. I have rejected this argument for three reasons: (1) some of the characters used, such as the minor anatomical ones, are of unknown phylogenetic significance, i.e., the primitive and derived conditions are unclear; (2) some characters used, e.g., presence of a terminal inflorescence, are plesiomorphic and are therefore not indicative of an evolutionary relationship; and (3) some similarities, particularly those of the androecium, may be due to convergence (Faden, 1975). Finally, it must be stressed that that a cladistic analysis of all of the genera that comprise the tribe Commelineae (sensu Faden, 1985), to which Aneilema and all of the above- mentioned genera, as well as others, belong, is urgently needed. Until that is undertaken, the results of this present analysis of the relationships among some of these genera must be considered preliminary. CITATIONS OF Aneilema Aneilema R. Brown, Prod. Fl. Novae-Hollandiae, 270, 1810, p.p.-P. de Beauvois, Fl. Owar., 256, 1818, p.p.-Bentham in Hooker and Bentham, Fl. Nigrit., 546, 1849, p.p,-- Bentham and Mueller, Fl. Austral., 7:85, 1878, pap. -Clarke, J. Bot., 19:199, 1881, p.p.-Clarke in DC, Monogr. Phan., 3:195, 1881, p.p.-Bentham and Hooker, Gen. Pl., 3:849, 1883, p.p.-Bailey, Synopsis Queensland Fl., 556, 1883, p.p.-Schumann in Engler, Pflanzenwelt Ost-Afrikas C, 135, 1895, p.p.-Durand and Schinz, Conspect. Fl. Africae, 5:429, 1895, p.p.-Durand and Schinz, Etud. F1. Congo, 1:270, 1896, p.p.--Clarke in Thiselton-Dyer, Fl. Capensis 7, 1: 12, 1897, p.p.-Durand and De Wildeman, Bull. SOC. Roy. Bot. Belgique, 36237, 1897, p.p.; 37:128, 1898, p.p.-De Wildeman and Durand, Ann. Mus. Congo Bot., S6r. 2, 1:62, 1899-1900, p.p.- Clarke in Thiselton-Dyer, Fl. Trop. Africa, 8:62, 1901, p.p.-Bailey, Queensland Fl., 5652, 1902, p.p.-Durand and Durand, Syll. F1. Congo., 579, 1909, p.p.-Bailey, Cat. Queensland Pl., 561, 1913, p.p.-Chevalier, Etudes Afr. Cent. Franc., 1:328, 1913, p.p.-Thonner, Fl. P1. Africa, 122, 1915, p.p.-Chiovenda, Result. Scient. Miss. Stefan- iniPaoli Somalia Italiana, 1:167, 1916.Ewart and Davies, Fl. Northern Tcrr., 69, 1917, pep.-Chevalier, Explor. Bot. Afr. Occident. Franc., 1:666, 1920, p.p.-Briickner, Bot. Jahrb. Syst., Beibl., 137:62,1926, p.p. maj.-Chiovenda,Fl. Somala, 1:3 16, 1929.-Briickner in Engler, Naturlich. Pflanzenfam., ed. 2, 15a:174, 1930, p.p. maj.-Hutchinson, Fam. F1. Pl., 2:55, 1934, p.p.-Hutchinson and Dalziel, Fl. W. Trop. Africa 2,2:312, 1936, p.p.-Perrier de la Bithie, Notul. Syst. (Paris), 5:193, 1936, p.p. min.-Denier de la B%thie, Fl. Madagascar, 23, 1938, p.p. min.-Chiovenda in Zavatti, Miss. Biol. Paese Borana, 4:304, 1939.-Pichon, Notul. Syst. (Paris), 12:238,1946, p.p. maj.-T%ckholm and Drar, Fl. Egypt, 2433, 1950.-Chiovenda, Webbia, 8:38, 1951.-Brenan, Kew Bull., 7: 180, 1952, p.p. maj.-Robyns and Tournay, Fl. Spermat. Parc Natl. Albert, 3:326, 1955.-Andrews, Fl. P1. Sudan, 3:238, 1956.-Troupin, Fl. Spermat. Parc Natl. Garamba, 1: 158, 1956.-Hutchinson, Fam. Fl. Pl., ed. 2, 2565, 1959, p.p.-Panigrahi and Kammathy, Proc. Natl. Acad. Sci. India, Sect. B, 33:491, 1963, speciebus exc1.-Brenan, Kew Bull., 19:63, 1964, p.p. maj.-Aristeguieta, Bol. Acad. Cien. Fls. Matemit. Nat. Caracas, 25134, 1965.-Compton, J. S. Afr. Bot., uppl., 633, 1966.-Morton, J. Linn. Soc., Bot., 59:431, 1966.- Schreiber in Merxmiiller, Prod. Fl. Siidwestafrika, 157:2, 1967.-Morton, J. Linn. SOC., Bot., 60:167, 1967.-Brenan, Kew Bull., 22:387, 1968.-Brenan in Hepper, Fl. W. Trop. Africa, ed. 2,3,1:29, 1968.-Rohweder, Ber. Schweiz. Bot. Ges., 79:216, 1969.-Cufodontis, Bull. Jard. Bot. Natl. Belgique, 41, suppl., 1516, 1971.-Lebrun, Andru, Gaston, and Mosnier, Inst. Elevage Med. VCt6rin. Pays Trop. Etude Bot., 1:185, 1972.-Ross, Fl. Natal, 117, 1972.Faden in Agnew, Upland Kenya Wild Fl., 664, 1974.-Faden, Bothalia, 12565, 1978.-Faden, Taxon, 27:289, 1978,- Faden, Fl. P1. Africa, 43: 1785,1979.Faden and Suda, Bot. J. Linn. SOC., 81:301,1980.-Faden in Morley and Toelken, Flowering Plants in Australia, 368,1983.-Faden, Bothalia, 14553, 1983.-Faden, Bothalia, 14:997, 1983.-Faden, Bothalia, 1589, 1984.4bermeyer and Faden in Leistner, Fl. Southern Africa, 4(2):23, 1985. SYNONYMY OF Aneilema Commelinu auct., pap. [non L. (175311. Anilema Kunth. Enum. Pl., 4:64, 1843 [ o h . var. of Aneilema R. Brown]. Lumprodithyras Hasskarl. Flora, 46:388, 1863; in Peters, Natunviss. Reise Mossambique, Bot.. 529, 1864. [Lectotype: L. petersii Hasskarl = Aneilema petersii (Hasskarl) C.B. Clarke.] Perosanthera Fend, Sitzungsb. Akad. Wien. 50:353. 1864 [nm. nud. in comb.] Baurchia Seubert in Warning, Vidensk. Meddel. Kj6b., 1872:123, 1872. [Type: B . bracteolata (Martius) Seubert = Aneilema umbrasum (Vahl) Kunth.] h l i n u C.B. Clarke, Commel. et Cyrt. Beng., 38. tab. 26, 1874. ['Qpe: A. wallichii C.B. Clarke = Aneilema aequinmtiale (P. de Beauvois) G. Don.] Aneilema subgenus Dicarpellaria C.B. Clarke in De Candolle, Monogr. Phan.. 3:196, 1881. Ballya Brenan, Kew Bull., 19:63, 1964. [%: 8. rebrina (Chiovenda) Brenan = Aneilema zebrinwn Chiovenda.] DESCRIPTION OF Aneilema Perennial and annual herbs of various habits. Roots fibrous or tuberous. Leaves spirally arranged or distichous, sheaths entire, usually ciliate at the apex, laminae petiolate or sessile, usually herbaceous (rarely succulent). Uppermost leaf (inflo- rescence bract) on the flowering shoot often very reduced. Inflorescences thyrses, terminal, terminal and axillary, or rarely, all axillary, consisting of a central axis bearing individual cincinni, rarely reduced to a single cincinnus. Cincinni always subtended by persistent bracts, which are often glandular near the apex. Bracteoles 2-ranked on the cincinnus, 58 SMITHSONTAN CONTRIBUTIONS TO BOTANY frequently cup-shaped, perfoliate, usually glandular near the apex, persistent. Flowers pedicellate, perfect, perfect, and staminate, or, rarely, perfect, staminate, and pistillate, all types produced in the same inflorescence. Sepals 3, free, sepaloid, persistent, convexo-concave, usually hooded apically, particularly the medial one, predominantly green, usually glandular near the apex, glabrous or puberulous. Petals 3, free, petaloid, deliques- cent, subequal and concolorous or, more commonly, the medial one much reduced (rarely enlarged) and discolorous; paired petals usually held upwards, always unguiculate, white to lilac or lavender, or yellow to orange, rarely red, claws glabrous, rarely puberulous; medial petal usually held downwards, cuneate at base, rarely unguiculate, commonly dull colored and inconspicuous, rarely puberulous basally. Staminodes 3(-2), opposite the medial sepal and paired petals, equal or, more commonly, the medial (antesepalous) one reduced (sometimes vestigial or absent), when present its antherode often different in size and/or shape from those of the lateral staminodes; antherodes bilobed (rarely unlobed), the lobes stipitate or sessile, filaments always glabrous. Fertile stamens 3, opposite the paired sepals and medial petal, equal or, more commonly, the medial (antepetalous) one much shorter than the lateral (antesepalous) ones, all filaments glabrous, or the laterals sparsely to densely bearded with uniseriate, non-moniliform hairs; anthers basifixed, dorsifixed or versatile, dehiscence longitudinal, usually introrse, rarely latrorse or extrorse; anther of the medial stamen usually different in size and/or shape from those of the lateral stamens, its pollen often also different in color and sometimes inviable. Ovary stipitate or sessile, bi- or trilocular, glabrous or, more commonly, puberulous with glandular hairs (occasionally mixed with few to many hook-hairs), dorsal locule not developed or with one (rarely more) ovule(s), ventral locules each with 1-6 uniseriate ovules; style simple, stigma capitate or not enlarged. Fruit capsular, usually dehiscent, rarely k indehiscent, bi- or trilocular, bi- or (very rarely) trivalved, usually gray to brown, generally lustrous, glabrous, or puberulous, valves persistent or the dorsal one deciduous, dorsal locule (when present) empty or with one (rarely more) seed(s), ventral locules each with up to 6 uniseriate seeds. Seeds with a linear hilum and lateral embryotega, testa various, often with farinose granules. TYPE SPEcIES.-Aneilema biflorum R. Brown (see Faden, 1978b). DIVISION OF Aneilema INTO SECTIONS The first formal division of Aneilema into sections was proposed by Clarke (1881a:196), who used the following scheme: Subgenus I. Tricarpellaria. Capsule subequally 3-1wular, 3-valved. Section I. Euaneilema. Ovary with locules 2-many-ovulate; seeds 1-senate. Section 11. Dichaespermum. Ovary with locules many-ovulate; seeds 2-senate. Section 111. Dictyospermum. Ovary with locules 1 -ovulate. Subgenus 11. Dicarpellaria. Capsule 2-locular, 2-valved (a third smaller locule Section IV. Amelina. Capsule with apex truncate. 2-homed, with locules Section V. Lamprodithyros. Capsule with apex obtuse rounded, with locules rarely added). 3-5-seeded. 2-several-seeded (1 -seeded in A. siliculmum). Clarke?s subgenus Tricarpellaria does not include any species of Aneilema in the circumscription of the genus used here. Its species are assignable to Murdannia Royle, Dictyospermum Wight, Tricarpelema J.K. Morton, and Anthericopsis Engler. Subgenus Dicarpellaria equals Anei- lema in the present sense. Clarke?s sections Amelina and Lamprodithyros are the earliest published names for sections of Aneilema in the sense used here. Clarke?s divisions were accepted by Bentham and Hooker (1883) as ?apparently well limited although not always natural.? They were also used but evidently not understood by Schumann (1895), who placed two of his three just described species of Aneilema in section Euaneilema instead of section Lamprodithyros, where they should have been put according to Clarke?s characters. Clarke (1901:62-63) slightly revised subgenus Dicarpel- laria by dividing it into three instead of two sections as follows: Amelina: Panicles terminal. Capsule truncate, with 2 acute comers. Lamprodithyros: Panicles terminal. Capsule ellipsoid without angles at the top. Pseudo-axillares: Panicles appearing irregularly scattered, all terminal, but not rarely on remote sholt axiUary branches. Morton (1966) followed Clarke (1881a) in using two sections, Amelina and Lamprodithyros, defining them in the same way. He did not accept section Pseudo-axillares of Clarke (1901), however, stating that its species ?fall naturally into the section Lamprodit hyros.? I have found that the schemes proposed thus far for dividing Aneilema into sections are artifical and totally inadequate for expressing the natural relationships among the species in the genus. The single character used by Clarke (1881) and Morton (1966)-shape of the capsule apex-would in some cases separate closely related species, e.g., A. hockii and A. longirrhizum, and in others associate only distantly related ones, e.g., A. pedunculosum and A. acuminatum. My detailed investigation of characters of potential use in defining sections has revealed that, in general, floral attributes are more indicative of natural relationships in Aneilema than are capsular characters. However, capsule, seed, inflorescence, and, to a lesser extent, vegetative characters have also been found to be useful in some cases. These characters have been employed as much as possible because of the generally poor preservation of floral parts in dried specimens. In the following scheme, section Aneilema is recognized for the first time, Clarke?s section Amelina has been redefined and somewhat expanded, and his section Lamprodithyros has been greatly restricted in scope but still includes his section Pseudo -axillares. In addition, four new sections have been NUMBER 76 59 described. The seven sections proposed in this scheme are each held together by numerous morphological characters. Moreover, each section has a distinctive geographic distribution and generally only one or two basic chromosome numbers. Most sections also have characteristic patterns of floral hair types on the different floral whorls. No subsections or series have been recognized, although within some sections there are species groups that might warrant such formal taxonomic recognition on the basis of future investigations. Key to the Sections of Aneilema 1. Medial petal equal to or larger than a lateral petal, usually concolorous with it; stamen filaments glabrous; all 3 anthers well developed; Australo-Oceanic, African, and Arabian Peninsular species. 2. Ovaries and capsules glabrous; medial petal not boat-, slipper-, or cup-shaped; filaments free; seeds with a rugose testa, lacking farinose granules; 2. Ovaries and capsules puberulous; medial petal boat-, slipper-, or cup-shaped; stamen filaments fused basally; seeds with a scrobiculate to reticulate testa, usually farinose in the depressions; continental African and Arabian Peninsular 1. Medial petal usually much smaller than a lateral petal and differently colored from it (if subequal, then lateral stamen filaments bearded and medial anther reduced); stamen filaments glabrous or bearded; anther development various; African and American species. 3. Capsules glabrous; medial anther large, elliptic to ovate, with a strongly convex, maroon-spotted connective; lateral stamen filaments of staminate flower and Australo-Oceanic and Madagascan species . . . . . . . . 1. Section Aneilema species . . . . . . . . . . . . . . . . . . . . . . . 5 . Section Lamprodithyros style of perfect flower J-shaped; filaments glabrous; stigma not enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Section Rendlei, new section 3. Capsules puberulous or glabrous; medial anther various, but not as above; curvature of lateral stamen filaments and style various, but not J-shaped, filaments bearded or glabrous; stigma usually distinctly capitate (not capitate in section Somaliensia). 4. Inflorescences lax with cincinni mostly subopposite or subverticillate (mostly alternate in A. longirrhizum); flowers usually medium to large, 12-40 mm wide (if less, then paired petals yellow to orange); lateral stamen filaments straight or undulate, not geniculate . . . . . . . . . . . 2. Section Amelina 4. Inflorescences dense to moderately lax (rarely lax) with mostly alternate cincinni; flowers small to medium, 4-15(-20) mm wide; lateral stamen filaments S-shaped, k geniculate. 5 . Stamen filaments glabrous; tuberous-rooted perennials generally of dry bushland (habit and roots of A. grandibracteolatum unknown); flowering shoots often disarticulating at the nodes at the end of the growing season . . . . . . . . . . . . . . . . . . . .4. Section Somaliensia, new section 5. Stamen filaments bearded; perennials or annuals, fibrous or tuberous-rooted, generally of grassland or forest, rarely of bushland; flowering shoots not disarticulating at the nodes. 6. Lateral stamens usually sparsely and inconspicuously bearded; bracteoles cup-shaped, generally perfoliate, lacking linear apices; cincinnus axes usually elongate (except in A. paludosum and A. mortonii); tuberous roots sometimes present. . . . . 6 . Section Brevibarbata, new section 6. Lateral stamens densely and conspicuously bearded; bracteoles neither cup-shaped nor perfoliate, often with linear, gland-tipped apices; cincinnus axes usually not elongate, rarely somewhat elongate; tuberous roots absent . . . . . . . . . . . 7. Section Pedunculosa, new section 60 SMITHSONIAN CONTRIBUTIONS TO BOTANY Section 1: Aneilema Perennial or annual herbs with fibrous or tuberous roots. Leaves spirally arranged, with sheaths usually lacking long cilia at the apex. Inflorescences thyrses, terminal on the main or short lateral shoots, lax to moderately lax. Inflorescence axes puberulous with hook-hairs of uniform length. Bracteoles herbaceous, cup-shaped, perfoliate. Flowers all perfect or perfect and staminate. Fruiting pedicels erect, laterally spreading, or sharply decurved. Petals subequal, concolorous, white to blue-violet, glabrous, medial petal neither cup-shaped nor retaining the lateral stamens. Filaments free. Staminodes equal or subequal, antherodes unlobed or bilobed, yellow or white to violet. Stamens equal or the medial shorter than the laterals and its anther of a different form, filaments straight to very gently S-shaped, glabrous. Ovary sessile, glabrous, dorsal locule not developed or 1--5-ovulate, ventral locules each 1 --5-ovulate; stigma capitate. Capsules sessile, dehiscent, bi- or trilocular, bi- or trivalved, lustrous, glabrous, apex acute or rounded to truncate or emarginate, valves persistent and spreading to -180' or dorsal valve deciduous, dorsal locule (when present) 1 --heeded, ventral locules each 1--5-seeded, cells of the capsule wall transversely elongate. Seeds with hilum short, testa rugose- tuberculate, lacking farinose granules. TYPE SPECIES.-Aneilema biforum R. Brown (see Faden, 1978b). BASIC CHROMOSOME NUMBER.+ = 16. DISTRIBUTION.-'krnate Island (Indonesia) to the Solomon Islands, S to Northern Territory, N and E Queensland, E New South Wales, Australia, New Caledonia and the New Hebrides; also in Madagascar. Key to the Species of Section Aneilema 1. Leaves sessile; roots tuberous; capsules with ventral locules 1-seeded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. siliculosum 1. Leaves petiolate; roots fibrous; capsules with ventral locules 2-several seeded. 2. Sepals and pedicels puberulous; ventral locules of capsule 2-seeded. 3 . Plants not rhizomatous; inflorescences 10-25 cm long, with (4-)8-18 cincinni; dorsal capsule valve persistent; Madagascar. . . . . . . . . . . A. aparine 3. Plants rhizomatous; inflorescences 2.5-6 cm long, with (1-)2-6 cincinni; dorsal capsule valve deciduous; Australia . . . . . . . . . A. sclerocarpum 4. Inflorescences consisting of 1-4 cincinni, borne on short lateral shoots; fruiting pedicels deflexed. 5. Leaves linear to linear-lanceolate; flowers 1.2-2.0 cm wide; anthers linear to linear-oblong; pollen white . . . . . . . . . . . . . . A. neocaledonicum 5. Leaves lanceolate; flowers less than 1 cm wide; anthers ovate to 4. Inflorescences consisting of 4-many cincinni, terminal on the main and major lateral shoots; fruiting pedicels horizontal to erect . . . . . A. acuminatum 2. Sepals and pedicels glabrous; ventral locules of capsule 2-5-seeded. ovate-elliptic: pollen yellow . . . . . . . . . . . . . . . . . A. biflorum (including A. papuanum) SPECIES CITATIONS AND DISTRIBUTIONS Aneilema acwninaium R. Brown, Prod. Fl. Novae-Hollandiae, 270, 1810 [temate to the Solomon Islands S to Australia (Queensland. NSW) (Plate Aneilema aparine H. Pemer de la Bathie, Notul. Syst. (Pans), 5:1%, 1936 Aneitema bijlorum R. Brown, Prod. Fl. Novae-Hollandiae, 270,1810 [Australia Aneilema neocaledonicum Schlechter, Bot. Jahrb. Syst., 3928, 1906 [New Aneilema scterocarpum F. Mueller, Frag., 861,1873 [Australia (Queensland)]. Aneilemu siliculosum R. Brown. Prod. Fl. Novae-Hollandiae, 270, 1810 la,b)l. [Madagascar]. (Qucensland, NSW)]. Caledonia and the New Hebrides (Plate lc)]. [Australia (N 'lkmtory, N Queensland)]. DISCUSSION The diversity in morphology and wide geographic distribu- tion of this section suggest an old radiation. The major taxonomic problem is the final disposition of A. papuanum, which is tentatively included within A. acuminatum herein. Aneilema papuanum is geographically almost completely discrete from typical A. acuminatum and thus merits at least subspecific rank, but further study is required to determine whether it should be accepted as a full species. The placement of the Madagascan A. aparine in section Aneilema is based on a congruence of many characters, including floral morphology NUMBER 76 61 as determined by dissection of buds on a herbarium specimen. Section 2: Amelina (C.B. Clarke) C.B. Clarke Aneilema section Amelina (C.B. Clarke) C.B. Clarke in De Candolle. Monogr. Phan., 3:197. 1881; in Thiselton-Dyer, Fl. Trop. Africa, 8:62, 1901.- Morton, J. Linn. Soc., Bot.. 59:439, 1966. [Lectotype: Aneilema aequinoc- t ide (F? de Beauvois) G. Don.] Amelina C.B. Clarke, Commel. et Cyrt. Bengal., 39, tab. 26, 1874, pro gen. [type: Amelina wallichii C.B. Clarke (= Aneilema aequinoctiale (p. de Beauvois) G. Don).] Perennial or rarely annual herbs with fibrous or tuberous roots. Leaves spirally arranged or distichous. Inflorescences thyrses, terminal on the main shoots and occasionally on shorter lateral shoots, lax, with mostly subopposite or subverticillate (rarely mostly alternate) cincinni. Inflorescence axis and cincinnus peduncles and axes usually puberulous with hook-hairs of two sizes (rarely all glabrous). Cincinni elongate. Bracteoles usually herbaceous and cup- shaped, perfoliate (rarely membranous and not perfoliate). Flowers mostly relatively large (10-40 mm wide), perfect and staminate (rarely pistillate, perfect, and staminate). Fruiting pedicels erect (rarely recurved 270?), persistent. Lateral sepals narrow, generally lanceolate to lanceolate-elliptic, commonly puberulous with hook-hairs of two sizes (rarely glabrous or with uniform hook-hairs). Paired petals lilac to lavender or yellow to orange, claws glabrous or puberulous. Medial petal reduced, usually inconspicuous, rarely concolorous with the paired petals. Filament bases usually free (rarely fused). Staminodes with yellow, bilobed antherodes; medial staminode very short, its lobes usually larger than those of the lateral staminodes (rarely all three staminodes short and their antherodes equal). Lateral staminodes with filaments straight to arcuate, sometimes strongly recurved near the apex. Lateral stamens with filaments straight or undulate, parallel or divergent, glabrous or bearded. Medial stamen with filament strongly ascending apically, anther very reduced or large and saddle- or shield-shaped, when large, commonly with a contrasting dark spot or bar on the yellow connective. Ovary sessile or stipitate, very sparsely to densely puberulous, usually glabrous on the lateral and middorsal sutures, dorsal locule not developed or 1 -ovulate, ventral locules each 2-6-ovulate; style straight or undulate, sometimes strongly recurved apically, usually held k in the floral midplane, stigma capitate. Capsules sessile or stipitate, dehiscent, bivalved, bi- or trilocular, lustrous or dull, apex acute to rounded, truncate or emarginate, valves persistent and spreading up to 180?, or occasionally dorsal valve deciduous, dorsal locule (when present) 1-seeded or, by abortion, empty; ventral locules up to 6-seeded (rarely regularly empty by abortion), cells of the capsule wall isodiametric or transversely elongate. Seeds usually oblong to trapezoidal (occasionally ovate or elliptic), apical ventral generally rounded apically, basal ventral gener- ally rounded to truncate basally, the apical and basal seeds therefore usually little differentiated in shape (rarely strongly differentiated), hilum commonly not in a groove (occasionally in a groove), often slightly extended onto apical and basal surfaces, testa smooth to alveolate, shallowly foveate, reticu- late-foveate or scrobiculate, farinose granules present or absent. BASIC CHROMOSOME NUMBERS.+ = 13, 15, and 16. DIsTRIBUn0N.-Throughout the continental African range of the genus except for the extreme N and NE; better represented on the eastern side of the continent than in the west. Key to the Species of Section Amelina 1. Inflorescence axis and cincinnus peduncles and axes glabrous; paired petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. johnstonii 1. Inflorescence axis and cincinnus peduncles and axes puberulous; paired petal color, stamen filament pubescence and habit various, but not in the above combination. 2. Bracteoles not perfoliate, membranous; capsule glabrous or subglabrous, apex 2. Bracteoles perfoliate, herbaceous; capsule puberulous, apex rounded to truncate or emarginate; stamen filaments glabrous or bearded, if bearded, then not solely in basal half. 3. Leaf margins smooth: paired petals pale lavender; fruiting pedicels recurved 270'; seeds 2 per ventral locule, densely covered with white farinose granules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. longirrhizum 3. Leaf margins scabrid; petal color various; fruiting pedicels erect or spreading laterally; seeds 0-6 per ventral locule, lacking white farinose granules. 4. Paired petals lilac or lavender; stamen filaments bearded: capsules strongly compressed laterally, rounded apically, dorsal locule with 1 very large, smooth seed -6 mm long, ventral locules abortive, cells of the capsule wall isodiametric . . . . . . . . . . . . . . . . . . . . . . . . A. plagwcapsa yellow-orange; stamen filaments glabrous; tuberous rooted perennials . . . . . . acute; stamen filaments finely bearded in basal half. . . . . . . . . A. gillettii 62 SMITHSONIAN CONTRIBUTIONS TO BOTANY 4. Paired petals lavender or yellow to orange; stamen filaments bearded or glabrous; capsules not laterally compressed, truncate to emarginate apically, dorsal locule not developed or with 1 seed less than 3 mm long, ventral locules 1-&seeded, cells of the capsule wall isodiametric or transversely elongate. 5. Paired petals lavender; stamen filaments glabrous; roots tuberous or fibrous; capsules with ventral locules 4-6-seeded (occasionally less by abortion), cells of the wall transversely elongate . . . . . . . A. hockii 5. Paired petals yellow to orange-yellow; stamen filaments bearded or glabrous; roots fibrous; capsules with ventral locules l-3-seeded, cells of the wall isodiametric. 6. 6. Perennials with distichous leaves; stamen filaments bearded. Annuals with spirally arranged leaves; stamen filaments glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ephemerum 7. Sheaths with whitish hairs at the apex; flowers -18-30 mm wide; claws of the paired petals bearded; pollen orange: seeds 3 per ventral locule . . . . . . . . . . . . . . . . . . . . . . . A. aequinoctiale 7. Sheaths with orange or reddish orange hairs at the apex; flowers -10-15 mm wide; claws of the paired petals glabrous; pollen white; seeds 1-2 per ventral locule . . . . . . . . . . . . . . A. nyasense SPECIES CITATIONS AND DISTRIBUTIONS Aneilema aequinoctiale (P. de Beauvois) G. Don in Loudon, Hort. Brit., 15, 1830 Wsionym: Commelina aequinoctialis P. de Beauvois, Fl. Owar, 1:65, tab. 38, 1806; S Ethiopia to Guinea S to Angola, Zimbabwe, and South Africa (Natal, Cape, Transvaal) (Plate li)]. Aneilema ephemerum Faden, ined. [Faden & Faden 741794; N Kenya]. Aneilema gillettii Brenan, Kew Bull., 15212, 1961 [S Ethiopia and N Kenya Aneilema hockii De Wildeman, Feddes Repert. Spec. Nov. Regni Veg., 12:290, 1913 [S Ethiopia S to Mozambique.South Africa (Transvaal), Botswana, and Namibia (Plate lg,h,l)]. Aneilema johnrfonii K. Schumann in Engler, F'flanzenwelt Ost-Afrikas C, 135, 1895 [S Ethiopia S to Mozambique, Zimbabwe and Botswana (Plate Id)]. Aneilema longirrhizum Faden, Botharia, 12(3):565, 1978 [South Africa (Transvaal) (Plate lk)]. Aneilema nyasense C.B. Clarke in Thiselton-Dyer, Fl. Trop. Africa, 8:66,1901 [w Zaire to Uganda S to Burundi. Malawi, and Mozambique (Plate lj)]. Aneilema plagiocapsa K. Schumann in Warburg, Kunene-Sambesi-Exped., 184. 1903 [Zambia, S Zaire, and Angola]. (Plate le,f)l. DISCUSSION The species of section Amelina are very diverse, and their relationships to one another are often difficult to ascertain. This variability suggests that either the section is not monophyletic or, more likely, I believe, the extant species represent endpoints of an old radiation. This section is the only one that includes some species with glabrous lateral stamen filaments and others with bearded filaments. The occurrence of three basic chromosome numbers in section Amelina is shared only with section Brevibarbata, which has far more species. A number of unique characters for the genus occur in one or two species of section Amelina, e.g., paired petals claws puberulous (A. uequinoctiale and A. ephemerum). Indeed, nearly every species of the section has at least one attribute that is unique for Aneilemu, e.g., stamen filaments bearded only in the basal half (A. gillettii); stamen filaments with branched hairs (A. nyasense); pollen dimorphic (A. hockii); capsules strongly compressed laterally with only the single dorsal locule seed developing (A. plagiocapsa). Many other characters present in section Amelina are uncommon in the genus as a whole, e.g., yellow petals, pistillate flowers, scented flowers, basally fused stamen filaments, hook-hairs on the stamen filaments, and isodiametric capsule wall cells. Section Amelina is the most widespread section within Africa. This is due in large measure to the broad ranges of two species, A. aequinoctiale in forests throughout tropical Africa, and A. hockii in non-forest habitats in eastern and southern Africa. Section 3: Rendlei Faden, new section Staminodium medium erectum. Staminodia lateralia versus apicem valde recurvata. Stamina lateralia florum staminatorum filamentis prope basin valde deflexis, saepe formis litterae J, approximatis, parallelis, glabris. Stamen medium connectivo antherae valde convex0 maculis sanguineis ornato. Ovarium glandulifero-microtrichomatibus sparsis, appressis praeditum, aliter glabrum. Stylus forma litterae J. Capsulae glabrae. TYPE SPECIES.-Aneilemu rendlei C.B. Clarke. Perennial or rarely annual herbs with fibrous or tuberous roots. Leaves spirally arranged. Inflorescences terminal (and occasionally axillary) thyrses with predominantly alternate cincinni. Inflorescence axis and cincinnus peduncles and axes glabrous or puberulous, uniseri- ate hairs absent. Bracteoles herbaceous, cup-shaped, perfoliate, prominently glandular near the apex, margin thickened (glandular?) towards the fused edge. NUMBER 76 63 Rowers perfect and staminate. Fruiting pedicels usually erect, occasionally spreading laterally or decurved. Sepals prominently glandular, the gland of the medial sepal incon- spicuously bilobed, those of the lateral sepals usually unlobed. Petals glabrous, the medial reduced, inconspicuous. Filament bases free. Medial staminode short, erect, antherode bilobed with lobes stipitate, generally reniform, often spotted with maroon adaxially, always larger than the lobes of the lateral staminodes, connective elongate. Lateral staminodes with filaments thickened basally, strongly deflexed, then strongly recurved towards the apex, antherodes bilobed with stipitate lobes. Lateral stamens dimorphic in perfect and staminate flowers, filaments J-, U-, or S-shaped, parallel in the staminate flowers, glabrous, anthers lanceolate to elliptic, pollen different in color from that of the medial stamen. Medial stamen with anther elliptic to ovate, connective very convex and maroon- spotted. Ovary sessile, oblong-elliptic, glabrous except for sparse, appressed, glandular microhairs mainly or exclusively on the lateral sutures and ventral surface, dorsal locule usually not developed, occasionally with an ovule, ventral locules each 2-6-ovulate; style J-shaped, held in the floral midplane, sharply deflexed at the base, stigma small. Capsules sessile, oblong to oblong-elliptic, dehiscent, bi- valved, usually bilocular, lustrous, glabrous, valves persistent, dorsal locule (when present) empty or 1-seeded, ventral locules each 2-5-seeded, cells of the wall transversely elongate, in regular files. Seeds trapezoidal, the apical rounded apically, the basal angular basally, hilum raised within a groove, testa scrobiculate, white-farinose in all the depressions and around the embryotega and hilum. BASIC CHROMOSOME NUMBER.-% = 16. DISTRIBUTION.CW Ethiopia S to NE Tanzania (Map 4). - D 100 .oo KILDUEiERB 0 10 1 1 h l A ~ 4.-Distribution of Aneilema section Rendlei. new section. SMITHSONIAN CONTRIBUTIONS TO BOTANY Key to the Species of Section Rendlei 1. Plants not rhizomatous; roots fibrous; leaves petiolate; inflorescence axis glabrous; flowers lilac to lavender. 2. Shoots decumbent; fruiting pedicels to 8 mm long; capsules 3-6 mm long: seeds 1.3-1.6 mm wide. 3. Cincinni 1-13 per inflorescence; leaves to 10 cm long; capsules stramineous, locules 2-3-seeded . . . . . . . . . . . . . . . . . . . . . . . 1. A. taylorii 3. Cincinni 14-20 per inflorescence; leaves to 16 cm long; capsules chocolate brown, locules 4-seeded . . . . . . . . . . 2. A. usambarense, new species 2. Shoots mostly erect or ascending; fruiting pedicels to 6.5 mm long; capsules 6-10 mm long; seeds 1.5-2 mm wide . . . . . . . . . . . . . . . . . 3. A. rendlei 1. Plants rhizomatous; roots tuberous; leaves sessile; inflorescence axis puberulous; flowers white or nearly so . . . . . . . . . . . 4. A. brenanianum, new species DISCUSSION Section Rendlei is very natural and homogeneous. The most distinctive features are the moderately to strongly dimorphic lateral stamen filaments in perfect and staminate flowers; strongly convex, maroon-spotted medial anther connective; and lack of hairs (other than inconspicuous glandular mi- crohairs) on the ovaries. The only taxonomic problems are the possible division of A. taylorii into infraspecific taxa and the final determination of the status of the plant described below as A. usambarense. Resolution of these problems would require obtaining living material of these plants. 1. Aneilema taylorii C.B. Clarke Aneilema taylorii C.B. Clarke [?Taylori?] in Thiselton-Dyer, Fl. Trop. Africa, 8:79, 1901 [type: Kenya (East Africa on label), Rabai Hills, Mornbasa (Mornbaz on label), Fimbine? (probably = Fumbini on Kilifi Creek), Jul-Nov 1885, W. E. Taylor s.n. (BM)]. Aneilema rendlei Brenan, Kew Bull., 7:196. 1952, p.p. [non C.B. Clarke (1 90 1 )]. Aneilema sp. nov. aff. A. umbrosum (Vahl) Kunth, Morton, T. Linn. Soc., Bot., 59:464, 1966 [in adnot]. Perennial or annual herbs (habit type IA, Figure 1). Roots fibrous. Shoots decumbent, much branched, flowering shoots ascending, 15-30(-60) cm tall. Internodes 2-8(-9.5) cm long, puberulous, occasionally sparsely pilose as well. Leaves spirally arranged, sheaths 0.5-1.3 cm long, puberulous and frequently sparsely pilose, ciliate at the apex, laminae petiolate, lanceolate to lanceolate-elliptic or ovate, 2- 10 cm long, 0.7-3(-4.3) cm wide, apex acuminate to acute, base cuneate, margin sometimes undulate, scabrid, adaxial surface patently hirsute-puberulous, uniseriate hairs 1-2.3 mm long, abaxial surface patently pilose-puberulous, uniseriate hairs very vari- able in number, sometimes confined to the veins, always shorter than those of the adaxial surface; petioles to 1.X-2) cm long, ciliate on the margins. Inflorescences thyrses (Figure 449, terminal and occasion- ally axillary from the uppermost leaves, lax to moderately lax, ovoid, 1.5-5.5(-7) cm long, 1.5-6(-12) cm wide, with (1-)2-lo(-13) cincinni, alternate to subopposite, patent to ascending (rarely the lower somewhat declinate). Peduncles (1.5-)2-53-73 cm long, green, puberulous, often glabrous apically. Inflorescence bract medial, herbaceous, or occasion- ally membranous, generally very reduced, occasionally foli- aceous. Inflorescence axis glabrous, slightly zigzag. Cincinni to 5(-6.5) cm long and 17(-18)-flowered. Cincinnus bracts herbaceous to membranous, lowermost sometimes perfoliate, appressed to the cincinnus peduncles, lanceolate or lanceolate- elliptic to oblong or ovate, 1.5-3 mm long, glandular near the cucullate apex, glabrous. Cincinnus peduncles relatively uniform in length within the inflorescence, exceeding the cincinnus bracts, (2.5-)3.5-8 mm long, glabrous. Cincinnus axes glabrous. Bracteoles attached (1.2-)2-3.5(-4.5) mm apart, herbaceous, eccentrically cup-shaped, perfoliate, 1- 1.5 mm long, 0.4-0.8 mm high, with a prominent gland near the apex, glabrous, margin slightly thickened (glandular?) near the fused edge. Flowers perfect and staminate (Plate lm,n), odorless, (7-)9-15 mm wide. Pedicels 3.5-5.5(-6.5) mm long in flower, to 8 mm long in fruit, straight or gently arcuate in flower, erect, spreading laterally or decurved in fruit (some- times all curvature types within the same inflorescence), persistent, glabrous. Sepals with a subapical, moderatcly prominent gland, glabrous; medial sepal elliptic to lanceolate- elliptic or lanceolate-ovate, 2.5-3.7 mm long, 1.8-2.4 mm wide, gland unlobed or bilobed; lateral sepals elliptic to oblong-elliptic or ovate-elliptic, 2-3.4 mm long, 1.6-2 mm wide. Paired petals (5.2-)5.5-9.5 mm long, (3.7-)4-7 mm wide, limb ovate to ovate-elliptic or ovate-suborbicular, 4-6.5 mm long, lilac to lavender (RHS colors: 87D, Magogo in EA14203, Faden & Faden 74t371 and 77t611; 76B-C, Faden & Faden 74/371 and 77t611), apex rounded, claw 1.5-3.5 mm long, concolorous with limb, glabrous. Medial petal reflexed, ovate to ovate-elliptic, 3-5 mm long, 2.3-3.7 mm wide, greenish white, sometimes tinged with lilac, margins colorless. All filament bases free, Medial staminode filament 1-2 mm long, reddish pink or reddish purple, antherode bilobed, lobcs stipitate, flat, reniform or sickle-shaped, (0.3-)0.6- 1 mm long, NUMBER 76 65 i I I I h I d +-&& FIGURE 44.-Comparism of Aneilem rendlei, A. tuylorii, and A . rendlei x tuylorii: inflorescences, leaves, capsules, and seeds. A. rendlei C.B. Clarke (A): 4,inflorescence; b, leaf; c, capsule; d , seed (from Kubuye & Evum163). A. rendlei C.B. Clarke x tuylorii C.B. Clarke (4): e, inflorescence;$ leaf; g , capsule; h, seed (from Kubuye & Evum 163 (rendlei) x Faden & Fuden 741371 (tuylorii3). A. tuylorii C.B. Clarke (i-l): i, inflorescence; j , leaf; k, capsule; I , seed (from Fuden & Fuden 741371). (Bar = 10 mm for u,b,e,$i,j; 2 mm for c,g,k; 0.5 mm for d M . ) 66 SMITHSO? CONTRIBUTIONS TO BOTANY 1.1 - 1.7 mm wide, yellow with small maroon or brown spots on the adaxial surface (these rarely absent), connective elongate. Lateral staminodes with filaments 3-6 mm long, arcuate- decurved then sharply recurved into a vertical position (Plate 5n), tapering from a thick base, reddish pink or purple, shading to yellow near the apex, antherodes bilobed, lobes stipitate, oblong, elliptic, elliptic-subreniform or C-shaped, (0.25-p.4- 0.6 mm long, yellow, sometimes with small maroon or brown spots on the adaxial surface, connective not at all or scarcely elongate. Lateral stamens moderately dimorphic (or sometimes not dimorphic?) in perfect and staminate flowers, shorter and more divergent in the former than in the latter (Figure Se,f,k,I); filaments 4-10 mm long, decurved near the base, J- or S-shaped (Plate 5n), glabrous, anthers lanceolate to lanceolate- elliptic or elliptic, 0.9-1.4 mm long, 0.5-0.8 mm wide, connective whitish, creamy yellow or gray-green, pollen yellow or orange-yellow. Medial stamen with filament 3-6 mm long, anther broadly elliptic, 1-1.7 mm long, 0.7-0.9 mm wide (Figure 31ef). usually held perpendicular to the ground, connective very convex, creamy white blotched with maroon, sometimes mostly maroon, pollen yellow or white or a mixture of yellow and white grains. Ovary not at all or scarcely stipitate, oblong-elliptic to ovate-oblong, 1-1.8 mm long, 0.8-1.1 mm wide, glabrous except for a few inconspicuous, appressed, glandular microhairs on the lateral sutures and sometimes also on the ventral surface (Figure 426 ), apex truncate or rounded, dorsal locule represented by a low, middorsal ridge, empty, ventral locules each 2-3-ovulate; style 3.5-8.5 mm long, J-shaped (Plate 5n), sharply deflexed at base, held in floral midplane, k uniform in thickness for most of its length, tapering near the apex, stigma small, whitish or yellowish, held below the stamens. Capsules (Figure 44k) sessile, elliptic to broadly elliptic or oblong-elliptic, dehiscent, bivalved, bilocular, (2.8-)3-4.5 (-6) mm long, 2-3.2 mm wide, stramineous, lustrous, glabrous, apex emarginate to truncate, valves persistent, locules 2-3-seeded, cells of the capsule wall transversely elongate. Seeds (Figure 441; Plate 5a) trapezoidal, apical rounded apically, basal angular basally, 1.2-2 mm long, 1.3-1.6 mm wide, hilum dark brown, prominently raised within a groove, not at all to only slightly extended onto basal or apical surface, testa pale grayish brown, scrobiculate on all surfaces, white- farinose in all depressions, around the embryotega and in the groove around the hilum, the farinose granules rarely confined to the vicinity of the hilum and embryotega (Verdcourt 1921 and Faden & Faden 771611). HABITAT.-bwland and submontane forests, sometimes associated with limestone outcrops, frequently on boulders along streams and at the bases of waterfalls in the Eastern Usambara Mountains, described as ?the first invader in a cleared area in evergreen rain forest,? on Greenway 1668; partial or dense shade; 30-900 m. FLowEmNcx-Flowering specimens have been collected in March, April, June, August-October, and December. Based on two populations in cultivation, the flowers appear to open before to just slightly after dawn. They were observed to fade in the field about 1200 hrs. CHROMOSOME NUMBER.-n = 16,2n = 32. D I S T R I B U T I O N . ~ O ~ S ~ ~ ~ SE Kenya, coastal and subcoastal NE Tanzania (Map 5). SPECIMFNS EEN.- KENYA. KILIFI: Rabai Hills, Mombaz. (Mombasa), Fimbine? (probably = Fumbini on Kilifi Creek), Jul-Nov 1885, WE. Taylor s.n. (BM); Kaloleni district (i.e., area; erroneously recorded as Kwale District), Verdcourt 2410 (EA, FT, K). KWALE: Muhaka Forest, 4?20?S, 39?312, Faden & Faden 77/611 (EA, F, K, MO, US); Marenji (Marenje) Forest Reserve, Verdcourt 1921 (BR, EA, K, PRE). TANZANIA. Without data: Warnecke 323 (EA). LUSHOTO: Amani, Kwam Kujv (Kwamkuyu), Engler 3416 (EA); Eastern Usambara Mts., Muheza-Amani road, crossing of Kwamkuyu River, 5?06?S,38?39%, Faden & Faden 741371 (C, EA, K, MO, US, WAG); Eastern Usambara Mts., Amani, Greenway 1668 (EA, K, PRE); Eastern Usambara Mts., Sigi, near Amani, cultivated in Nairobi, Magogo in EA14203 (BR, EA, K, MO); Eastern Usambara Mts., Kwamkuyu Falls, Derema, Verdcourt 161 @A, K , MO, PRE); Amani, Winkler 3638 (WRSL-2 sheets); same locality, Zimmermann 8231 @A). TANGA: Magila near Muheza, Archbold 1408 (EA); Kiomoni Quarries near Tanga, cultivated in Nairobi, Organ in EA15079 (EA). DISCUSSION I could not read the locality starting with ?F? on the type specimen. Clarke (1901) interpreted the Rev. Taylor?s hand- writing as ?Fimbine.? S.A. Robertson (personal communica- tion) has stated that it is probably Fumbini on Kilifi Creek. This species has sometimes been confused with others. Brenan (1952) cited two specimens of A. taylorii, Greenway 1668, and Verdcourt 161 (cited as 151), under Aneilemu rendlei, noting that they had smaller capsules with fewer seeds than other specimens of A. rendlei. Morton (1966) identified another collection of A. taylorii, Verdcourt 2410, which had been determined in the Kew Herbarium as A. umbrosum var. ovato-oblongum, as an undescribed species closely related to A. umbrosum. Although pressed specimens of A. taylorii bear a striking resemblance to some West African plants of A. umbrosum, this similarity is superficial. The two species differ in at least 20 floral characters, including lateral stamen filament pubescence and medial stamen anther size, shape, and color. Herbarium specimens of A. umbrosum are readily separated by the inflorescence axis and pedicel apex puberulous and the seeds reticulate-foveate. Aneilema taylorii exhibits a great deal of morphological variation, particularly in size of floral parts. I previously suggested that different-size flowers might be taxonomically significant in this species, with smaller flowers occurring in gracile, narrow-leaved, coastal plants, and larger flowers in more robust, broader-leaved plants from the Eastern Usambara NUMBER 76 67 I I I . 0 0 4?i/ I I I I I I I I I I I I no 0 . . 7 0 . \ 0 L 0 :, \ o \ \ 0 100 MILES 100 KILOMETERS 36 38 I I MAP 5.-Distribution of Aneilema brenuniunwn Faden, new species (open squares), A. rendlei C.B. Clarke (solid circles). A . luylorii C.B. Clarke (open circles), and A. usomburerne Faden. new species (solid square). 68 SMITHSONIAN CONTRIBUTIONS TO BOTANY i b I FIGURE 45.4ompanson of Aneilem rendlei, A. taylorii, and A. rendlei x taylorii: lateral stamens and gynoecium. A. rendlei C.B. Clarke (a,b,g,h): a, perfect flower, front view; b, perfect flower, side view; g, staminate flower, front view; h, staminate flower, side view (from Kabuye & Evans 163). A. rendlei C.B. Clarke x taylorii C.B. Clarke (c,d,i,j): c, perfect flower, front view; d , perfect flower, side view; i , staminate flower, front view; j 8 staminate flower, side view (from Kabuye & Evans 163 (rendlei) x Faden & Faden 74/371 (taylorii]). A . taylorii C.B. Clarke (e,Jk,l): e , perfect flower, front view;f, perfect flower, side view; k, staminate flower, front view; I , staminate flower, side view (from Faden & Faden 74/37]). (Bar = 2 mm for all illustrations.) Mountains (Faden, 1975). A new collection from the Kenya coast, Faden & Faden 77/611, has tended to support this hypothesis. First, by being annual-all of the plants found were completely dead, and plants subsequently grown from seeds collected from the dead plants behaved as annuals-this collection agreed with the presumption that all of the Kenya collections, as well as Organ in EA15079 from Tanzania, were annuals, in contrast to plants from the Usambaras. Second, flower size and floral parts? dimensions in the cultivated plants of Faden & Faden 77/611 much better agreed with Organ in EA15079 than with living plants of two Usambara collections, Magogo in EA14203 and Paden & Faden 74/371. While the NUMBER 76 69 new data gained from Faden & Faden 77/61 1 would support the recognition of two subspecies in A. tuylorii, hard and fast characters usable in herbarium specimens have remained elusive. Until such time as some are found or the weight of the floral evidence becomes overwhelming, there is no compelling reason to recognize subspecies that can only be identified by the provenance of the collection, when flowers are not available. The variation in color of the pollen grains of the middle anther is a phenological phenomenon. When the flowers open all of these pollen grains are yellow. During the course of flowering they change to white, and so, when the flowers begin to fade, only white pollen grains can be found in the middle anther. No such color change occurs in the pollen of the lateral anthers. The gaps in the monthly flowering records mostly reflect the small number of collections of this species. On the other hand, the lack of flowering specimens from January and February, the driest months through out the range of A. tuylorii, may well indicate that the species is not in flower during that period. All of the Tanzanian collections of A. tuylorii from Lushoto District are from a small area of the Eastern Usambara Mountains centered around Amani, the former German colonial agricultural research station. The species is apparently fairly common in this region between 500-900 m altitude. It is decidedly uncommon throughout the rest of its range. Aneilemu tuylorii is undoubtedly most closely related to A. usumburense, which may ultimately prove to be conspecific. For distinctions see that species. It is also closely related to A. rendlei, and crossing experiments have shown the two species to be interfertile, at least through the F1. Some of the differences between these species are given in the key and under A. rendlei (see Figures 44,45). To these distinctions the following characters of A. tuylorii can be added: leaf sheaths shorter; inflorescence axis, cincinnus peduncles and axes, pedicels and medial sepal consistently green: lateral staminode filaments less strongly curved, with bases more slender; lateral anthers always facing the floral midplane. Moreover, A. tuylorii and A. rendlei are completely isolated ecologically as well as geographically (Map 5) . 2. Aneilema usambarense Faden, new species Aneilema rendlei C.B. Clarke.-Brenan, Kew Bull., 7:196, 1952, p.p. Herbae perennes decumbentes caulibus ad 90 cm altas, foliis lanceoato-ellipticis, 6.5-16 cm longis, 2-4.5 cm latis, petiolis ad 3.5 cm longos. Inflorescentiae thyrsi ovoidei, glabri, 4-7 cm longi, 4.5-6 cm lati, cincinnis 14-20 compositi. Cincinni pedunculis 5-10 mm longis, bracteolis (2-)3-4(-5) mm distantibus. Pedicelli 6-8 mm longi, glabri; sepala glabra; stamina filamentis glabris. Capsulae 5-6 mm longae, 2.5 mm latae, chocolatinae, glabrae, loculis ventralibus uterque semini- bus 4 praeditis. Semina 1.1-1.7 mm longa, 1.5-1.6 mm lata, testis scrobiculatis (Figure 46). TYPE.-Tanzania, Eastern Usambara Mts., W slope of Mt. Mlinga, 4 Dec 1940, Creenway 6058 ( K , holotype; EA, PRE, isot ypes) . FIGURE 46.-Aneilema usumburense Faden, new species: u, leaf; b, inflorescence; c, capsule; d, seed (all from Greenway 6058). (Bar = 10 mm for u,b; 2 mm for c; 0.5 mm for d. ) 7 0 SMITHSONIAN CONTRIBUTIONS TO BOTANY Perennial herbs (habit type IA, Figure 1 ) . Roots not seen, probably thin, fibrous. Shoots decumbent, the flowering shoots ascending, to 90 cm tall. Internodes 3.5-10 cm long, puberulous to glabrescent. Leaves spirally arranged, sheaths 0.7-2 cm long, puberulous, ciliate at the apex, laminae petiolate, lanceolate-elliptic, 6.5-16 cm long, 2-4.5 cm wide, apex acuminate, base cuneate, margin planar, scabrid, adaxial surface patently hirsute-puberulous, the scattered uniseriate hairs 1.5-2.5 mm long, abaxial surface pilose-puberulous, the hairs more numerous and shorter than those of the adaxial surface and f confined to the lateral veins, petioles to 3.5 cm long, ciliate on the margins. Inflorescences thyrses (Figure 46b), terminal (rarely also axillary from the inflorescence bract), moderately dense, ovoid, 4-7 cm long, 4.5-6 cm wide, with 14-20 ascending cincinni. Peduncles 1.5-2.8 cm long, puberulous. Inflorescence bract medial, herbaceous, very reduced. Inflorescence axis glabrous. Cincinni to 4 cm long and 9-flowered. Cincinnus bracts herbaceous, lowermost sometimes perfoliate, appressed to the cincinnus peduncles, lanceolate-elliptic to ovate, 1-3.5 mm long, glandular near the apex, glabrous. Cincinnus peduncles relatively uniform in length within the inflorescence, exceeding the cincinnus bracts, 5-10 mm long, glabrous. Cincinnus axes glabrous. Bracteoles attached (2-)3-4(-5) mm apart, herba- ceous, eccentrically cup-shaped, perfoliate, 1.2-1.5 mm long, 0.4-0.7 mm high, with a prominent subapical gland, glabrous, margin somewhat thickened (glandular?) near the fused edge. Flowers perfect and staminate. Pedicels 6-8 mm long, aligned with the cincinnus axis in flower, erect in fruit, glabrous. Sepals glandular near the apex, glabrous; medial sepal -3 mm long, lateral sepals oblong-elliptic, -3 mm long and 1.5 mm wide. Paired petals -7 mm long, mauve. Medial petal ovate, 4-5 mm long. All filament bases free. Medial staminode filament -1.5 mm long, antherode bilobed, lobes reniform, -1-1.5 mm wide, connective elongate. Lateral staminodes with filaments question-mark-shaped, -5 mm long, antherodes bilobed, lobes stipitate, -0.7 mm wide. Lateral stamens with filaments declinate, -7-8 mm long, glabrous, anther lanceolate-elliptic, 1.5 mm long, 0.7 mm wide. Medial stamen with filament straight, then gently recurved, -5 mm long, anther 1.8 mm long, with a broad connective. Ovary sessile, -2 mm long, apparently glabrous, style J-shaped, 8.5 mm long, stigma small. Capsules sessile (Figure 46c). oblong-elliptic, dehiscent, bivalved, bilocular, 5-6 mm long, 2.5-3 mm wide, chocolate broken, lustrous, glabrous, apex emarginate, valves persistent, dorsal locule obsolete, ventral locules 4-seeded, cells of the capsule wall transversely elongate. Seeds trapezoidal (Figure 46d) , apical rounded apically, basal angular basally, 1.1-1.7 mm long, 1.5-1.6 mm wide, hilum black, prominently raised within a groove, not extended onto apical and basal surfaces, testa pale grayish brown, scrobiculate on all surfaces, white-farinose in all depressions, around the embryotega and in the groove around the hilum. HABITAT.-Shady path sides in evergreen forest with Parimri excelsa, Funtumia latifolia, Sterculia appendiculata, Allanblackia stuhlmannii, Isoberlinia, Scheflera, growing with Sacciolepis curvata; 620 m. FLOWERING.-The single collection is from December and is flowering and fruiting. The sizes of the inflorescences indicate that the plant was in flower in November and perhaps earlier. CHROMOSOME NUMBER.-Unknown. DISTRIBUTION.-Known only from the isolated massif Mt. Mlinga in the Eastern Usambara Mts. in NE Tanzania (Map 5). SPECIMENS EEN.-TMZANIA. TANGA: Eastern Usam- bara Mts., W slope of Mt. Mlinga, Greenway 6058 @A, K, PRE). DISCUSSION The single collection of A. usambarense was treated without comment as A. rendlei by Brenan (1952). This is hardly surprising in view of his inclusion of two collections of A. taylorii under A. rendlei. Brenan?s confusion of these three taxa is understandable considering the small number of collections available to him and the somewhat intermediate appearance of Greenway 6058 between A. taylorii and A. rendlei. Two relatively well-preserved flowers on the PRE sheet of Greenway 6058, one perfect, the other staminate, suggest that dimorphism in lateral stamen filaments is present and is no greater than that in A. taylorii. Aneilema usambarense is most closely related to A. taylorii, but differs in a number of characters. Most importantly, A. usambarense has taller flowering shoots, longer leaves with longer petioles, more numerous cincinni in the inflorescence, and darker brown capsules with more numerous seeds per locule. It also has somewhat longer cincinnus peduncles and more widely spaced bracteoles than A. taylorii. Possibly other distinguishing characters are present in the flowers, but they could not be determined from the dried specimens. The generally larger size in all of these characters in A. usambarense than in A. taylorii suggests that Greenway 6058 might be merely a specimen of the latter from a very favorable habitat or perhaps an ecotype of that species. Because plants of A. taylorii (Faden & Faden 741371) have been cultivated under favorable conditions and have failed to significantly reduce the morphological differences between the two species, ecological factors effecting greater size seem insufficient to account for the observed differences between Greenway 6058 and A. taylorii. Genetically based, ecotypic variation cannot be ruled out, but the plant association described on Greenway 6058 (see habitat above) is the dominant one in the Amani area of the Eastern Usambara Mountains where A. taylorii is frequent. There is little reason to expect climatic differences in these two areas, only about 5 km apart, to be great enough to account for the observed morphological differences. Greenway 6058 might also be considered a possible NUMBER 76 71 autopolyploid of A. taylorii or a hybrid between it and A. rendlei. Studies of the morphological differences in other supposed autopolyploids in Aneilema have not been suffi- ciently detailed to provide a basis for comparison. In the absence of living material or any clearly supportive data of a polyploid origin, there is no compelling reason for treating the Greenway specimen as having had one. Aneilema usamburense, from a casual inspection, appears intermediate between A. taylorii and A. rendlei. It is somewhat intermediate only in the number of cincinni per inflorescence, capsule size and number of seeds per locule. In most other characters, including habit, leaf shape, cincinnus peduncle and pedicel length, seed width, distance between bracteoles and degree of lateral stamen dimorphism, A. usambarense is much more similar to A. taylorii than to A. rendlei. Its distribution and habitat also suggest a closer relationship with A. taylorii; a hybrid origin is unlikely. Artificial hybrids (Figures 44,45) made between cultivated plants of A. taylorii (Faden & Faden 741371) and A. rendlei (Kabuye & Evans 163) differed from Greenway 6058 in having broader, stramineous capsules (3.2-3.9 mm wide) and broader seeds (1.65-1.75 mm wide), which further supports the conclusion that A. usambarense is not a hybrid between those two species. Greenway 6058 is best treated as a closely related and probably recent derivative of A. taylorii. It could be considered either a subspecies of A. raylorii or a closely related species. The latter course has been followed largely for convenience. It was felt that inclusion of Greenway 6058 in A. taylorii would have resulted in a blurring of the distinctions between that species and A. rendlei. The peculiarly colored capsules of Greenway 6058 separate it from both of these species and support its separation at the specific level. Final determination of the proper status for A. usumburense must await the availability of further collections, particularly living material, so that the floral structure and variability can be studied. An attempt by the author to collect this plant in March 1974 was unsuccessful. Unfortunately, the habitat was being destroyed rapidly. 3. Aneilema rendlei C.B. Clarke Aneilema rendlei C.B. Clarke in Thiselton-Dyer. H. Trop. Africa, 8:68, 1901 [Domldron Smith 346 excl.].4hiovenda, Webbia, 8:38. 1951, p.p. maj.-Brenan, Kew Bull., 7: 196. 1952. p.p.4ufodontis. Bull. Jard. Bot. Natl. Belgique, 41, suppl., 1518,1971 [DoMIdson Smith 346 excl.1.-Faden in Agnew, Upland Kenya Wild H., 666,1974 [type: Ethiopia (Somaliland on label), 23 Apr 1895. DoMkfson Smilh s.n. (BM)]. Aneilema octospermum C.B. Clarke, in sched. ~octosperma?].-Brenan, Kew Bull., 7:196, 1952 [nom. nud. pro syn.]. Perennial herbs (habit types IB, IC, Figure 1). Roots fibrous, generally produced only at the lower nodes. Shoots erect to ascending or occasionally decumbent, generally 25-100 cm high, much branched at the base, unbranched or sparsely branched above, shoot bases swollen. Internodes (2.5-)4-6 (-9) cm long, glabrescent to puberulous. Leaves spirally arranged, sheaths 1-2.5(-3) cm long, puberulous, pilose or pilose-puberulous, ciliate at the apex, laminae petiolate, narrowly lanceolate to lanceolate-elliptic, elliptic or occasion- ally ovate-elliptic, 2.5-11 cm long, 1-3.5 cm wide, apex acuminate or occasionally acute, base cuneate, margin undulate or planar, scabrid and sometimes ciliate, both surfaces dull, the adaxial patently hirsute-puberulous, with uniseriate hairs to 1.5-3 mm long, abaxial patently pilose-pubemlous, with uniseriate hairs shorter and more numerous than those of the adaxial surface, petioles to 1.5(-1.8) mm long, with ciliate margins. Inflorescences thyrses (Figure 44a; Plate 2d) , terminal and occasionally axillary from the uppermost leaves, moderately dense, ovoid, (2-)3-5(-5.5) cm long, (1.5-)2-3(-6) cm wide, with (9-)13-20(-32) cincinni, alternate to subopposite, ascending. Peduncles (1.5-)4-6(-10,5) cm long, puberulous basally, glabrous apically. Inflorescence bract medial, supra- medial or occasionally subapical, herbaceous or less commonly membranous, bract-like or occasionally foliaceous. Inflores- cence axis glabrous. Cincinni to 3.5 cm long and 13-flowered. Cincinnus bracts herbaceous, lowermost sometimes perfoliate, appressed to the cincinnus peduncles or rarely reflexed with age, lanceolate or lanceolate-elliptic to oblong or ovate, 1.5-3(-5) mm long, cucullate and glandular subapically, glabrous. Cincinnus peduncles relatively uniform in length within the inflorescence, exceeding the cincinnus bracts, (3-)4-6(-8.5) mm long, glabrous. Cincinnus axes glabrous. Bracteoles attached (1-)2-23-3) mm apart, herbaceous, eccentrically cup-shaped, perfoliate (rarely not perfoliate), 1.2-2.2 mm long, 0.4-1.2 mm high, with a prominent, subapical gland, glabrous, margin sometimes apparently glandularly thickened near the fused edge. Flowers perfect and staminate, slightly fragrant, 9.5-17 mm wide (Figure 17; Plate log). Pedicels 2.5-4(-5.5) mm long, to 6.5 mm long in fruit, generally held above the cincinnus axis in flower, erect in fruit, persistent, glabrous. Sepals with a prominent subapical, green, inconspicuously bilobed gland, glabrous; medial sepal lanceolate-ovate to ovate-elliptic, 3-4.5 mm long, 2-2.6 mm wide; lateral sepals elliptic to oblong- elliptic, 3-4.5 mm long, 1.7-2.2 mm wide. Paired petals 7.5-1 1 mm long, 4.5-8 mm wide, limb ovate to ovate-elliptic or suborbicular, 5.5-8 mm long, lilac (RHS colors: 76B, Kabuye & Evans 163, cultivated, and Faden et al. 691322, cultivated; 76A-B, Faden & Faden 741232 and Faden et al. 701157, cultivated; 76C, Faden & Faden 721240, cultivated; 75A, 76C. 84C, Faden & Faden 741281; 76A, 82C-D, Faden & Faden 741503), veins contrastingly darker than background color, apex rounded, claw 2.5-3 mm long, colorous with limb at apex, shading to greenish yellow at base, glabrous. Medial petal not strongly reflexed, ovate to ovate-elliptic or lanceolate- ovate, 4.5-5.5 mm long, 3-3.5 mm wide, greenish white, sometimes flushed with lilac. All filament bases free. Medial 72 SMITHSOMAN CONTRIBUTIONS TO BOTANY staminode filament 1.2-2 mm long, yellow, antherode bilobed (Figure 43c), lobes stipitate, flat, reniform, 1-1.3 mm long, 1.8-2.5 mm wide, yellow abaxially, yellow with small maroon spots (rarely lacking) adaxially, connective elongate. Lateral staminodes (Plate l p ) 4.5-5.5 mm long, question-mark- shaped, very thick at the base, tapering apically, entirely yellow or, greenish yellow at base, flesh pink or pinkish buff in middle, yellow at apex, antherodes bilobed (Figure 43c), lobes stipitate, reniform, 0.5-0.8 mm long, entirely yellow or sometimes spotted with maroon on adaxial surface, connective not elongate. Lateral stamens strongly dimorphic in perfect and staminate flowers (Figures 16b, 17,45), filaments glabrous, in perfect flowers parallel or slightly divergent, 5.5-8 mm long, broadly U-shaped, in staminate flowers closely parallel, 9-13 mm long, J-shaped, anthers narrowly elliptic to lanceolate- elliptic, in staminate and some perfect flowers with anther sacs facing the filament bases, in some perfect flowers facing the floral midplane, 1.5-1.7 mm long, 0.6-0.8 mm wide, connective creamy white to gray-green, pollen orange-yellow. Medial stamen with filament 3.5-5 mm long, anther elliptic, 1.7-2 mm long, 1-1.4 mm wide, usually held perpendicular to the ground, sometimes pendulous, connective very convex, creamy white, greenish white or greenish yellow, always with numerous maroon spots, pollen yellow. Ovary sessile, oblong- elliptic, 2-2.5 mm long, 0.9-1.1 mm wide, glabrous except for some very inconspicuous, appressed, glandular microhairs along the lateral sutures and sometimes on the ventral surface (Figure 42u), apex rounded, dorsal locule usually represented by a low, middorsal hump or ridge, usually empty, occasionally l(-3)-ovulate, ventral locules each 3-6-ovulate; style 7.5-10 mm long, J-shaped, sharply deflexed at base, thickest below middle, tapering towards base and apex (Figure 17u; Plate l p ) , held in floral midplane, stigma small, bluish purple, held below the stamens but at some distance from them. Capsules sessile (Figure 444, oblong to oblong-elliptic, dehiscent, bivalved, 2(-3)-locular, (5-)6- 10 mm long, (3.1-)3.5-4.2 mm wide, stramineous, lustrous, glabrous, apex rounded to truncate or emarginate, valves persistent, dorsal locule obsolete or, if present, usually empty, rarely 1-seeded, ventral locules each 3-5-seeded, rarely less by abortion, cells of the capsule wall transversely elongate. Seeds (Figure 44d; Plate 5f) trapezoidal, apical ventral rounded apically, basal ventral angular basally, 1.5-2.2 mm long, 1.5-2 mm wide, hilum blackish brown, prominently raised within a groove, not at all to only slightly extended onto apical or basal surfaces, testa grayish brown to flesh-pink, scrobiculate on ventral and dorsal surfaces, white-farinose in all depressions, around the embryotega and in the groove around the hilum. HABITAT.-Dry deciduous bushland and bushland thicket, often on rocky slopes, occasionally along seasonal streams; usually in sandy soil; generally in partial shade; 300-1200 m. FLOWERING.-Flowering specimens have been seen from all months except August and September. In Kenya and Tanzania there are two chief flowering periods, December-February (following the short rains) and April-June (during and after the long rains). In the field, flowers were observed to open at about 0800 hrs. They probably close 1330-1430 hrs. CHROMOSOME NUMBER.-n = 1 6 . h = 32. DISTRIBUTION.-SW Ethiopia N and SE Kenya, NE Tanza- nia (Map 5). SPECIMENS EEN.-ETHIOPIA. GEMU-GOFA: Rive del Ghizo, Corrudi 2159, p,p. (FT) and 2160 F). SIDAMO: 23 Apr 1895, Donaldson Smith s.n. (BM). KENYA. KWALE: 2.8 km Samburu-Mackinnon Road, 3'47'S, 39'152, Fuden & Evans 701788 (EA, K); same locality, Fuden & Fuden 721240 (EA, MO); Taru, Scott Elliot 6179 (K). LAIKIPIA: Colcheccio Ranch, NE comer, between Oo40'N, 36'54'E and Oo44'N, 36'572, A. Faden 27185 ( U S ) . LAIKIPIA~ORTHERN FRONTIER: Near meeting point with Samburu District, Nolaleta River, -Oo44'N, 36'572, A. Fuden 16185 ( U S ) . MACHAKOS: Kibwezi, Dummer 5020 (K); Bush- wackers, planted in Nairobi, Gilbert s.n. (EA); Machakos, Katumani Experimental Farm, Kaiti, Thomas 1061b (EA); Kibwezi, Tweedie 4036 (K). NORTHERN FRONTIER: Mathews Range, Ngeng, Newbould 3184 (K). TEITA: Voi, Mazinga (Beehive) Hill, Bully 88534 (EA, K); Mt. Kasigau, lower slopes, Bully B12692 (G, K) and Bully 813592 (EA); Maktau, 26 Oct 1915, Buchanun s.n. (BM); Taveta-Voi road, at mile post Taveta 36Noi 36, -395's. 38'102, Fuden et al. 691322 (DSM, EA, K, MO, US); 2-3 mi Maungu Station-Rukanga road, Fuden et ul. 691403b (EA); Maungu Hills, S of Maungu Station on Nairobi-Mombasa road, 3'38'S, 38'44'E, Fuden et al. 701157 (EA); Tsavo National Park East, Simba Hill, 3'223, 38'35'E, Fuden & Fuden 741232 (EA, K, MO); 7.3 km on Voi-Taveta road from Nairobi-Mombasa road, 3'269, 38'30'E, Fuden & Fuden 741281 (EA, FT, K, MO, P, PRE, US); 1.5 km from Nairobi-Mombasa road on eastern turnoff to Voi, 3'24'S,38'35'EE, Fuden & Faden 741503 (C, EA, K, LISC, MO, US); Nairobi-Mombasa road, 11.7 km towards Mombasa past Maungu Station, 3'373, 38'50%. Fuden & Fuden 7411285 (EA, K, MO); Taita Hills, 20 Mar 1906, Grenfell s.n. (B, K); Tsavo National Park East, Irima, 3'179, 38'22'E, Hucks 669 (EA, Tsavo Research Centre) (mixture with A. petersii subsp. pallidiflorum); Maungu, Msafiri 307A (EA); Voi, Nupier 914 (EA, K); Tsavo National Park East, 10-15 mi from Aruba on road to Buchuma, Verdcourt 3901 (BR, EA). TANZANIA. BAGAMOYO: Pongwe, 6'19'S,38'142, Harris et ul. BJH4377 (EA, K). HANDENI: Kwa m o n o [Kwamkono], Archbold 713 (K); Kideleko, 7 km S of Handeni, Archbold 1897 (EA). LUSHOTO: Western Usambara Mts., Lushoto- Mombo road, 1.5 mi SW of Gare turnoff, Drummond & Hemsley 2928 (BR, EA, K). MOsHI: Djalla-Krater [Chala], Geilinger 4783 (K). PARE: E South Pare Mts., Kisuani-Same, Greenwuy 2193 (BM, EA, K, PRE); Kiruru, Huurer 447 (EA, K, MO) and I445 (EA, K); 3 km Hedaru-Same road, -4'30'S, 37'54'E, Kubuye & Evans 163 (BR, EA, K, MO); Masandari Hill, 6 km SSW of Same, Wingfield 2899 (EA). NUMBER 76 73 DISCUSSION The type of A. rendlei is recorded by Clarke (1901). Brenan (1952), and Cufodontis (1971) as coming from Somaliland, undoubtedly because of the printed label, "Somaliland, 1895, A. Donaldson Smith," on it. However, there is also a handwritten label with the date "23-4-95." From the account of his journey (Donaldson Smith, 1897), it can be determined that on that date Donaldson Smith was not in Somaliland but in Ethiopia at the foot of the Amaro Mountains (5'29'N, 37"56'10"E, 3670 ft). Aneilema rendlei has not been collected in Somalia. On the same day that he collected the type of A. rendlei, Donaldson Smith also collected his number 346, which is cited by Clarke (1901) under A. rendlei but noted to differ from the type in several characters. Donaldson Smith 346 (BM) is not related to A . rendlei; it is A. sebitense, which is described below in section Lamprodithyros. Corradi 2159, cited by Chiovenda (1951) as A. rendlei, is a mixed collection, consisting mainly of that species but also including a shoot of A. recurvatum. Brenan (1952) supplied a detailed description of A. rendlei but unfortunately based it in part on several specimens that do not belong to this species. Greenway 1668 and Verdcourt 161 (cited as 151) are A. taylorii. Greenway 6058 is described above as A. usambarense. Aneilema octospermum was intended by Clarke to be based upon Grenfell s.n. 6). This specimen falls well within the range of A. rendlei. Among collections with atypical features, Verdcourt 3901 and Newbould 3184 have unusually narrow leaves. Faden & Evans 701788 and Faden & Faden 721240, both from the same population, have bracteoles that regularly are not perfoliate, a feature otherwise unknown in this species. In all other characters these four collections are typical of A. rendlei. The two populations in which variation in lateral petal color was noted were both large. In contrast, the color in Faden & Faden 741232 was determined from a single plant in a small population. Those in Kubuye & Evans 163 and the other cultivated collections were based on material that was probably genetically uniform. The yellow flower color recorded on Bully B8534, is obviously an error. Capsules are almost always confined to the lower cincinni in the inflorescence. This indicates that the upper cincinni rarely produce perfect flowers. Although plants cultivated in Nairobi did occasionally produce some perfect flowers on the upper cincinni, further field observations are required to determine whether this occurs under natural conditions. Small, erect, unbranched plants 1.5-5.5 cm tall have been found associated with larger plants of this species. These small plants have the characteristic swollen shoot bases of A. rendlei and are considered to be first-year seedlings. Such seedlings are able to survive long, dry periods because they store water in the stem and have reduced transpiration in the dry season due to the die off of the laminae and to the presence of overlapping, papery, persistent sheaths covering the stem. A number of such plants, apparently in good condition, were found in the Tsavo area of Kenya in March 1974, at the end of a longer than usual dry season. Plants of this species may take several growing seasons to mature. This species most commonly occurs from 500-700 m altitude in Tanzania and southeastern Kenya. The five collections from above 1000 m have come mainly from northern Kenya and Ethiopia. The altitude recorded on Greenway 2193 should be 3000 ft, not 300 ft. Aneilema rendlei has several attributes that are rare in the genus. The contrasting veins in the paired petals has not been observed in any other species. Other unusual characters of A. rendlei and their distribution within the genus are swollen perennial shoot bases (also in A. petersii subsp. petersii); maroon-spotted staminode lobes (A. taylorii]; J-shaped lateral stamen filaments and style (A. brenanianum, A. taylorii, and probably A. usambarense); a large ellipsoid medial anther with a very convex, maroon-spotted connective (A. brenanianum, A. taylorii, and probably A. usambarense); only glandular microhairs on the ovary (A. brenanianum, A. johnstonii, and A. taylorii). The high degree of stamen dimorphism in A. rendlei is approached only in A. brenanianum. Aneilema rendlei is most closely related to A. brenanianum, A. taylorii, and A. usambarense. Aneilema brenanianum differs from A. rendlei in its habit rhizomatous, roots tuberous, shoots more erect, sheaths becoming papyraceous and completely covering the older parts of the shoots (occurring in A. rendlei only in seedlings), laminae sessile and usually lacking long, uniseriate hairs, paired petals white to very pale bluish purple, pedicels longer, capsules broader and seeds larger. Aneilema taylorii diverges from A. rendlei in its habit decumbent, shoot bases not swollen, thyrses less dense and with fewer cincinni, lateral stamens less dimorphic, fruiting pedicels often decur- ved, capsules smaller and with fewer, smaller seeds. For differences between A. usambarense and A. rendlei see the key. 4. Aneilema brenanianum Faden, new species Aneilema sp. A Faden in Agnew. Upland Kenya Wild Fl., 664,666, 1974. Herbae perennes rhizomatosae radicibus tuberosis, parte inferiore perenni caulium aeriorum vaginis foliorum papyraceis imbricatis inclusa. Folia laminis lineari-lanceolatis ad anguste lanceolato-ellipticas, (3-)6-15 cm longis, (0.4-)0.6-2.1 cm latis, sessilibus. Inflorescentiae thyrsi, 2-5(-7.5) cm longi; 1.5-4(-5) cm lati, cincinnis (5-)7-20(-24) compositi. Axis inflorescentiae puberulus. Cincinni pedunculis (2-)4-7(-8) mm longis, bracteolis (1-)1.5-4 mm distantibus. Pedicelli 5-12 mm longi. Petala lateralia alba vel albida, 4-8.5 mm lata. Capsulae 6-11 mm longae, 4-5 mm latae, glabrae, loculis ventralibus uterque 3(-4)-seminalibus. Semina 2.1-2.5(-2.9) mm longa, 1.8-2.4 mm lata, testa scrobiculatis (Figure 47). TYPE.-Kenya, Masai District (Kajiado District on label), 74 SMITHSONIAN CONTRIBUTIONS TO BOTANY P FIGURE 47.-Aneilema brenanianwn Faden. new species: a, habit; b, inflorescence; c, perfect flower, side view; d, perfect flower, front view; e, staminate flower, side view; f, staminate flower, front view; g, calyx, front view: h, lateral petal; i, medial petal; j , medial staminode; k, lateral staminode; I , medial stamen anther, m, ovary and base of style, side view: n, capsule; 0. seed, ventral view; p . seed, dorsal view (a,c-f,j,m-p, from Faden & Faden 721166; b,g-i,k,l, from Faden et al. 69/2069). (Bars = 20 nun for a; 10 mm forb; 2 mm for c-f,n; 1 mm for g-k; 0.5 mm for I,m,o,p.) NUMBER 76 75 mile 35-36 on Nairobi-Magadi road, 14 Dec 1969, Faden, Evans & Lye 69/2069 (US, holotype; EA, K, MO, isotypes). Perennial herbs (habit type IVB2, Figures 3,47a; Plate 2a). Roots tuberous, brown, to 7 mm thick at base, slightly tapering from there or uniform in thickness for most of their length, to 20 cm long, distal swellings absent. Rhizomes subterranean, unbranched or little branched, moniliform, each segment producing 1-several roots and usually only one aerial shoot. Aerial shoots erect to ascending, rarely straggling, sparsely branched, 15-60(-100) cm tall, the ends dying back to a stiff, perennial, sometimes semi-woody, basal portion. Internodes (1-)1.5-6 cm long, green, those in lower, perennial part of shoots glabrous, completely covered by overlapping, persistent leaf sheaths (Plate 2u), those near the inflorescences puberu- lous, exposed. Leaves spirally arranged, sheaths 1-8.5 cm long, those in lower part of shoot usually greatly overlapping, ribbed longitudinally, becoming gray and papyraceous, persis- tent, puberulous or glabrescent, sparsely ciliate at the apex in young leaves, laminae sessile, linear-lanceolate to narrowly lanceolate-elliptic, (3-)6-15 cm long, (0.4-)0.6-2.1 cm wide, apex acute to acuminate, base narrowly cuneate, margin usually planar, rarely somewhat undulate, scabrid with a mixture of hook-hairs and prickle-hairs, except near the apex where only prickle-hairs present, both surfaces puberulous, abaxial rarely also with a few longer, uniseriate hairs less than 1 mm long. Inflorescences thyrses (Figure 47b; Plate 2c). terminal, moderately dense, ovoid to ovoid-ellipsoid, 2-5(-7.5) cm long, 1.5-4(-5) cm wide, with (5-)7-20(-24) cincinni, alternate or subopposite, ascending. Peduncles 2.5-8(-10.5) cm long, puberulous. Inflorescence bract medial or suprame- dial, herbaceous or membranous, generally very reduced, sometimes foliaceous: Inflorescence axis puberulous. Cincinni to 3 cm long and 9-flowered. Cincinnus bracts membranous, lowermost sometimes perfoliate, patent or appressed to the cincinnus peduncles or sometimes reflexed, lanceolate to ovate, 2.5-4(-7) mm long, glandular near the apex and frequently along the margin toward the base, glabrous or lower ones sparsely puberulous basally. Cincinnus peduncles relatively uniform in length within the inflorescence, usually exceeding the cincinnus bracts, (2-)4-7(-8) mm long, puberulous. Cincinnus axes puberulous. Bracteoles attached (1-)1.5-4 mm apart, herbaceous, eccentrically cup-shaped, perfoliate, 1.8-2.3 mm long, 0.8-1 mm high, with a prominent gland near the apex and usually smaller glands along the margins especially towards the fused edge. Flowers perfect and staminate (Figure 4 7 ~ 9 ; Plate 2b), slightly fragrant, 12-14.5 mm wide. Pedicels 5-7.5 mm long in flower, to 12 mm long in fruit, aligned with the cincinnus axis or above it in flower, erect in fruit, persistent, glabrous or with a few minute hairs at the apex. Sepals with a prominent, subapical, green gland; medial sepal ovate, 3-3.7 mm long, 2-3 mm wide, glabrous or rarely with a few minute hairs at the base, gland inconspicuously bilobed; lateral sepals ovate- elliptic to oblong-elliptic, 3-3.7 mm long, 1.9-2.3 mm wide, glabrous, gland unlobed (rarely bilobed). Paired petals 7- 10.5 mm long, 4-8.5 mm wide, limb ovate or broadly ovate to suborbicular, 5-7.5 mm long, white or whitish (very pale bluish purple), the latter color often noticeable (or appearing?) only when the flowers fade, apex rounded, claw 2-4 mm long, concolorous with limb, glabrous. Medial petal reflexed, ovate to ovate-elliptic, 3-5 mm long, 2-4 mm wide, greenish white. All filament bases free. Medial staminode (Figure 47j) entirely yellow, filament 0.6-1.1 mm long, antherode bilobed, lobes stipitate, flat, ovate or reniform, 0.5-0.9 mm long, 0.5-1.3 mm wide, connective elongate. Lateral staminodes (Figure 47k) with filaments 6.5-8 mm long, question-mark-shaped, very thick at base, tapering apically, entirely yellow or tinged with flesh pink subapically, antherodes bilobed, yellow, lobes stipitate, obovate or elliptic, 0.2-0.4 mm long, connective not elongate. Lateral stamens dimorphic in perfect and staminate flowers (Figure 47cj;. Plate 2b), parallel or slightly divergent, 5-7 mm long (somewhat longer in staminate flowers than perfect flowers), J-shaped, k pointing forward and downward, then recurved at the apex in the perfect flowers, similarly curved but strongly deflexed near the base and with a larger hook near the apex in the staminate flowers, glabrous, anthers elliptic to lanceolate-elliptic, 1.3- 1.9 mm long, 0.7- 1 mm wide, pollen creamy white. Medial stamen with filament 3-4 mm long, tapered from a thick base to apex, anther broadly elliptic or ovate, 1.4-1.9 mm long, 1-1.4 mm wide, usually pendulous with the pollen sacs facing upward, connective very convex, pearly white or cream with maroon spots, pollen creamy yellow. Ovary sessile, oblong-elliptic, 2-2.3 mm long, 1- 1.3 mm wide, glabrous except for some inconspicuous, appressed glandular microhairs scattered all over or k confined to the ventral surface and lateral sutures, apex rounded, dorsal locule represented by a low middorsal hump or ridge, empty, ventral locules each 3-4-ovulate; style 6-7 mm long, J-shaped, held in floral midplane, in lateral view thickest in the middle and tapering towards base and apex, stigma small, held below the anthers but at some distance from them. Capsules sessile (Figure 47n), oblong-elliptic, dehiscent, bivalved, bilocular, 6-11 mm long, 4-5 mm wide, stramine- ous, lustrous, glabrous, apex obtuse to rounded or truncate, valves persistent, dorsal locule obsolete, ventral locules each 3(-4)-seeded, cells of the capsule wall transversely elongate. Seeds (Figure 4709; Plate 5b) trapezoidal, apical ventral rounded apically, basal ventral angular basally, 2.1-2.5(-2.9) mm long, 1.8-2.4 mm wide, hilum black, prominently raised within a groove, slightly cxtcnded onto apical and basal surfaces, testa tan, scrobiculate on all surfaces, the depressions often partially uniting on the dorsal surface, forming larger, irregular depressions, white-farinose in all depressions, around the embryotega and in the groove around the hilum. HABITAT.-Rocky hillsides with numerous boulders and covered by Acacia -Grewia or Acacia-Commiphora, thorny, deciduous bushland, or occasionally dry plains; sandy soil; partial shade; (600?-)1150-1500 m. 76 SMITHSONAN CONTRIBUTIONS TO BOTANY FLOwERING.-FlOWering specimens have been seen from December-April and June. Flowers of Faden & Faden 721166 opened at 1145 hrs in the field on a hot, sunny day. Those of Faden & Evans 711498 did not open until 1700 hrs on a cool, overcast day. The normal opening time is probably about 1200 hrs. Fading time has not been noted. CHROMOSOME NUMBER.-% = 32. DISTRIBUTION.-Very local in S central Kenya (eastern edge of Rift Valley) and adjacent Tanzania (Map 5). SPECIMENS EEN.-KENYA. MASAI: Nairobi-Magadi foad 55.4 km, -l03O?S,36?36?, Faden & Evans 711513 (BR, C, EA, E, K, MO, PRE, US, WAG); mi 35-36 on Nairobi-Magadi road, Faden et al. 6912069 (EA-2 sheets, K, MO, US); Nairobi-Magadi road 55.4 km, Faden & Faden 721166 (BR, EA, K, MO, US); 16 mi from Kiserian towards Magadi on Nairobi-Magadi road, Faden & Napper 691238 (EA, K); 17 mi from Kiserian towards Magadi on Nairobi-Magadi road, Faden & Napper 691251 (EA, K); Magadi, Jan 1962, Heriz Smith s.n. (EA); Nairobi-Magadi road mi 32, Napper et al. 1877 (EA, K); Nairobi-Olorgasailie [Olorgesailie], mi 38, lo33.5?S, 36?292, Napper et al. 1999 (EA, K); 37 mi from Nairobi on Magadi road, Verdcowt et al. 2667 (EA, K, PRE). TANZANIA. MASAI: Longido, just below water tank of Longido (town), 2?43?S,36042?E, Faden & Evans 711498 @A, K, MO, US); bottom slopes of Mt. Longido near new water tanks, Verdcourt 2531 @A, K, PRE). DISCUSSION This species is named in honor of the late J.P.M. Brenan in recognition of his outstanding contributions to our knowledge of the Commelinaceae, especially of Africa, and for his encouragement of my studies of the family. The EA and K sheets of Verdcourt 2531 do not clearly exhibit the papery, overlapping sheaths characteristic of A. brenanianum. The flowering shoots in these specimens, however, have grown directly from the rhizome. This is evidenced by the lower leaves still having their laminae. Apparently the plants were collected fairly early in the growing season because only immature capsules are present. The sheaths have not yet turned gray or papyraceous. The PRE sheet of this collection is more typical of the species, the flowering shoots having arisen from an older shoot base covered by gray, papery sheaths. The habitats in the Kenyan and Tanzanian localities for A. brenanianum are quite similar, with Acacia brevispica, A. senegal, A . thomasii, and Acalypha fruticosa being common in both. The Kenyan localities also have Acacia mellifera and Grewia species as dominants, while the Tanzanian locality has Commiphora species common in addition to the Acacia species. The mean annual rainfall in these areas is 250-500 mm (Mean Annual Rainfall Map of East Africa, 1959), with the bulk of the precipitation falling in March to May and October to December (National Atlas of Kenya, 1970). Plants of A. brenanianum nearly always grow with their roots in crevices between boulders, making them almost impossible to extract whole. This adaptation allows them to accumulate additional moisture in the form of runoff and probably tends to reduce water loss in the dry season. It may also serve as a protection against root-eating rodents such as mole rats (Bathyergidae). The succulent roots and rhizomes function as storage organs between rainy seasons. The growth habit of A. brenanianum is an adaptation to short and unpredictable growing periods followed by long dry seasons. The overlapping, papery leaf sheaths that cover the lower internodes of the aerial shoots probably reduce transpira- tion and enable these shoot bases to survive long, dry periods. This covered-internode growth precedes the exposed-internode growth that leads to flowering, although a certain amount of the latter is produced concurrently from the previous growing season?s persistent shoots. The exposed internode portions of the shoots, whether flowering or not, die back at the end of the rainy season. If the rainy season is unusually short, most of the plant?s growth will have been covered-internode growth and is thus protected against the ensuing dry season. The Kenyan collections, with the possible exception of Heriz Smith s.n., are all from an 8 km section of the Nairobi-Magadi road. The Heriz Smith specimen might also have come from this area, as the name ?Magadi? is often used in a general sense for the whole region of Lake Magadi and the dry, low altitude area to the north of it. If the Heriz Smith collection came instead from the vicinity of the town of Magadi, then the 600 m altitude, recorded with uncertainty above, would be correct. The local distribution of A. brenanianum is not just an artifact of limited collections. The presence of a local endemic in this part of East Africa is consistent with the isolation of the very dry habitats of the region from equally dry conditions to the east and north by about 200 and 400 km, respectively. Other endemics in this area include species of Turraea (Meliaceae: Dale and Greenway, 1961) and Barleria (Acanthaceae: Napper and Agnew, 1974). Aneilema brenanianum is most distinctive because of its persistent shoot bases covered by overlapping, papery sheaths. These shoot bases can become very stiff and semi-woody, with the consistency of dead grass culms. Overlapping sheaths, but without stiff shoots, have been observed only in A. longir- rhizum, occasional plants of A. hockii, and seedlings of A. rendlei. The aspect of the plants of A. brenanianum suggests affinities with A. longicapsa, A. obbiadense, A. somaliense, and A. longirrhizum. However, it differs from all of them in many floral characters, and their vegetative similarity is undoubtedly due to convergence. The floral characters of A. brenanianum indicate that it is most closely related to A. rendlei, A . taylorii, and A. usambarense. All of them differ in lacking rhizomes and NUMBER 76 77 tuberous roots, and in having petiolate leaves with long uniseriate hairs on both surfaces, a completely glabrous inflorescence, lilac or lavender flowers, narrower capsules, and smaller seeds. Aneilema brenanianum further differs from A. rendlei, its closest relative, in having smaller antherode lobes that always lack maroon spots, shorter medial staminode filaments and styles, longer lateral staminode filaments, less-dimorphic lateral stamen filaments, and generally fewer- seeded capsules. Its flowers also open considerably later than those of A. rendlei. Section 4: Somaliensia Faden, new section Pedicelli fructuum erecti ad recurvatos. Petalum medium redactum. Stamina lateralia filamentis basi libris, sigmoideis, glabris, infra medium geniculatis. Stamen medium anthera ephippiomorpha. Ovarium puberulum loculo dorsali plerum- que bene formato et 1-ovulato, loculis ventralibus uterque 2(-4)-ovulatis. Capsulae puberulae. TYPE SPECIES-Aneilema somaliense C.B. Clarke. Tuberous-rooted perennials with annual flowering shoots often disarticulating at the nodes at the end of the growing season (roots and habit of A. grandibracteolatum unknown). Leaves usually spirally arranged, sheaths mostly short, and laminae sessile. Inflorescences terminal thyrses with mostly alternate cin- cinni. Inflorescence axis puberulous with uniform-length hook-hairs, lacking uniseriate hairs. Bracteoles generally membranous, cup-shaped, perfoliate or not, glandular near the apex. Flowers perfect and staminate, or pistillate, perfect, and staminate. Fruiting pedicels erect or recurved to -270?, persistent. Sepals puberulous with uniform-length hook-hairs. Petals glabrous, the medial reduced. Filament bases free. Medial staminode with antherode lobes sessile or shortly stipitate and connective slightly to not at all elongate. Lateral staminodes with filaments straight, then sharply decurved near the apex, antherode lobes stipitate. Lateral stamens with filaments S-shaped, geniculate (or probably so) below the middle, glabrous. Medial stamen with anther saddle-shaped. Ovary densely and uniformly eglandular- or glandular- puberulous, dorsal locule generally well developed and 1-ovulate, ventral locules each 2(-4)-ovulate, stigma usually small. Capsules sessile or stipitate, dehiscent, bivalved, generally trilocular, lustrous, dorsal lccule usually developed, 1 -seeded or empty, ventral locules each 2(-4)-seeded. Seeds with a short or moderately long hilum, testa with pits and/or grooves on the dorsal surface radiating from the embryotega, farinose or not. BASIC CHROMOSOME NUMBERS.-x = 14,16. DISTRIBUTION.~omali Republic and E Ethiopia, S to central Kenya (Map 6). Key to the Species of Section Somaliensia 1. Inflorescences composed of (8-) 15-50 cincinni; cincinnus peduncles (2-)5-27 mm long. 2. Bracteoles not perfoliate; capsules (5-)6-9 mm long, apex usually rounded to truncate; seeds 2-3(-4) per ventral locule, 1.6-2.7 mm long, white-farinose in the depressions . . . . . . . . . . . . . . . . . . . . . . . . 1. A. sornaliense 2. Bracteoles usually perfoliate (rarely not); capsules (10.5-)12-17 mm long, apex f rostrate; seeds 2 per ventral locule, 3.4-4.55 mm long, not farinose (mature seeds unknown in A. obbiadense). 3. Bracteoles (2.5-)3-3.5 mm long, attached (2-)2.5-3.5 mm apart; cincinnus peduncles 7-13 mm long . . . . . . 2. A. grandibracteolaturn, new species 3. Bracteoles (1.5-)1.9-3 mm long, attached 1.5-22 mm apart: cincinnus peduncles 10-27 mm long. 4. Cincinnus peduncles 15-27 mm long; bracteoles attached 9-22 mm apart; lamina lanceolate to ovate, 1-2 cm wide . . . . . . . . 3. A . obbiadense 4. Cincinnus peduncles 10-20 mm long; bracteoles attached 1.5-7.5 mm apart: lamina usually linear-lanceolate and 0.35-0.8 cm wide, rarely lanceolate to lanceolate-elliptic and 0.8-1.9 cm wide . . . . . . . . . . . . . . . . . 1. Inflorescences composed of 2- 11(- 13) cincinni; cincinnus peduncles (1.2-)1.5- 5.5(-8.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . 5. A. pusillurn . . . . . . . . . . . . . . . . . . . . . . . 4. A. longicapsa, new species 78 SMITHSOMAN CONTRIBUTIONS TO BOTANY MAP 6.-Distribution of Aneilema section Somalienria, new section. DISCUSSION The five species recognized in section Somaliensia are mostly inadequately known, with only A. somaliense and A. pusillum being represented by more than seven collections each. In particular, the relationships among A . grandibracteola - turn, A . obbiadense, and A. longicapsa, collectively known from nine gatherings, need further investigation. Similarly, the treatment of A. pusillurn as a species with four subspecies should be considered provisional, for it is based on only 14 collections. The species in section Somaliensia fall into three distinct groups, the first consisting of A . somaliense, the second of A. grandibracteolaturn, A. obbiadense, and A. longicapsa, and the third of A. pusillum. These groups are characterized mainly by inflorescence, capsule, and seed characters, which are given in the key and in the discussions under the individual species, and perhaps also by chromosome number. In view of our incomplete knowledge of most of these m a , particularly of floral morphology, it would be premature to recognize subsections at this time. 1. Aneilema somaliense C.B. Clarke Aneilema somalienre C.B. Clarke in Thiselton-Dyer, Fl. Trop. Africa. 8:69, 1901.-Chiovenda, Resultati Scientifici Missione Stefanini-Paoli Somalia Italiana, 1:167,1916; Webbia 8:38,1951.-Cufodontis, Bull. Jard. Bot. Natl. Belgique, 41, suppl., 1518, 1971. [Type: Ethiopia (Somaliland on label, see below), Harradigit, Mar (1885). James & Thrupp s.n. (K).] NUMBER 76 79 Aneilema smithii C.B. Clarke in Thiselton-Dyer, FL Trop. Africa, 8:70, 1901.-Cufodontis, Bull. Tard. Bot. Natl. Belgique, 41, suppl., 1518, 1971. [Syntypes: Ethiopia, Lake Rudolf to Gondokoro. Jara. 23 Oct 1899, Donaldson Smith s.n. (BM, lectotype); Ethiopia, Lake Rudolf to Gondokoro, Anole, 21 Oct 1899, Donaldson Smith s.n. (BM).] Aneilema tacazzeanwn sensu Chiovenda, Missione Biol. Paese Borana, 4:305. 1939 [pro Cufodonh 319, non Hochstetter ex A. Richard (ISSO)]. Perennial herbs (habit types IVBI, IVB2, Figure 3). Roots tuberous, to 60 cm long, with distal fusiform to cylindrical tubers to 24 cm long. Perennial shoot bases subterranean, short, erect, sometimes a few connected, forming a short rhizome; flowering shoots annual, sparsely to profusely branched, erect to ascending, lower parts sometimes prostrate or looping on ground but not rooting at nodes, shoots disarticulating at base and nodes at end of growing season, 7.5-40 cm tall. Internodes (0.7-)1.5-8(-10) cm long, green, puberulous and rarely sparsely pilose. Leaves spirally arranged, sheaths 0.3-1(-1.5) cm long, puberulous and rarely sparsely pilose along the fused edge, ciliate at the apex, laminae sessile, lanceolate-elliptic to elliptic, ovate-elliptic or ovate, rarely broadly elliptic, (1.5-)2.5-7.5(-13) cm long, (1-)2-3.5(-5.5) cm wide, apex acuminate to acute, rarely obtuse or rounded, and then mucronate, base cuneate to rounded, rarely subcordate, margin planar to strongly undulate, scabrid and frequently ciliate, especially towards the base, both surfaces gray-green, lustrous, puberulous, usually with few to many, patent, uniseriate hairs less than 1 mm long, more numerous on abaxial surface (occasionally lacking on adaxial surface); midvein impressed on adaxial surface, prominent on abaxial surface. Inflorescences thyrses (Plate 2e). terminal, moderately dense, ovoid to ellipsoid or cylindrical, (2-)3-9.5 cm long, (1.5-)2-4.5 cm wide, with (8-)17-40(-50) cincinni, alternate to subopposite or occasionally subverticillate, patent to ascending; occasionally a solitary cincinnus produced from the axil of the inflorescence bract. Peduncles 3-9 cm long, puberulous. Inflorescence bract medial to supramedial, com- monly herbaceous and foliaceous, sometimes membranous and much reduced. Inflorescence axis puberulous. Cincinni up to 2.5 cm long and 8-flowered. Cincinnus bracts usually membranous, appressed to the cincinnus peduncles, ovate to lanceolate-ovate (rarely lanceolate), 1-2.5(-5) mm long, hooded and glandular apically, puberulous basally or entirely glabrous. Cincinnus peduncles shortest in lower cincinni, longest in middle cincinni, slightly reduced in uppermost cincinni, exceeding the cincinnus bracts, (2-)5-11(-13) mm long, greenish violet, puberulous. Cincinnus axes puberulous. Bracteoles attached 1-33-4) mm apart, membranous, ovate, cup-shaped, amplexicaul but not perfoliate, 0.9-1.4(-1.7) mm long, with a prominent subapical gland, often puberulous, at least basally, rarely with a few uniseriate hairs along the margin towards the base, sometimes glabrous. Flowers pistillate, perfect and staminate (Plate 2f,g), slightly fragrant, (7.5-)lo-12.5(-17) mm wide. Pedicels 2.8-4.5(- 5.5) mm long in flower, to 7 mm long in fruit, held in line with the cincinnus axis or slightly below in flower, erect to slightly recurved (rarely up to 180') in h i t , persistent, glabrous, or occasionally with a few hook-hairs at the apex. Sepals with a prominent, green, subapical gland, puberulous; medial sepal ovate, (2-)2.8-3.5 mm long, 1.8-3.1 mm wide, gland inconspicuously bilobed; lateral sepals ovate-elliptic to lanceo- late-ovate, (2-)2.4-3.1 mm long, 1.6-2.7 mm wide, gland inconspicuously bilobed or unlobed. Paired petals 4.8-9 mm long, 3.2-7 mm wide, limb ovate, 2.8-6 mm long, white to very pale lilac or pale blue (Bally B10133). apex rounded, usually slightly cucullate, claw 2-3 mm long, concolorous with limb, glabrous. Medial petal reflexed, ovate to lanceolate, elliptic or obovate-elliptic, 2.8-6 mm long, 1.6-4.5 mm wide, green medially with whitish margins, sometimes tinged with red apically. All filament bases free. Medial staminode with filament 1.3-2.7 mm long, yellow, sometimes purplish basally, antherode bilobed, yellow, lobes shortly stipitate, lanceolate, elliptic or broadly reniform, 0.4-1 mm long, connective scarcely elongate. Lateral staminodes with filaments 2.8-4.5 mm long, sharply decurved near the apex, purplish at base, yellow above, antherodes bilobed, yellow, lobes stipitate, obovate-cuneate to ellipsoid or suborbicular, 0.4-0.9 mm long, connective not elongate. Lateral stamens with filaments divergent in basal half, more strongly divergent apically (sometimes parallel or slightly convergent near the apex), 5-6.5 mm long, gently S-shaped, pendulous, glabrous, anthers elliptic to ovate, 0.6-1.3 mm long, 0.5-1 mm wide, pollen yellowish white. Medial stamen with filament 3.5-5 mm long, anther saddle-shaped, 0.9-1.1 mm long, 0.9-1 mm wide, connective broad, greenish white, pollen yellow or orange- yellow. Ovary sessile, oblong-elliptic, 0.9-2.5 mm long, 1.1-1.3 mm wide, densely and uniformly (except for a narrow V extending basally from the style base along the middorsal line) covered with spreading glandular hairs (Figure 42c), apex acute, dorsal locule represented by a prominent hump, empty or 1-ovulate, ventral locules (2-)3(-4)-ovulate; style 5.5-6.2 mm long, arcuate-decurved for most of its length, recurved near the apex, passing through the upper notch of the medial anther, held in floral midplane, sometimes with a few glandular hairs, like those of the ovary, at Lhe base, stigma small, held in front of the anthers. Capsules shortly stipitate, oblong-elliptic, bi- or rarely somewhat uivalved, bi- to trilocular, (5-)6-9 mm long, 3.2-4(-4.8) mm wide, stramineous to brown, lustrous, sparsely puberulous, apex rounded to truncate, occasionally emarginate or acute, valves persistent or occasionally dorsal valve tardily deciduous, dorsal locule occasionally obsolete but usually represented by a middorsal hump overlying the first and second pairs of seeds (from the base), empty or 1-seeded, ventral locules each 2-3(-4)-seeded. Seeds (Plate 5g) trapezoi- dal, rectangular, triangular or transversely ellipsoid, ventral apical rounded apically, ventral basal angular basally, 1.6-2.7 mm long (1.2-)1.3-1.6(-2) mm wide, hilum dark brown to dark gray, prominently raised within a groove, not at all to 80 SMITHSONIAN CONTRIBUTIONS TO BOTANY MAP 7.--Distnhtion of Aneilema somalieme C.B. Clarke (solid circles), A. grandibracieoIatwn Faden, new species (open circle), and A. Iongicapsa Faden, new species (solid squares). slightly extended onto apical and/or basal surfaces, testa pinkish brown to grayish tan or tan, intermptedly furrowed on the dorsal surface in a radiating pattern from the embryotega, some of the furrows continuing onto apical, basal and lateral surfaces, whitish farinose in all of the grooves and depressions and around the embryotega and hilum. HABITAT.-Acacia or Acacia-Commiphora bushland or woodland, dwarf shrubland or degraded montane scrub, sometimes on rocky slopes; sandy, silty or loamy soils; generally growing in partial shade; 750-1700 m. FLOwERING.-~OWeritIg specimens have been seen from March-May, July, and October-December. In cultivation the flowers are open in the morning and fade in the early afternoon, but exact opening and fading times have not been determined. CHROMOSOME NUMBER.-n = 16,2n = 32. DISTRIBUTION.-E and S Ethiopia, S Somali Republic and N Kenya (Map 7). SPECIMENS SEEN.-ETHIOPIA. HARAR: Ogaden Reserved Area, 37 mi E of Awareh, Bully B10133 [in CAI222981 (EA, K); Harradigit, Mar [1885], James & Thrupp s.n. (K) . SIDAMO: 50 km S of Neghelli on Wachelli road, Ash 823 (K); El Banno (Tcertale), Corradi 2196 0; h e r o (Meta-Gafersa), Cufodon- tis 319 (FT); Lake Rudolf to Gondokoro, Jara, 23 Oct 1899, Donuldson Smith s.n. (BM); Lake Rudolf to Gondokoro, Anole, 21 Oct 1899, Donaldson Smith s.n. (BM); Moyale area, 4"23'N, 38'252, Everurd in EA11456 (EA) [wrongly recorded as coming from Kenya on the label]; 10 km N of Neghelle, along road to Kebre Mengist, Friis et al. 2670 (K); Old Airfield -15 km NNE of Yavello, 4'59", 38'132, Gilbert & Jefford 4559 (K); 46 km from turning to Mega on road from Negele, near Bulbula River, -4"58'N, 39'372, Thulin et al. 3539 (K). SOMALI REPUBLIC. UPPER JUBA: Piana di El UalAc, Paoli 1085 (IT-2 sheets). KENYA. MERU: Isiolo, J. Adamson 599 (EA). NORTHERN FRONTIER: Mathews Range, Olkanto, J . Adamson 42 in [Bully] B4343 (K); Gof Choba, 11 km N of Marsabit, 2'24,5'N, 38'032, Carter & Stannard 652 (EA); 30 km S of Marsabit on Isiolo road, 2'08'N, 37'53'E, Carter & Stannard 692 (EA); South Horr Valley, Davey 079 (EA); 2 km WSW of Wamba, 0?57'N, 37'192, Faden & Faden 74/923 (BR, EA, FT, K, MO, US); Matakweni Hill, 3 km WSW of Wamba, 0?57'N,37'18'E, Fuden & Fuden 74939 (EA, lT, K, MO); Wamba-Isiolo road, near pass between Lekat and Olowa Werikoi Hills, 0?52'N, 37'24'E, Faden & Faden 74/951 @A, K, MO, PRE); Marsabit Mountain, Gugul 556 (EA); 01 Lolokwe (01 Doinyo Sabachi), E side of base, Oo52'N, 37'34'E, Gilbert 5018 (EA); 5 km E of junction of Banissa and Derkali roads, -3'55'N, 4Oo22'E, Gilbert & Thulin 1452 (EA, F, US); 48 km on Ramu-El Wak NUMBER 76 81 mad, -3?31?N, 41?00?E, Gilbert & Thulin 1619A 0; Subata Microwave Relay Station, NE part of 01 Lolokwe (01 Doinyo Sabachi), Oo53?N, 37?34?E, Gilbert et al. 5303 @A); Dandu, 3?26%. 39?54% Gillett 12651 (K, PRE; EA, photo) and 12702 (B, BR, EA, K, S); Moyale, 3?32?. 39?03?E, Gillett 14032 @A, K); [Marsabit District], Ngarunit fenced plot near outer airstrip, Herlocker 757 0; Archer?s Post, Samburu Park, Makin 233 @A, K); Wamba, Newbould 2957 (FT, K). DISCUSSION There has been difficulty in localizing the types of A. somaliense and A. smithii. The type of the former has ?Somaliland? written on the sheet, but the route followed by James and Thrupp, given in Chiovenda (1929) as ?James 1885,? shows that part of their journey was within the present boundaries of Ethiopia. The locality of the type, ?Harradigit,? is undoubtedly the same as ?Harar Digit,? for which Bally (personal communication) provided the coordinates 7?4734 44?27?E. This is close to the route shown by Chiovenda and is in Ethiopia, not the Somali Republic. The Donaldson Smith collections that are the syntypes of A. smithii were recorded from ?British East Africa, Lake Rudolph? by Clarke (1901). From the collector?s account of his journey (Donaldson Smith, 1900) and the dates on the specimens, it has been determined that he was ENE of Moyale, near the present Kenya-Ethiopia border, not at Lake Rudolf (now Lake Turkana), as written on the labels, about the time he made these collections. Both specimens were probably col- lected in Ethiopia between Eimole (4?03?N, 40?12%) in Kenya and El Der (3?52?30?N, 39?46%) in Ethiopia. The types of A. somaliense and A. smithii are very different looking vegetatively. The former, although consisting of only a single flowering shoot with the upper three leaves, is obviously a much larger plant with longer, proportionally narrower leaves than the latter. The only reproductive characters that appcar to distinguish the types are the longer sepals and pedicels of A. somaliense. When a wider range of material was studied, it became apparent that these vegetative and reproductive differences were not taxonomically signifi- cant. Aneilema somaliense and A. smithii were described in the same work (Clarke, 1901) and have not been combined previously. The former name has been selected because it suggests the general range of this species. Aneilema somaliense is a very variable species. Most of that variation, however, is in the vegetative parts, and almost none of it is geographically based. The only exception is the collection Paoli 1085, the sole gathering from the Somali Republic, which is unusual in several of its shoots having broadly elliptic leaves that are sometimes rounded- or obtuse-mucronate at the apex. It is also unusual in some of its reproductive features: bracteoles frequently with uniseriate hairs on the margin towards the base and seeds radiately pitted instead of furrowed. In all other characters, Paoli 1085 falls within the normal range of A. somaliense. There are at present insufficient grounds to separate it taxonomically. Some variation observed in cultivated plants has not been found in field collected specimens. A cultivated plant of Faden & Faden 74/923 produced small lateral shoots from many of its nodes. These shoots generated from their bases short, thick roots that perforated the leaf sheaths. When detached, these shoots could be grown as independent plants. This formation of plantlets has not been observed in specimens of A. somaliense nor in any other species of Aneilema, cultivated or wild. The dimensions of many structures in the cultivated plants of Faden & Faden 74/923 exceeded those of all other collections of this species. The measurements of the floral parts, capsule width, and seed length and width have been considered reasonable extensions of the range of variation and have been included in the description. The following have been omitted as being unlikely to occur in the field: inflorescences to 6 cm wide; cincinni all strongly ascending, up to 5 cm long and 12-flowered, with bracteoles attached up to 5 mm apart. Within a small geographic area A. somaliense can occupy quite diverse ecological sites. Faden & Faden 74J939 was growing on a steep rocky slope beneath a massive granitic hill. It was found under shrubs of Acalypha fruticosa and an undescribed, spiny species of Barleria and was associated with the fern Actiniop- teris radiuta. Only one kilometer away, the same collectors? 74J923 was found in level Acacia tortilis subsp. spirocurpa-A. brevispica bushland growing in sandy soil with much bare ground between the trees and thicket clumps. These collectors? third collection in the same region (74/951) came from Commiphora bushland, in which the shrubs Euphorbia cuneata and Boscia coriaceu, both absent from the other two localities, were common. In all three of these habitats, plants of A. somaliense occurred in protected spots under trees and shrubs. The two annual flowering periods for A. somaliense coincide with the rainy seasons in its range. The Faden and Faden collections confirm that the flowering shoots die off completely during the long, dry season (June-September). It is unknown whether individual plants flower in both rainy seasons. The relationships of A. somaliense have been difficult to assess. In overall appearance it shows similarities to some species in section Brevibarbata, e.g., A. lanceolatum and A. welwitschii. It differs from all of them in its glabrous stamen filaments and basic chromosome number. It also occurs entirely outside of the range of all species in that section. The similarities are probably due to convergence. Aneilema somaliense also shares a number of features with species in section Rendlei, particularly A. rendlei and A. brenanianum: inflorescence form, scented flowers, glabrous filaments, small stigma, number of seeds per capsule locule, farinose testa, basic chromosome number, and ecology. However, those species have quite different habits, lacking distal tubers on their roots and having at least the basal part of the aerial shoots persisting during the dry season. They further 82 SMITHSONIAN CONTRIBUTIONS TO BOTANY differ by lacking pistillate flowers, having a totally different androecial morphology, lacking conspicuous hairs on the ovaries and capsules, rarely having a well-developed dorsal locule in the capsule, and having seeds with a scrobiculate testa. The affinities of A. somaliense with section Rendlei species probably are not close. As much through the absence of important discordant characters as through the presence of strong similarities, A. somaliense appears to be most closely related to the four other species placed in section Somaliensia. Although the data that have accumulated on these taxa since the decision to recognize section Somaliensia was made (Faden, 1975) have tended to demonstrate the more isolated position of A. somaliense within the section, important similarities have also been confirmed: tuberous roots, disarticulating, annual flowering shoots, short sheaths, sessile laminae, S-shaped, glabrous, lateral stamen filaments, saddle-shaped medial anther, well-developed dorsal locule in the ovary, small stigma, and radiately pitted or grooved testa. Aneilema somaliense differs from the other species in the section in having non-perfoliate bracteoles, glandular hairs on the ovary, more numerous ovules per ventral locule, basally angular ventral locule basal seeds, hilum in a groove that is not cut off at either end, and farinose testa. 2. Aneilema grandibracteolatum Faden, new species Caules floriferi ad nodos disarticulati. Folia laminis lanceo- latis vel lanceolato-ellipticis, 2.5-7.5 cm longis, 1-2.5 cm latis. Inflorescentiae thyrsi, 2-6 cm longi, 2-7 cm lati, cincinnis -20-30 compositi. Bracteae cincinnorum (2.3-)3.5- 5.8 mm longae. Cincinni pedunculis 7-13 mm longis, bracteolis (2-)2.5-3.5 distantibus. Bracteolae (2.5-)3-3.5 mm longae. Capsulae stipitatae, rostratae, (10.5-)12-14 mm longae, 3.5-3.8 mm latae, loculis ventralibus uterque 2- seminalibus. Semina 3.4-3.9 mm longa, 1.6-1.9 lata; testa aurantiaco-brunnea foveis sulcisque non profundis radiatis, granulis farinosis destituta. TYPE.-Ethiopia (British Somaliland on label), between Walwal and Bulleh (Sirauw), 26 Nov 1944, Glover & Gilliland 408 (K, holotype, photo F; EA, isotype). Herbs, probably perennial (habit type unknown, probably NB2, Figure 3). Roots unknown. Flowering shoots probably annual, erect to ascending, disarticulating at the base and nodes when dry, to 15 cm tall. Internodes 0.7-4 cm long, puberulous. Leaves spirally arranged, sheaths to 5 mm long, puberulous, sparsely ciliate (with short uniseriate hairs) or ciliolate with hook-hairs) at the apex, laminae sessile, lanceolate to lanceo- late-elliptic, 2.5-7.5 cm long, 1-2.5 cm wide, apex acute to obtuse, base cuneate to rounded, margin usually undulate, scabrid with prickle hairs near the apex, otherwise beset with numerous hook-hairs, both surfaces puberulous, uniseriate hairs absent, except for some, poorly differentiated from prickle-hairs, midvein impressed on adaxial surface, prominent on abaxial. Inflorescences thyrses, terminal, moderately dense to dense, ovoid to broadly ellipsoid, 2-6 cm long, 2-7 cm wide, with about 20-30 cincinni, alternate to subopposite, the lowermost often subverticillate, ascending to patent, the lowermost becoming declinate. Peduncles 1 M . 2 cm long, puberulous, distance between inflorescence bract and lowest cincinnus 0.6-1.6 cm. Inflorescence bract supramedial to medial, foliaceous or reduced and bract-like. Inflorescence axis puberulous. Cincinni up to 3.5 cm long and 8-flowered. Cincinnus bracts scarious, appressed to the cincinnus pedun- cles, becoming reflexed with age, lanceolate to lanceolate- elliptic (2.3-)3.5-5.8 mm .long, glandular near the apex, glabrous to sparsely puberulous. Cincinnus peduncles rela- tively uniform in length within the inflorescence, or the lowermost slightly reduced, exceeding the cincinnus bracts, 7-13 mm long, puberulous. Cincinnus axes puberulous. Bracteoles attached (2-)2.5-3.5 mm apart, sometimes slightly overlapping, herbaceous, eccentrically cup-shaped, perfoliate or sometimes not, (2.5-)3-3.5 mm long, to 2.5 mm high, inconspicuously glandular near the apex, puberulous at least towards the base. Flowers probably perfect and staminate (only the latter seen). Pedicels 2.5-4.2(-5.7) mm long in flower, to 8.5 mm long in fruit, aligned with the cincinnus in flower, erect to recurved (to -150?) in fruit, persistent, glabrous or sparsely puberulous at the apex. Sepals puberulous; medial sepal -3.2 mm long, cucullate and glandular apically; lateral sepals elliptic to oblong-elliptic, -3.5 mm long x 2.2 mm wide, inconspicuously glandular apically. Paired petals ?pale blue,? -6-6.5 mm long, claw -1.5 mm long. Medial petal reduced, -4-4.5 mm long. All filament bases free. Medial staminode yellow, filament 0.7-1.5 mm long, slightly broadened apically, antherode bilobed, the lobes sessile, obovate to suborbicular, -0.3 mm long x 0.4 mm wide, broadly or narrowly attached to the filament, connective slightly elongate. Lateral staminodes yellow, filaments -4 mm long, antherodes bilobed, lobes stipitate, obovate- to suborbicular-cuneate, -0.3 mm long x 0.25 mm wide, stalks -0.15 mm long, attached at about right angles to the filament, connective not elongate. Lateral stamens with filaments -8.5 mm long, geniculate below the middle, glabrous, anther elliptic, 1-1.5 mm long, -1 mm wide. Medial stamen with filament -5-5.5 mm long, anther saddle-shaped, 1.3-1.6 mm long. Ovary not seen but, extrapolating from capsule, probably stipitate, puberulous, apex acute, tapering into the style, dorsal locule l-ovulate or empty, ventral locules each 2-ovulate; style (on young capsule) -9 mm long, stigma small. Capsules stipitate, oblong-elliptic, tapering at both ends, slightly falcate, dehiscent, bivalved, bi- to trilocular, (10.5-)12-14 mm long, 3.5-3.8 mm wide, stramineous, lustrous, evenly puberulous on both surfaces and all sutures, glandular hairs occasionally mixed with a few hook-hairs, apex f rostrate, valves persistent, slightly spreading, dorsal locule obsolete or 1-seeded, ventral locules each 2-seeded, cells of the NUMBER 76 83 capsule wall transversely elongate, in regular files. Seeds (Plate 5c) transversely elliptic, the apical ventral rounded apically, the basal ventral rounded basally, 3.4-3.9 mm long, 1.6-1.9 mm wide, hilum dark gray, much shorter than the seed, prominently raised within a groove, groove cut off apically and basally by ridges, with apical, basal and lateral edges crenate, embryotega pale pinkish gray, testa orange-brown, with very shallow pits and grooves radiating from the embryotega on the dorsal surface, the pits and grooves becoming deeper with distance from the embryotega, farinose granules lacking. HABITAT.-"Open areas on sand" (Glover & Gilliland 408), probably in bushland. FLOWERING.-The single collection from late November has flowers and mature capsules. It had probably been flowering for at least 3-4 weeks and was beginning to dry up, judging from the scarcity of large flower buds. CHROMOSOME NUMBER.-Unknown. DISTRIBUTION.-E Ethiopia (Map 7). SPECIMENS EEN.-ETHIOPIA. HARAR: Between Walwal and Bulleh (Sirauw), Glover & Gilliland 408 (EA, K). DISCUSSION The Glover and Gilliland collection records the country as British Somaliland, but the coordinates for the two localities mentioned on the label indicate that the plants actually came from the Ogaden region of Ethiopia. The locality Sirauw (= Bulle Sirau) appears only on the EA specimen. This species is clearly closely related to A. obbiudense and A. longicapsa because of its longly stipitate, roseate, curved capsules and large,. transversely elliptic seeds with a short hilum prominently raised within a groove that is cut off at one or both ends by ridges. Aneilemu grandibracteolatum differs from A. obbiadense by its longer leaves and much smaller, denser inflorescences, from A. longicapsa by its broader leaves and smaller, more orange seeds, and from both by its shorter cincinnus peduncles and larger, less widely spaced bracteoles. Apparently it is also geographically isolated from them, based on the limited number of collections available. Floral charac- ters, when they can be determined from living plants, will probably be decisive in establishing the final taxonomy of these species. The large, pointed capsules in these three species and their northeastem African distributions, suggest a possible affinity with A. gillettii of section Amefina. However, that species has mostly subverticillate cincinni, bearded lateral stamen fila- ments, concolorous petals, sessile, more or less glabrous capsules with more seeds per locule, and farinose seeds of a different structure. The resemblance of the capsules is almost certainly due to convergence. A recent collection from the Somali Republic, Thulin 5667, is somewhat intermediate between A. grandibracteolatum and A. longicapsa. It is discussed under the latter species. 3. Aneilema obbiadense Chiovenda Aneilemu obbiadense Chiovenda ["Obbiadensis"], P1. Novae Aethiopia, 10, 1928; Fl. Somala, 1:316,1929.-Cufodontis, Bull. lard. Bot. Natl. Belgique, 41, suppl., 1517. 1971. [Type: Somali Republic (Somalia centrale on label), Sultanato di Obbia, Tobungiib. 4 May 1924, Puccioni & Stefanini 592 [646] O v . 1 Perennial herbs (habit type IVB2, Figure 3). Roots tuberous, -4 mm thick at base, then abruptly narrowed and uniformly 2-3 mm thick for rest of their length, to 28 cm long (or longer?), distal swellings apparently absent but roots in specimens incomplete. Perennial shoots rhizomes, subterra- nean, moniliform, unbranched or sparsely branched, 7.5 cm long in specimen, each segment producing several roots and probably only one aerial shoot. Flowering shoots annual, erect, sparsely branched, disarticulating at base and nodes on drying. Internodes 1-2 cm long, puberulous. Leaves spirally arranged, sheaths often split to base or nearly so, to 0.5 cm long, puberulous, sparsely ciliolate with short uniseriate hairs or hook-hairs at the apex, laminae somewhat succulent, sessile, lanceolate to ovate, 2-5 cm long, 1-2 cm wide, apex acute, base rounded to broadly cuneate, margin undulate, scabrid with prickle-hairs near the apex and hook-hairs below, both surfaces puberulous, lacking uniseriate hairs. Inflorescences thyrses, terminal, lax (or moderately lax), ovoid to subspherical, 4.5-11 cm long, 5.5-12(--20) cm wide, with 15-21 cincinni, mostly alternate (or a few subopposite), except the lower, which are subverticillate, ascending, patent, or declinate. Peduncles 3-4 cm long, puberulous. Inflorescence bract medial, bract-like or foli- aceous. Distance between inflorescence bract and lowest cincinnus 1-2 cm. Inflorescence axis puberulous with uniform- length hook-hairs, sometimes blue-tinged. Cincinni to 12 cm long and 6-flowered. Cincinnus bracts scarious, patent to ascending, ovate to ovate-elliptic, 2.5-3.5 mm long, glabrous or subglabrous. Cincinnus peduncles relatively uniform within the inflorescence, 15-27 mm long, sometimes blue-tinged, puberulous. Cincinnus axes puberulous with uniform-length hook-hairs, sometimes blue-tinged. Bracteoles attached 9-22 mm apart, scarious, cup-shaped, perfoliate, 2-2.7 mm long, 1.1-2 mm high, not clearly glandular, glabrous or with a few minute hairs near the base. Flowers perfect and staminate. Pedicels 4.5-7 mm long in flower, to 10 mm long in fruit, aligned with the cincinnus axis in flower, strongly recurved (-180'-270') in fruit, glabrous or sparsely puberulous. Sepals 3-6 mm long, glandular near the apex, but not very prominently so, (sparsely) puberulous with uniform-length hook-hairs. Paired petals lilac, 6 mm long and wide (fide type description). Medial petal narrow. All filament bases free. Medial staminode -2.5 mm long, yellow, filament broadened apically, antherode (sometimes not developed) bilobed, lobes sessile, lanceolate-elliptic, -1 mm long, asym- metric, attached near the base and parallel to filament, connective slightly elongate. Lateral staminodes -5 mm long, 84 SMITHSONIAN CONTRIBUTIONS TO BOTANY MAP I.--Distribution of Aneilema obbiadense Chiovenda (solid circle with star), A. pusillwn Chiovenda subsp. pusillwn (solid squares). A . pusillwn subsp. gypsophilwn Faden, new subspecies (open circles), A . pusillwn subsp. thulinii Faden, new subspecies (open squares), and A. purilllun subsp. variabile Faden. new subspecies (solid circles). yellow, antherodes bilobed, lobes stipitate, obovate-cuneate. Lateral stamens with filaments S-shaped, -10 mm long, glabrous, anthers (in bud) ovate. Medial stamen with anther saddle-shaped, subequal to the lateral anthers (in bud). Ovary sessile, densely and apparently uniformly puberulous, apex acute, tapering into the style, dorsal locule 1-ovulate, ventral locules each 2-ovulate, style 11-12 mm long, stigma small. Submature capsule stipitate, fusiform, tapering at both ends, slightly falcate, dehiscent, bivalved, trilocular, 15 mm long, -3.5 mm wide, smmineus, lustrous, uniformly puberulous, apex rostrate, dorsal locule 1-seeded, ventral locules each 2-seeded, cells of the capsule wall transversely elongate, in regular files. Immature seed (only one seen) transversely elliptic 4 . 7 x 1.5 mm, hilum much shorter than the seed, raised within a groove, groove cut off apically or basally (but not both) by a ridge, testa apparently smooth. HABITAT.-Stabilized, rocky sand dunes, and grassy, fixed dunes (with limestone exposures) dominated by prostrate bushes; -50 m. FLOWERING.-May. DISTRIBUTION.rentrd Somali Republic (Map 8). CHROMOSOME NUMBER.-Unknown. SPECIMENS EEN.-SOMALI REPUBLIC. 18 Km SW of Obbia on road to Wisil, 5"17'N, 48"22'E, Gillett et al. 22229 @A, K, US); Tobunghb, Puccioni & Stefunini 592 [646] (FI'). DISCUSSION Some of the dimensions given here differ from those in Chiovenda's type description. Several of his measurements, e.g., leaf width and cincinnus bract length, appear to have been made inaccurately. Others seem to have come from a soaked bud and are therefore much too small, e.g., petal length and width. On the other hand, his measurement for sepal width is obviously too great, the sepals actually being much narrower than long. In the above description, Chiovenda's dimensions are given only when they accord with those that I have obtained or when they are the only dimensions available. Based on the seeds in the capsule, the ovary should have one ovule in the dorsal locule and two in each ventral locule. I do not believe Chiovenda's description of the ovary as bilocular with each locule triovulate. The most distinctive feature of this species is the very wide spacing of the bracteoles on the cincinni, greater than in any NUMBER 76 85 other species in the genus. This character combined with the unusually long cincinnus peduncles gives the inflorescence a most distinctive facies. The disarticulating shoots are also unusual in the genus. The rhizome and roots in A. obbiadense are very similar to those of A. brenanianum. That species differs in having much longer leaves with very long sheaths, the flowering shoots not disarticulating, the basal part persistent, a denser inflorescence with shorter cincinnus peduncles and internodes, the medial staminode with stipitate, reniform antherode lobes, the medial anther ellipsoid or ovoid with a very convex connective, the ovary glabrous with more numerous ovules per ventral locule. The floral differences show that the similarity in the subterra- nean parts of these two species is due to convergence and is not indicative of a close relationship. The true relationships of A. obbiadense are clearly with A. grandibracteolatum and A. longicapsa (see discussion under section Somaliensia). 4. Aneilema longicapsa Faden, new species Aneilema obbiadense var. angustifoliwn Chiovenda [(Langustifolia??l, P1. Novae Aethiopia, 10, 1928; Fl. Somala, 1:316, 1929. [Type: Somali Republic [Somalia centrale on label], Sultanato di Obbia, tra Scermarca Hassam e Tobungab, 4 May 1924, Puccioni & Sfefunini 579 [6M] F)]. Herbae perennes foliis spiraliter dispositis, laminis linearo- lanceolatis (ad lanceolato-ellipticas), 2-8 cm longis, 0.35- 0.8(-1.9) cm latis. Inflorescentiae thyrsi ovoidei, 3.5-7.5 longi, 3-6.5(-9) cm lati, cincinnis (1 1-)13-36 compositi. Pedunculi cincinnorum 10-20 mm longi, bracteis longiores. Bracteolae 1.5-7.5 mm semotae. Sepala 3-5 mm longa, puberula. Petala 7-8.5 mm longa, 6.5-8 mm lata. Ovarium dense eglanduloso-puberulum, loculo dorsali 1-ovulato vel casso, loculis ventralibus 2-ovulatis. Capsulae stipitatae, rostratae, (12-)14- 17 mm longae, loculis ventralibus plerum- que uterque 2-seminalibus. Semina transverse elliptica ad oblongo-elliptica, 3.75-4.55 mm longa, 1.55-1.7 mm lata. TYPE.-Somali Republic, Central Somalia, 17 km S of Mugakori, 3?57?N, 46?112, 6 June 1981, Gillett & Beckett 23289 (K, holotype; EA, US, isotypes). Perennial herbs (habit type IVA; Figure 3). Roots k tuberous, 3-4 mm thick at base, then abruptly narrowed and 1.5-2 mm thick for rest of their length (possibly broadened distally in Gillett et al. 22609-US), to 28 cm long or longer. Aerial shoots annual, erect to ascending, sparsely branched, disarticu- lating at base when drying, to -30 cm tall. Internodes 1-4.5 cm long, puberulous. Leaves spirally arranged, sheaths often split to the base or nearly so, to 0.5 cm long, puberulous, sparsely ciliate (with uniseriate hairs) or ciliolate with uniseriate hairs and/or hook-hairs at the apex, lamina sessile (to shortly petiolate in cultivation), linear-lanceolate (to lanceolate- elliptic), 2-8 cm long, 0.35-0.8(-1.9) cm wide, apex acuminate to acute, base cuneate (to rounded), margins planar, scabrid with prickle-hairs near the apex and hook-hairs below, both surfaces scabrous, puberulous, lacking uniseriate hairs. Inflorescences thyrses, terminal, moderately dense to moder- ately lax, ovoid, 3.5-7.5 cm long, 3-6.5(-9) cm wide (to 10 cm wide in cultivation), with (1 1-)13-36 cincinni, mostly alternate, with the lower several subverticillate, ascending to patent. Peduncles 2-3.5(-5.5 in cultivation) cm long, puberu- lous. Inflorescence bract supramedial, usually foliaceous (occasionally bract-like). Distance between inflorescence bract and lowest cincinnus 0.5-1.3(-1.7) cm. Inflorescence axis puberulous with uniform hook-hairs, green tinged with blue. Cincinni to 5 cm long and 8-flowered (to 6 cm long and 9-flowered in cultivation). Cincinnus bracts scarious, ascend- ing to reflexed, lanceolate to lanceolate-oblong or ovate- oblong, 2-4.5 mm long, not evidently glandular, glabrous or subglabrous. Cincinnus peduncles relatively uniform within the inflorescence (or the uppermost the longest), greatly exceeding the cincinnus bracts, 10-20 mm long, sometimes blue-tinged, puberulous. Cincinnus axis puberulous with uniform-length hook-hairs, sometimes blue-tinged. Bracteoles spaced 1.5-7.5 mm apart, scarious, eccentrically cup-shaped, perfoliate, (1.5-)1.9-3 mm long, 0.6-1.6 mm high, glandular subapi- cally, sparsely puberulous basally or glabrous, margin usually erose basally. Flowers perfect and staminate (Plate 2h,l), odorless, 15-18 mm wide. Pedicels 3.5-6.5 mm long in flower, to 9 mm long in fruit, ascending in flower, recurved in fruit at the base 120?-2250(mostly -180?), green, glabrous or with a few hairs at the apex (occasionally puberulous on one side). Sepals convexo-concave, green, tinged with reddish purple basally, glandular near the apex but not prominently so, puberulous with uniform hook-hairs; medial sepal with a strongly cucullate apex, ovate to ovate-elliptic, 3-4.5 mm long, 2.1-2.5 mm wide; lateral sepals oblong-elliptic to ovate-elliptic, 3-5 mm long, 2-2.2 mm wide. Paired petals not overlapping, 7-8.5 mm long, 6.5-8 mm wide, limb ovate, 5.5-7 mm long, lilac (RHS color: 77C-D, Gillett & Beckett 23289), apex rounded, cucullate, base nearly symmetric, broadly cuneate; claw 1.5-2.5 mm long, glabrous. Medial petal lanceolate to lanceolate-elliptic or ovate-elliptic, 5-7 mm long, 2.5-3.3 mm wide, hyaline white faintly tinged with lilac in center, margins colorless. All filament bases free. Medial staminode 1.5-2 mm long, yellow, filament -0.5-1 mm long, broadened apically, antherode bilobed, lobes sessile, oblong-elliptic to obovate, obliquely attached to filament, -1-1.4 mm long, connective slightly to greatly elongate. Lateral staminodes yellow, filaments 4.5-6 mm long, antherodes bilobed, lobes stipitate, transversely elliptic to obovate, -0.5 mm long, stalks 0.5-0.7 mm long, meeting filament at an acute angle, connective not elongate. Lateral stamens with filaments divergent, distinctly longer in staminate than perfect flowers but of similar curvature, 8.5-12 mm long, S-shaped, glabrous, white at base, pale pink above, anthers facing towards the flower base or towards the floral midplane and slightly forwards, ovate, 0.6-1.8 mm long, connective whitish, sutures blue-black, pollen dirty yellow. Medial stamen with filament 5-6.5 mm 86 SMITHSONIAN CONTRIBUTIONS TO BOTANY long, arcuate ascending, white basally, pale pink medially, yellow apically, anther saddle-shaped, ovate, 1-1.8 mm long, connective yellow, pollen golden yellow (distinctly discolorous from lateral anther pollen). Ovary sessile, oblong-elliptic to elliptic, 2.5-3 mm long, -1 mm wide, densely and uniformly covered with patent, colorless, eglandular hairs, apex acute, tapered into style, dorsal locule prominent or not, l-ovulate or empty, ventral locules each 2-ovulate; style 9-12.5 mm long, arcuatedecurved then recurved, lilac, tapered at apex, stigma very slightly enlarged over the style apex, lilac. Capsules stipitate, oblanceolate to oblong-oblanceolate, tapering at both ends, slightly curved, slightly constricted between the seeds, dehiscent, bivalved, bilocular (sometimes trilocular?), (12-)14-17 mm long (including stipe and beak), -2.5-3 mm wide, chocolate brown or stramineous, lustrous, uniformly glandular-puberulous, apex rostrate, valves spread- ing less than go?, cells of the capsule wall transversely elongate, in regular files. Seeds transversely elliptic to transversely oblong-elliptic, rounded to somewhat truncate at both ends (in 1-seeded locules) or rounded to somewhat truncate at one end and truncate at other end (in 2-seeded locules), 3.75-4.55 mm long, 1.55-1.7 mm wide, hilum dark brown, straight or slightly curved, much shorter than the seed, raised within a deep groove that is cut off at both ends by ridges, with margins of the groove crenate to lobed, testa light brown, scrobiculate, with the shallow pits and short grooves in more or less radiating lines from the embryotega, farinose granules lacking, embryotega tan. HABITAT.-Acacia-Commiphora bushland on red, sandy soil, sometimes with limestone outcrops, also ?elevated meadows?; 150-300 m. FLowERING.--Flowering specimens have been seen from May, June, and November. In greenhouse cultivation the flowers fade by mid morning. CHROMOSOME NUMBER.-2n = 64. DISTRIBUTION. Sudan to N Kenya and S Somali Republic S to Uganda and N and SE Tanzania (Map 10). SPECIMENS SEEN.-SUDAN. EAST EQUATORIA: 10 km S of Mongalla, by Juba Road, Lock 811275 (K), cultivated Smith- sonian Institution (US). SOMALI REPUBLIC. BALAD: Hilweyne area along Balad- Mogadishu road, Thulin & Warfa 5280 (UPS). UGANDA. BUNYORO: 23 mi N of Butiaba turnoff on Butiaba-Murchison Falls National Park road, Faden et al. 6911068 (BR, EA). KENYA. BARINGO: Chemolingot borehole area, Achepto- pokwa River, -0'58'N. 35'572, Timberlake 344 (EA); approximately same locality, Timberlake 156a (EA). EMBU: Thuchi R. crossing on Embu-Meru road, 0"25'S, 37'52'E, Gillett & Mathew 19068 (EA, K) & 19073 (EA); by Tana R., -0'28's. 37'55'E, Gillett & Mathew 19090 (BR, EA, K). FORT HALL: 31 mi Embu-Kangonde, -1 mi S of suspension bridge, Oo50'S, 37'402, Faden et al. 69/395 (EA, K, MO, US). KITUI: Thika-Garissa road, 6 km Nguni-Enziu, Oo50'S, 38'17'E, Faden & Faden 741751 (BR, EA, FT, K, MO, PRE, US, WAG): Thika-Garissa road, 6.2 km Ukazzi-Garissa, Oo48'S, 38'35'E, Faden & Faden 74/753 (B, BM, C, DSM, EA, G, K, M, MO, P, UPS, US); Mutomo Hill Plant Sanctuary, 1'519, 38013'EE, Gillett 18604 (EA, K); Nairobi-Garissa road, -5 km E of Ukazzi, -0?48'S,38'37'E, Gillett & Gachathi 20488 (BR, EA, K, MO, PRE); Tsavo National Park E, Ithumba, Hucks 221 (EA, K, Tsavo Research Centre); Mutomo Hill, Mathew & Hanid 6061 (EA, K). K I T ~ A N A RIVER: Katumba Hill, 92 km on the Garissa-Nairobi road, -0'443, 38'483, Gilbert & Thulin 1730 (EA, F). MACHAKOS: Near Hunter's Lodge, first rocky hill with baobabs on Nairobi-Mombasa road, 2'139, 37'433, Faden & Faden 771310 (EA, F, K, US); Tsavo National Park W, Chyulu Hills foothills, -2'45'S, 37'583, Faden et al. 69/61 7 (BR, EA, K, MO); Tsavo National Park W, 3 km WNW of Kyulu Station, 2'56'S, 38?23'E, Gillett 17271 (BR, EA, K); "Bushwackers Safari Camp" (Masaleni), 2'199, 98 SMITHSONIAN CONTRIBUTIONS TO BOTANY 38?07?E, Napper & Kanuri 2087 (EA, K, LISC, WAG); Tsavo National Park, S of Mtito Andei, Strid 2406 (EA); first baobab rock outcrop few km beyond Simba, Verdcourt 5234 (MO). m u : Meru National Park, S of Point 40, new road crossing Kiolu River, Ament & Magogo 263 (EA). NORTHERN FRON- TIER: Dandu, 3?26?N, 39?54?E, Gillett 13132 (B, BR, EA, K-2 sheets, PRE, S). TErrA: Tsavo National Park E, Irima Rock above Irima Water Hole, Faden & Faden 721284 (B, EA, FT, K, MO) & 741237 (BR, EA, K, MO, PRE, US, WAG) & 771342 (EA, F, US); Tsavo National Park E, Simba Hill, 3?22?S, 38?3575 Faden & Faden 741231 (MO) & 741231A (BR, EA, K, MO, PRE, US, Z); 1.5 km from Nairobi-Mombasa road on E turnoff to Voi, 3024?S,38?352, Faden & Faden 741246 (BR, C, EA, K, MO, US); 2.8 km on Voi-Taveta road from Nairobi-Mombasa road junction, 3?25?S, 38?322, Faden & Faden 741274 @A, MO); 7.3 km on Voi-Taveta road from Nairobi-Mombasa road junction, 3?263, 38?302, Faden & Faden 741529 (EA, K, MO, PRE, US); Tsavo National Park E, Voi Gate, Gillett 17204 (BR, EA, K); Tsavo National Park E, Mudanda Rock, Hucks & Hucks 15 (EA); Tsavo National Park E, Irima, 3?17?S, 38?322, H u c h 669 (Tsavo Research Centre) (mixture with A. rendlei); Tsavo National Park E, Worsessa, -3?20?S, 38?332, Hucks 690 @A, Tsavo Research Centre); Tsavo East National Park, Voi Gate camp site, Msaj?ri 256 (EA); Ndi Mountain, opposite Manga Mountain, Verdcourt & Polhill 2708 (BR, EA, K, PRE). TANZANIA. KILWA: Selous Game Reserve, Kingupira, 8?283, 38?332, Vollesen 1964 (EA). MBULU: Lake Manyara National Park, Bagayo River, Greenway & Kanuri 11871 (BR, EA, K, PRE). SHINYANGA: Shinyanga, Bau 368 (BR, K). DISCUSSION The unambiguous typification of Aneilema petersii is possible only because of the fortuitous survival of the type in the Berlin Herbarium. Despite its detail, the description of Lamprodithyros petersii by Hasskarl (1864a) would in itself have been inadequate to distinguish A. petersii from the closely related A. indehiscens, which also occurs in Mozambique. The smaller packet on the Peters? type contains a capsule of A. schlechteri. The rarity of that species-it is known from only four collections (see Faden, 1984)-suggests that the capsule probably came from the type (Schlechter I 1 748), because that is the only collection of A. schlechteri known to have been in the Berlin Herbarium. Schlechter?s collection was made in 1897 and, therefore, it could not have been the basis for any part of Hasskarl?s description of Lamprodithyros petersii. Both Aneilema tetraspermum K. Schumann and A. sacleuxii Hua are clearly synonyms of A. petersii subsp. petersii. The three specimens seen of Volkens 175, isosyntypes of A. tetraspermum, all have ?Aneilema leptasperma K. Schum.? on the label, a name Schumann never published. Clarke has written on the Kew sheet that ?K. Schumann.. .altered his new specific name before publication, but forgot to alter his old specific name on the set distributed to Kew.? Because all the original material of A. tetraspermum in the Berlin Herbarium was destroyed, the Kew sheet of Volkens 175 is here selected as the lectotype. Hua (1895) did not cite any specimen when he described A. saclew*i. It is clear that Sacleux 1142-the ?4? can easily be misread as a ?9?-is the specimen on which the name was based because it matches Hua?s description and apparently is the only collection of this species made by Sacleux. Aneilema chrysanthum K. Schumann, the type of which did not survive in Berlin, is probably a synonym of A. petersii subsp. petersii. The type locality and certain characters in Schumann?s description of A. chrysanthum agree with A. petersii, especially the narrow leaves, lax ?panicle? with well-spaced branches, short pedicels, stipitate, subplano- convex, lustrous, mother-of-pearl colored, strongly hump- backed, obtuse capsules, and relatively large seeds. Among the characters that disagree, the single staminode reported is not found in any Aneilema, so this was probably inaccurately observed; the bilocular capsule is not consistent with the description of the capsule as humpbacked, which strongly suggests a trilocular capsule; and the yellow petal color may just have been erroneous. The only yellow-flowered species from the same general area, A. nyasense, occurs in forest, not in ?sandy bush,? as recorded for A. chrysanthum. It further differs from the description of A. chrysanthum in so many significant ways, that it is unlikely to have been the same species. Tweedie 3365 and Allen 94 have been included in this species with some doubt because both collections are of plants that have just begun to flower and lack capsules. The former has been included in this species because of the flower color, recorded by the collector as pale mauve, and the form of the antherodes, as determined from a dissected bud. Allen 94 agrees with A. petersii in more attributes than it does with A. indehiscens, the only other possibility, and it was collected within the known range of the former but not the latter. This specimen has therefore been treated as A. petersii. The subspecific determinations of the four collections from the Kenya-Ethiopia border region are also somewhat doubtful. Bully & Smith B14873, Cufodontis 705, and Gillett 14091 are morphologically similar to one another. Because preserved flowers of Gillett 14091 from the Kew Herbarium are in the size range of subsp. petersii (sepals 3.9-4.9 mm long; lateral stamen filaments 9 mm long; style 7.7 mm long) but are too large for subsp. pallidflorum, all three of these collections have been treated as subsp. petersii. The most striking features of the fourth collection, Gillett 13132, that immediately separate it from the other three collections are its very large leaves (to -13.5 x 4.5 cm) and capsules (to 9 x 5.5 mm), the maximum dimensions for both of these organs in A. petersii. While capsule size shows considerable variation within both subspecies, the leaf width of Gillett 13132 is approached only by specimens of subsp. pallidiflorum. The Gillett collection also agrees with this subspecies in flowering sepal length (3-3.2 mm), as measured in preserved flowers from Kew. Only the lack of perforating NUMBER 76 99 inflorescence-shoots would distinguish this collection from typical subsp. pallidiflorum, but perhaps this is merely a function of age, as has been noted in other collections of this subspecies, e.g., Ament & Glover 263. Overall, Gillett 13132 much better conforms with subsp. pallidiflorum than with subsp. petersii. The occurrence of both subspecies of A. petersii in the Kenya-Ethiopia border region at first seems unlikely in view of their apparently different precipitation requirements and nearly total geographic isolation in other areas. However, closer study reveals that Gillett 13132, treated as subsp. pallidiflorum, is geographically separate from the other three collections that were considered subsp. petersii, and that its locality, Dandu, is situated in a zone of lower mean annual rainfall (less than 255 mm) than Moyale (687 mm), where Gillett 14091 was collected (National Atlas of Kenya, 1970). Thus the pattern of distribution for these populations of the two subspecies conforms with that found in other parts of their ranges, at least with regard to rainfall. The two Mafia Island collections (Greenway 5236; Wallace 810) have unusually short bracteoles and narrow capsules. They are considered to represent a minor geographic variant unworthy of taxonomic recognition. The recent collection from Somalia (Thulin & Warfa 5280) is unusual in its inflorescences composed of only two to three cinicnni; bracteoles all cup-shaped and perfoliate, with at most a single uniseriate hair; and capsule locules one-seeded. Further collections are needed in order to determine whether this is an atypical specimen, a local variant, or even possibly an undescribed species. If this is A. pefersii, then it is closer to subsp. pallidiflorum because of its short, perforating inflores- cence-shoots and short sepals and style. The great variation in vegetative morphology in A. petersii is due to the Occurrence of both annuals and perennials in the species. Subspecies petersii is always perennial. It typically has a definite base and produces ascending shoots. Occasionally it may also form longer, decumbent shoots that give rise to new plants. The main shoots are thickened at the base, at least during the dry season. Subspecies pallidiforum is consistently annual despite some collectors? notes (including the writer?s on Faden & Faden 721284) to the contrary. Although all plants in some popula- tions are very reduced and clearly annual, some from other populations may have decumbent shoots with their habits impossible to determine by casual inspection. Repeated field observations indicate that even these decumbent plants live less than a year and flower only once before dying. Their phenology is discussed below. Distichously arranged leaves have been noted in A. petersii only in two populations, both of subsp. pallidiflorum (Faden et al. 691617 and Faden et al. 6911068). All the lateral shoots of the latter have this arrangement, at least in plants raised from seeds. The leaves of subsp. petersii are typically narrower (to 3.2 cm wide) than those of subsp. pallidiflorum (to 4.5 cm wide). They are also proportionally narrower, never being ovate. Subcordate- to cordate-amplexicaul leaves are present only in subsp. pallidiflorum. The presence of numerous, short, lateral, inflorescence- bearing shoots in subsp. pallidiflorum but not in subsp. petersii is concordant with the former?s annual habit, which requires greater emphasis on seed production. This is accomplished in subsp. pallidiflorum partly through an increased number of inflorescences and flowers and partly through self-pollination. The short shoots usually emerge from the mouths of the sheaths of the upper leaves but perforate those of the lower leaves. The inflorescence bract of the terminal inflorescence of the main shoot is generally longer in subsp. pallidiflorum than in the typical subspecies. In populations of the former in the Vol area of Kenya the inflorescence bracts are borne close to the inflorescences and are held erect. They thus form a dark green background for the white flowers, increasing the lattre?s visibility (Plate 20). The size ranges of floral parts are greater in A. petersii than in any other species of section Lamprodithyros. This is duein large measure to the reduced flower size in subsp.pallidiflo- rum, an attribute commonly associated with self-pollination in Aneilema. The last flowers produced in plants of this subspecies just prior to their death are particularly small Flowers of the two subspecies differ consistently in size and petal color. The most reliable distinctions are included in the key. Differences in the capsules and seeds between the two subspecies are largely qualitative. Their reliability is uncenain due to insufficient material from some critical areas such as Mozambique (subsp. petersii), northern Tanzania (subsp. pallidiflorum), and the Kenya-Ethiopia border region (both subspecies). On the basis of available material, the capsules of subsp. pallidiflorum are more obovate (versus subquadrate) more deeply emarginate apically and browner (versus grayer) than those of subsp. petersii. The ventral locule seeds of subsp. pallidiflorum are regularly blackish farinose, while those of subsp. petersii often complctcly lack farinose granules. The apical ventral seeds are usually more pointed in the former. The difference in flowering phenology between the two subspecies is striking and is largely due to the rainfall distribution patterns in the regions in which they occur (Jackson, 1961; National Atlas of Kenya, 1970). Throughout the range of subsp. petersii there is no consistently dry month Hence it may be expected in flower in every month in some part of its range. The lack of flowering specimens from December is almost certainly an artifact. Faden et al. 72138, collectcdon 10 January, has mature fruits and must have been flowering in December. For subsp. pallidiflorum, the flowering phenology is well understood only in eastern Kcnya. There are two distinct rainy seasons, March to May and October to December. Observa- tions in the Tsavo area indicate that the plants come up with the October to December rains but do not flower then. Many of them survive the following dry season and flower during the 100 SMITHSOMAN CONTRIBUTIONS TO BOTANY March to May rains. All of them die during the subsequent dry season. In the Ugandan locality the rainy season spans the coincident dry months of the Kenyan and Tanzanian sites, so it is not surprising that the single collection from Uganda was flowering at a very different time of year from the other collections. A single exception to this pattern has been recorded. In 1977, the rains in Kenya, which should have ended in December 1976, continued into January. Consequently, a few plants of subsp. pallidiflorum in Tsavo National Park were found in flower in February 1977 in what would normally have been the height of the dry season. Five collections made since my original treatment of this species (Faden, 1975) have greatly extended the known range of subsp. pallidiflorum: to the Sudan (Luck 81/275), to the Somali Republic (Thulin & Warfa 5280), to western Kenya (Timberluke 344 and 156a), and to southern Tanzania (Vollesen 1964). Aneilema petersii is a highly distinctive species. It can be separated from all other taxa of section Lamprodithyros by its mostly non-perfoliate bracteoles, some or all with long, uniseriate hairs basally, crossed lateral stamen filaments not retained by the medial petal, longly stipitate, dehiscent, broad capsules with a humpbacked, deciduous, dorsal valve, and strongly dimorphic seeds, the dorsal locule seed particularly large (largest seed in the section). Both subspecies have all of these attributes and are easily separated in the field using the key characters, many of which, unfortunately, do not preserve well in herbarium specimens. Aneilema petersii appears to be most closely related to A. indehiscens, the only other species in section Lamprodithyros (except for A. rebrinum) known to lack a stamen retention mechanism. Aneilema indehiscens is also similar in its bracteoles sometimes not perfoliate and frequently with one or more uniseriate hairs on the margin, and in the antherodes of all three staminodes of similar form (to one another). For differences between these species, see the discussion of A. indehiscens and the key to the species. 2. Aneilema indehiscens Faden Aneilema indehiscens Faden, Bothalia, 15:96,1984.-Obermeyer and Faden in Leistner, Fl. Southem Africa, 4(2):40, fig. 9-2a,b. 1985. [Type: Kenya, Tana River District, Garsen-Malindi road, 1.5 km towards Malindi from tumoff to Oda. 2032'S, 4O007'30"E, 22-24 July 1974, Faden & Faden 7411184 (US, holotype; BR, EA, R, K, MO, PRE, WAG, isotypes).] Aneilemo pefersii (Hasskarl) C.B. Clarke in De Candolle, Monogr. Phan., 3225, 1881 [pro Kirk s.n.1; in Thiselton-Dyer, F1. Trop. Africa, 8:70, 1901 [pro Kirk s.n.1. Aneilema dregeanwn sensu Compton, Fl. Swaziland, 33. 1966. p.p. [non Kunth. 18431. Perennial herbs (habit type IIA3, Figures 2, 48a). Roots fibrous. Vegetative shoots sparsely branched, trailing and often looping along the ground, occasionally rooting at the nodes, sometimes straggling through shrubs, to 3 m long (or longer?), flowering shoots produced irregularly, unbranched or sparsely branched, erect to ascending, to -60 cm tall (reaching a greater height when straggling through shrubs). Internodes (1-)3-15 (-19) cm long, to 5 mm thick, glabrous or very sparsely puberulous and glabrescent. Leaves spirally arranged, sheaths 0.5-2 cm long, puberulous and sometimes sparsely pilose, ciliate at the apex, laminae shortly petiolate, gradually reduced towards the terminal inflorescence, narrowly lanceolate to lanceolate-elliptic, lanceolate-ovate, or ovate-elliptic, rarely ovate, (2.5-)3-lo(-13) cm long, (0.7-)1-2.5(-3.5) cm wide, apex acuminate or acute, margin frequently undulate, scabrid, also ciliate at least basally and on the petiolate, both surfaces lustrous, puberulous, rarely densely pilose-puberulous, abaxial surface with with longer, more numerous, uniseriate, hairs, veins pale on the adaxial surface. Inflorescences thyrses (Figure 48b), terminal and frequently axillary from the upper leaves on the flowering shoots, lax to moderately dense, ovoid to broadly ovoid, (2-)2.5-5(-8) cm long, (1.5-)2-5(-7) cm wide, with (1-)3-9 cincinni, ascend- ing (the lower sometimes patent), mostly alternate (frequently some subopposite, rarely some subverticillate). Peduncles 2-5(-8) cm long, puberulous. Inflorescence bract supramedial to medial, herbaceous and foliaceous or membranous and bract-like. Inflorescence axis slightly zigzag, puberulous. Cincinni to 7.5 cm long and 27-flowered. Cincinnus bracts membranous, appressed to the cincinnus peduncles or patent, lanceolate-elliptic to linear-lanceolate or ovate, (1.5-)2-3(-5) mm long, glandular near the apex, glabrous to puberulous. Cincinnus peduncles relatively uniform within the inflores- cence or decreasing in the upper cincinni, exceeding the cincinnus bracts, 3-1 1(-12.5) mm long, puberulous. Cincin- nus axes puberulous. Bracteoles (Figure 48c) attached 1-3.5 (-4.5) mm apart, k herbaceous, eccentrically cup-shaped, usually perfoliate, 1.3-2.6 mm long, to 1 mm high, with a prominent subapical gland, puberulous at least basally or medially, frequently also with 1 -several, long, unisenate hairs on or near the fused edge, margin sometimes slightly thickened (glandular?) near the fused edge. Flowers perfect and staminate, odorless, (9-)13-17.5 mm FIOURE 48.-Aneilema indehiscens Faden. A. indehiscenr subsp. indehiscenr (u-g,bl): a, habit; b, inflorescence; c, bracteole; d, perfea flower, front view; e, perfect flower, side view;$ lateral petal; g, antherodes of staminodes: medial staminode (right) and lateral staminode (left); i , androeurn, top view, medial staminode omitted, gynoecium removed; j , ovaly and base of style, side view; k, capsule attached to uncinnus, side view; 1, dorsal capsule valve, dorsal view (Q-c, from Faden & Faden 741380; d-g,CI, from Faden & Faden 72172); A. indehircens subsp. lilacinurn Faden (h, m q ) : h, antherodes of staminodes: medial staminode (right) and lateral staminode (left); m, capsule, side view; n, dorsal capsule valve, dorsal view; 0, dorsal locule seed, dorsal view; p , ventral locule seed, ventral view; q, ventral locule seed, dorsal view (h, f r m Faden & Faden 741208; m q , from Faden & Faden 741202). (Bars = 40 mm for a; 10 mm for b; 2 mm for d-f,k,m; 1 mm for c,g-t',l,n; 0.67 mm forj; 0.5 mm for 0-q.) NUMBER 76 101 102 SMITHSOMAN CONTRIBUTIONS TO BOTANY wide (Figure 48d,e; Plate 3ab). Pedicels 3.8-6(-8) mm long in flower, to 10 mm long in fruit, erect or ascending in flower, k uniformly recurved in fruit, usually -180?, persistent, puberulous. Sepals glandular near the apex, puberulous; medial sepal ovate or ovate-elliptic to lanceolate-elliptic or lanceolate, 2.4-4.3(-4.9) mm long, 1.8-2.6 mm wide, subapical gland very prominent, k distinctly bilobed; lateral sepals broadly elliptic or ovate-elliptic to ovate-orbicular, oblong-elliptic or obovate-elliptic, 2.6-4.3(-4.6) mm long, 2.2-3.2 mm wide, subapical gland prominent or not, usually unlobed (rarely bilobed). Paired petals 7.3-9.5 mm long, 6-8.5 mm wide (Figure 48f). limb broadly ovate to ovate-deltate, 4.3-6.5 mm long, white to very pale lilac (RHS colors: 76C, Fuden & Fuden 741202; 84D, Faden C? Fuden 74/208), apex rounded or occasionally obtuse, claw 2-3.5 mm long, white or whitish, glabrous. Medial petal cup-shaped, obovate (occasionally ovate or suborbicular), broadest at the margin (viewed from the apex), 6-8 mm long, 4-6(-7.5) mm wide, 3-5 mm deep, concolorous with the limbs of the paired petals (Figure 23d-f). Stamen filaments shortly fused basally and also shortly fused with the lateral staminode filaments (Figures 26u, 48i). Medial staminode filament (1.5-)2.7-4.6 mm long, yellow above the white or whitish base, antherode (rarely absent) bilobed, yellow, lobes sessile to shortly stipitate, obovate-cuneate to sickle-shaped and decurved, 0.6-1.2 mm long, connective usually slightly elongate (Figure 48g, right, h, right). Lateral staminodes with filaments 4-5.6 mm long, white or whitish, shading to yellow at the apex, antherodes bilobed, yellow, generally similar in size and form to that of the medial staminode (Figure 48g, left, h, left). Lateral stamens with filaments usually k parallel or slightly divergent for their entire length, or sometimes convergent apically (Figures 26u, 48i; Plate 3u,b), 7.7-8.5 mm long, gently S-shaped, geniculate below the middle, glabrous, anthers ovate to ovate-elliptic or occasionally elliptic to lanceolate-elliptic, 0.65- 1.3 mm long, 0.6-1 mm wide, pollen yellow to orange or dirty white. Medial stamen with filament 5-7 mm long, anther ovate to ovate- elliptic, saddle-shaped, 1.5-2.4 mm long, 1.2-1.7 mm wide, pollen yellow to orange-yellow, concolorous with the pollen of the lateral anthers or different in color. Ovary substipitate (Figure 48~). obovate to oblong (rarely subquadrate), 1.5-2.3 mm long, 1.2-1.6 mm wide, densely and uniformly covered with patent, glandular hairs (very rarely mixed with a few hook-hairs), apex truncate to slightly emarginate, dorsal locule prominent, subequal to the ventral locules or distinctly smaller than them, l-(or rarely 2-)ovulate, ventral locules each 2-(or very rarely 3-)ovulate; style 8-9.3 mm long, straight or gently arcuate-decurved for most of its length and strongly curved laterally out of the floral midplane (Plate 3u,b), stigma capitate, usually held in front of and lateral to the anthers. Capsules (Figure 48k-n) subsessile to stipitate, obovate- elliptic to obovate-oblong, oblong or oblanceolate, dehiscent or indehiscent, when dehiscent, bivalved (occasionally partially trivalved), trilocular, (4-)4.5-6(-6.8) mm long, (1.9-)2.3-3 (-3.4) mm wide, chestnut brown or mottled dark and light brown or gray-brown, lustrous, puberulous, apex emarginate, valves persistent (very rarely dorsal valve deciduous), dorsal valve with dorsal locule truncate to rounded apically or sometimes terminating in a narrow ridge and subequal to the ventral valve (Figure 481), dorsal locule prominent, l-seeded or, by abortion, empty (very rarely 2-seeded), ventral locules each 2- or, by abortion, l-seeded (very rarely 3-seeded); cells of the capsule wall transversely elongate. Seeds (Figure 48o-q; Plate 6b) elliptic, 2-2.9 mm long, 1.35-1.65(-1.9) mm wide (dorsal locule seed) or ovate to trapezoidal with apical seed obtuse to rounded apically and basal seed rounded to slightly angular basally, 1.5-2.2(-2.5) mm long, 1.3-1.8 mm wide (ventral locule seeds), 0.65-1 mm thick, hilum dark brown, in a very shallow groove or not in a groove, not extended onto apical and basal surfaces, testa usually orange-buff (rarely buff or orange-brown), very shallowly scrobiculate, with white farinose granules sparse to dense around the hilum, sparse around the embryotega and very sparse or lacking in the depressions. Key to the Subspecies of Aneilema indehiscens 1. Petals white; capsule usually chestnut brown . . . , . . . . A. i. subsp. indehiscens 1. Petals pale lilac; capsules gray-brown or mottled light and dark brown. 2. Capsules with dorsal locule rounded to truncate apically, not terminating in a ridge; pollen of medial and lateral anthers discolorous; Mozambique and Zimbabwe to South Africa. , . . . . . . . . . . . . . . A. i. subsp. lilacinurn 2. Capsules with ventral valve usually terminating in a ridge; pollen of all 3 anthers concolorous; western Kenya , . . . . . A. i. subsp. keniense, new subspecies 2a. Aneilema indehiscens Faden subsp. indehiscens FLowEwNG.-Flowering specimens have been seen from January, March-May, July and October. In the field the flowers open 0600-0630(--0645) hrs and fade 1100-1230 hrs. HABITAT.-Bushland and thickets of varied species compo- sition; sandy or clayey soils; usually in partial shade; CHROMOSOME NUMBER.-" = 26,2n = 52. -10-1050 m. DISTRIBUTION.-E Kenya and NE Tanzania (Map 11). NUMBER 76 103 M A P 1 1.-Distribution of Aneilema indehiscens Faden subsp. indehiscens (solid circles), A. indehiscens subsp. lilacinum Faden (open circles), and A. indehiscens subsp. koniense Faden, new subspecies (solid squares). 104 SMITHSONIAN CONTRIBUTIONS TO BOTANY SPECIMENS EEN.-KENYA. TANA RIVER: Garsen, 2'169, 40?07'E, Fuden & Fuden 7411066 (B, EA, FT, K, MO, P, PRE, UPS, US, WAG); 0.8 km towards Garsen from turnoff to Kibusu on Malindi-Garsen road, 2'21'S, 4Oo07'E, Fuden & Fuden 7411173 (BR, C, EA, K, LISC, MO, US); Garsen- Malindi road, 1.5 km towards Malindi from turnoff to Oda, 2'323, 4O007'30"E, Fuden & Fuden 7411184 (BR, EA, FT, K, MO, PRE, US, WAG). TEITA: Voi-Taveta road, just E of Tsavo National Park East entrance gate, cult. University of Chicago, A. Fuden 3179 (F, US); Tsavo National Park East, Buchuma (Bachuma) Gate, 3'40'S, 38056'EE, Fuden & Fuden 72/72 (EA; cultivated Missouri Botanical Garden: BR, EA, K, MO, PRE, US); 28 km towards Taveta on Voi-Taveta road from turnoff on Nairobi-Mombasa road, 3'303, 38'19'E, Fuden & Fuden 741489 (BR, EA, K, MO, US); 18 km towards Taveta on Voi-Taveta road from turnoff on Nairobi-Mombasa road, -3"30'S,38'24'E, Fuden & Fuden 741532 (EA, MO); 26 km towards Taveta on Voi-Taveta road from Nairobi- Mombasa road turnoff, -3'31'S, 380212, Fuden & Fuden 741536 (EA, K, MO); 11.7 km towards Mombasa past Maungu Station on Nairobi-Mombasa road, 3'37'S, 38'50'E, Fuden & Fuden 7411284 (MO); mile post Taveta 36Noi 36 on Taveta-Voi road, -3"25'S, 38'10'E, Fuden et ul. 69J318 (EA, FT, K, MO); Maungu Hills, 3'38'S, 38'442, Fuden et ul. 701158 (EA, K); 3 km E of Bum Railway Station, Gillett 19562 (EA, K, MO); Voi, Nupier 973 (EA, K). TANZANIA. BAGAMOYO: 4.5 km towards Mbwewe from crossing of Milgoji River on Korogwe-Dar es Salaam road, -5'57's. 38O122, Fuden & Fuden 741380 (MO cultivated Missouri Botanical Garden: DSM, EA, K, MO); Mbwewe, Fuulkner 4471 (K). LUSHOTO: Mazinde, Drummond & Hemsley 2337 (K). TANGA: Magunga Estate, Fuulkner 1160 (K). ZANZIBAR: Chumbuni, Vuughun 1851 (EA, K). 2b. Aneilema indehiscens Faden subsp. lilacinum Aneilema indehiscens Faden subsp. lilacinum Faden, Bothalia, 15:97, 1984.4bermeyer and Faden in Leistner, Fl. Southem Africa, 4(2):40, fig. 9-2a,b, 1985. [Qpe: South Africa,Natal, Ingwavuma-Ndumu road, 15.5 km towards Ingwavuma from junction with Ndumu-Maputa road, -27'065, 32'12'E, 16 Feb 1974, Faden & Faden 741202 (US, holotype; BR, EA, K, LISC, MO, NH, NU, PRE, WAG, isotypes).] HABITAT.4pen forest, woodland, thickets, lowveld bush, and edges of marshes; sandy or clayey soils; usually in partial shade; -10-550 m. FLowERING.-Flowering specimens have been seen from all months except August. In the field all flowers of one population were found fully open at 0700 hrs, while those of a second, nearby population began to fade at 1130 hrs. In cultivation the flowers of two populations were found to open about sunrise; they began to fade 4 and 6l/4 hrs later, respectively. CHROMOSOME NUMBER.-2n = 52. DISTRIBUTION.-^ Mozambique and S Zimbabwe to N Transvaal, Swaziland, and N Natal (Map 11). SPECIMWS SEEN.-ZIMBABWE. District unknown: Bank of Lundi River, Buyliss BS7216 (MO). MOZAMBIQUE. LOURENcO MARQUES: Costa do Sol, Burbosu 655 (LISC); without precise locality, Borle 364 (G); near Lourenco Marques town (Costa do Sol), Gomes & Sousu 3441 (BR, K-2 sheets); Maputo, Sep 1930, Gomes & Sousa s.n. (LISC); Ricatla, Acajou wood, Junod 493 (LISC, PRE-2 sheets); Inhaca Island, 23 mi E of Lourenco Marques, Mogg 27469 (K); Inhaca, Picada Estaab Hotel, Mouru et ul. 399 (US) ; Costa do Sol, Pedro 109 (LMA); Matolla, Quintas 64 (COI-2 sheets). MANICA E SOFALA: Mouth of River Melambe, Zambesi Delta, 8 Jul 1861, Kirk s.n. (K). SUL DO SAVE: Vila de J o a a o Belo, Chipenhe, Burbosu & Lemos 8431 (COI, K, LISC, LMA, PRE); Benguerua Isle, central ridge, Mogg 28886 (SRGH). SWAZILAND. HLATIKULU: Ingwavuma Poort, Compton 2861 0 (K, PRE). SOUTH AFRICA. TRANSVAAL: 2230 (Messina): 18 mi NE of Sibasa on road to Sambandou, Codd 6891 (K, PRE). 2431 (Acornhoek): Manyeleti Game Reserve, Albatross koppie (-CB), Bredenkump 1795 (PRE). NATAL: 2632 (Bela Vista): Ndumu Game Reserve, near main Rest Camp, -26'55'S, 32'19'E (-CD), Fuden & Fuden 741208 (K, MO, NH, NU, PRE, US); grid reference only (-CD), Moll 4152 (EA, K, NH, NU, PRE); Ndumu Game Reserve, Ndumu Hill (-CD), Outley C6 (PRE); Ndumu Game Reserve, E of Polwe Pan (-CD), Pooley 1399 (NU). 2732 (Ubombo): Jozini below dam, Makatini flats (-AC), Brenun 14232 (K); Ingwavuma-Ndumu road, 15.5 km towards Ingwavuma from junction with Ndumu-Maputa road, -27OO6'S,32O12'E (-AA), Fuden & Faden 741202 (BR, EA, K, LISC, MO, NH, NU, PRE, US, WAG); Ubombo Flats (-AB), Strey 10326 (EA, K, NH, NU, PRE); Lake Sibayi (-BCD), Vuhrmeijer 693 (PRE); Mahatini Flats, Vuhrmeijer & Tolken 192 (PRE). 2831 (Nkandla): Eshowe, above reservoir (-CD), Lawn 1289 (NH). 2c. Aneilema indehiscens subsp. keniense Faden, new subspecies Ab subspecie indehiscenti petalis pallide lilacinis, capsulis griseo-brunneis vel pallide brunneis et atro-brunneis irregu- lariter maculatis differt; ab subspecie lilucino capsulis valva dorsali apice plerumque in crista terminanti, polline omnium trium antherarum concoloro differt. TYPE.-Kenya, West Suk West Pokotl District, Kerio Valley, Tot-Sigor road, 4.8 km before crossing of Weiwei River, -1'26'N. 35'32%. 14 Mar 1977: collected sterile, cultivated at the University of Chicago, pressed 17 Jul 1977, Fuden & Fuden 77J788 ( U S , holotype; EA, F, K, isotypes). HABITAT.-Thickets and thicket edges, -1000- 1250 m. FLowERING.--Flowering May and October to November. CHROMOSOME NUMBER.-2n = 52. DISTRIBUTION.-Western Kenya (Map 11). SPECIMENS EEN.-KENYA. BARINGO: Samatian Island, NUMBER 76 105 Lake Baringo, Birnie 9 (EA); same locality, Lavrunos 17063 (EA). ELGEYO: Tambach-Tot road, 10 km N of junction with Kabarnet-Tambach road, Oo37'N, 35'36'E, Faden et al. 70/894A (EA, K). SOUTH KAVIRONDO: Lambwe Valley, Riamkanga, 0?42'3(Y'S, 34'122, Faden 69/1300A (EA). WEST SUK: Kerio Valley, Tot-Sigor road, 4.8 km before crossing of Weiwei River, 1'26'N, 35'32'E, Faden & Faden 771788 (EA, F, K, US). DISCUSSION Specimens of Aneilemu indehiscens have usually been treated as other species. Clarke (1881a, 1901) considered Kirk s.n. (subsp. lilacinurn) as A. petersii, and Compton (1976) cited Compton 28610, which is also subsp. lilacinum, as A. dregeanurn. Ross (1972) did not cite any specimen referable to A. indehiscens, but he must have included the specimens he saw of this species, which is quite frequent in northern Natal, in A. dregeanum and/or other species. The two collections of A. indehiscens from western Kenya that were part of mixed collections with A . recurvaturn, Faden 6911300A and Faden et al. 70/894A, were treated with some doubt as subsp. indehiscens in Faden (1975) but were omitted when that species was described (Faden, 1984). A third collection from the same region, Faden & Faden 77/788, has confirmed that these plants are consistently distinguishable from both described subspecies of A. indehiscens, and, consequently, subsp. keniense is described above. Subspecies keniense differs from subsp. indehiscens by its pale lilac flowers and paler, mottled capsules that are more distinctly stipitate. In these characters it agrees with subsp. lilacinum, from which it differs by its capsules with the dorsal valve usually terminating in a ridge, scarcity of uniseriate hairs on the bracteoles, relatively short pedicels (4.5-6.5 mm long), and concolorous pollen in all three anthers. The long-trailing or straggling habit of A . indehiscens is quite characteristic and apparently constant. I have noted it in all 13 populations of subsp. indehiscens, both populations of subsp. lilacinum, and one population of subsp. keniense observed in the field. It is also maintained in cultivation. Plants persist during the dry season as thick, leafless shoots. A habit similar to that of A. indehiscens has been noted elsewhere in the genus only in A. recurvaturn and in a single sterile plant believed to belong to A. hockii. The inflorescences are often grouped into larger functional units. Such synflorescences (Troll, 1961) consist of a terminal thyrse, a secondary thyrse from the axil of the inflorescence bract, and frequently additional thyrses axillary to the uppermost foliage leaf (on the main shoot) and/or the inflorescence bract of the secondary inflorescence. The bracteoles are usually perfoliate. Occasionally, however, one or more may be split to the base by the pedicels. Faulkner 1160 and Vaughan 1851 are unusual in having all of the bracteoles regularly non-perfoliate and thus similar to A. petersii in this character. The number of uniseriate hairs on the bracteoles and the percentage of the bracteoles within an inflorescence to have such hairs show much variation. In general, uniseriate hairs are much more numerous and occur on more bracteoles in subsp. lilacinum than in the other subspecies. The medial petal usually does not retain the lateral stamens when the flower opens. In one population (Faden & Faden 74/489, subsp. indehiscens), however, the stamens were held briefly but not in the usual manner. Only the filaments were contained in the petal; the anthers were shortly exserted. Furthermore, when the margins of the petal unfurled, the stamens rose slowly, indicating that they had not been under tension. In one population of subsp. indehiscens (Faden & Faden 741 184), the lateral staminodes often have small anther sacs that never contain any pollen. Because this small population may well consist of a single genotype, the presence of such an atavism in it may not be significant. On the other hand, the tendency for the medial antherode to be lacking, noted in the same population, is more important, because it has been found in all three Tana River District populations of subsp. indehiscens encountered, but not in any other populations of the species. The lateral stamen filaments usually appear parallel. Com- monly they are slightly bowed outward in the middle and converge again towards the apex (Figures 26a, 48i). Very rarely they may cross near the apex. The lateral anthers are generally held very close together. The presence of two ovules in the dorsal locule of the ovary has been noted in two populations, both of subsp. indehiscens. Of the five examples that have been found, four had had only the basal seed develop while the fifth had had both ovules produce seeds. In the latter case these seeds were very similar to those of the ventral locules in size and shape. Three seeds were found in one ventral locule of a single capsule of Faden & Faden 741208 (subsp. lilacinum). Hook-hairs have been noted on the ovary of a single flower of subsp. indehiscens (Faden & Faden 741380) and on two of 13 capsules (one hook-hair on each) from cultivated plants of the same collection. Ovarian hook-hairs are otherwise un- known in this species. Capsules of subsp. indehiscens are frequently indehiscent or only partly dehiscent along the lateral sutures. Occasionally the dorsal locule also splits for a short distance along the middorsal suture near the apex. Capsules of subsp. lilacinurn seem to always dehisce along the lateral sutures but never along the middorsal suture. A specimen of subsp. keniense (Faden et al. 70/894A) appears to have regularly dehiscent capsules, while cultivated plants of another collection (Faden & Faden 77/788) have indehiscent capsules. The extent of dehiscence is difficult to determine in pressed specimens because their preparation often produces or increases the split. Dorsal capsule valves are almost always persistent. In two 106 SMITHSONIAN CONTRIBUTIONS TO BOTANY collections of subsp. lilacinurn (Cornpton 26824 and Codd 6891) and one of subsp. keniense (Faden et al. 701894A), however, they appear to be regularly deciduous. The three subspecies are allopatric (Map 11). Although subsp. lilacinurn is geographically separated by almost 1500 km from the other two subspecies, few characters distinguish the three subspecies unequivocally. There is some evidence that the gap between subsp. indehiscens and subsp. lilacinurn may be an artifact of inadequate collecting. Most of the intervening area between the known distributions is considered poorly or only moderately collected (A.E.T.F.A.T., 1965, ?Map of the Extent of Florisitic Exploration of Africa South of the Sahara?). Furthermore, the two southernmost mainland collec- tions of subsp. indehiscens show some morphological tenden- cies towards subsp. lilacinurn: Faulkner 4471 has more numerous uniseriate hairs on the bracteoles than any other collection of the subspecies; and Faden & Faden 741380 (from the same population) sometimes has petals with a faint pink tinge, at least in cultivation, a character otherwise unknown in subsp. indehiscens. This pattern suggests possible clinal variation between these subspecies. Aneilema indehiscens is most closely related to A. petersii and A. recurvaturn. Indeed it may prove to have arisen as an allopolyploid hybrid between the twb. All three species have been found in the Kerio Valley of western Kenya and, at two of the four known localities for A. indehiscens subsp. keniense, it was found growing with A. recurvaturn. In eastern Kenya subsp. indehiscens occurs not far from both subspecies of A. petersii but is ecologically distinct from each. Among the 13 populations of A. indehiscens subsp. indehiscens and 22 of A. petersii (both subspecies) observed in the field, only one instance has been found of the two species growing together. That was in a disturbed ecotonal situation in Tana River District (see Faden, 1983b). Aneilema indehiscens may be distinguished from A. petersii by the former?s trailing habit, often falcate antherode lobes, usually slightly elongate connec- tives, lateral stamen filaments usually not crossing, narrower, often indehiscent capsules, and only slight dimorphism between dorsal and ventral locule seeds. From A. recurvatum, A. indehiscens differs by the latter?s less symmetric, puberulous bracteoles that lack marginal glands, larger flowers with puberulous sepals, more rounded paired petals, less divergent and less strongly curved lateral stamen filaments that are usually not retained by the medial petal, and generally larger capsules that are often indehiscent. lanceolato-ellipticis vel lanceolatis ad ovato-ellipticas, 2.5-7 (-8) cm longis, 0.5-2.2(-2.9) cm latis. Inflorescentiae thyrsi ovoidei ad late ovoideos, 2-4.5 cm longi, 2-6.5 cm lati, cincinnis (1-)2-7(-9) compositi. Bracteolae herbaceae, plus minusve symmetrice cupulatae perfoliatae, 1-2 mm longae, 0.5-1 -25 mm altae, glandibus parvis marginalibus praeditae. Pedicelli 4.5-10 mm longi, tempore fructigero recurvati plerumque -180?. Petala lilacina, medio calceolato stamina lateralia retinenti anthesis initio. Capsulae dehiscentes trilocu- lares, (3-)3.5-5(-5.5) mm longae, 2-2.5 mm latae, loculo dorsali plerumque 1-seminali, loculis ventralibus uterque plerumque 2-seminalibus. Semina loculorum ventralium 1.4- 2.3 mm longa, 1.25-1.6 mm lata, testis scrobiculatis (Figure TYPE.-Uganda, Bunyoro District, 23 mi N of Butiaba turnoff on Butiaba-Murchison Falls Nat. Park road, 6 mi N of Sambiye River crossing, near Kisansya, 16 Sep 1969, Faden, Evans & Lye [in Faden] 69/1066 @A, sheet 1, holotype; BR, EA, sheet 2, K, MHU, isotypes; additional isotypes from material cultivated at the Missouri Botanical Garden, pressed 23 Jun 1973: MO, PRE, US). Perennial herbs (habit types IA and IIA3, Figures 1,2,49d). Roots fibrous. Vegetative shoots decumbent to subscandent, profusely branched; flowering shoots ascending, sparsely branched, to 30 cm tall or more. Internodes (1-)2-12(-15) cm long, green, glabrous (rarely sparsely puberulous and glabres- cent). Leaves spirally arranged, sheaths 0.5-1.5(-2) cm long, puberulous, ciliate at the apex, laminae shortly petiolate, gradually reduced towards the terminal inflorescence, narrowly lanceolate-elliptic or lanceolate to ovate-elliptic, 2.5-7(-8) cm long, 0.5-2.2(-2.9) cm wide, apex acuminate to acute, margin undulate, scabrid, occasionally also sparsely ciliate or ciliolate, both surfaces lustrous, sparsely puberulous, adaxial rarely also, with a few long, uniseriate hairs, abaxial with shorter, more numerous uniseriate hairs on the veins. Inflorescences thyrses (Figure 49b), terminal on the main flowering shoots and frequently on axillary inflorescence- shoots from the inflorescence bract and upper leaves (occasion- ally also terminal on short or long shoots from the lower nodes), moderately lax, ovoid to broadly ovoid, 2-4.5 cm long, 2-6.5 cm wide (to 11 cm long and 15 cm wide in cultivation), with (1-)2-7(-9) cincinni, alternate, subopposite or subverti- cillate (usually with a mixture of arrangements within the thyrse), ascending. Peduncles (1.5-)2.5-5(-6.8) cm long (to 10.5 cm long in cultivation), puberulous. Inflorescence bract 49). 3. Aneilema recurvaturn Faden, new species Aneilema rendlei sensu Chiovenda, Webbia, 8:38, 1951, pro Corradi 2159, p.p. Aneilema tacazzeanum sensu Lewis, Sida. 1:279, 1964 [non Hochstetterex A. Aneilema sp. C.-Faden in Agnew, Upland Kenya Wild Fl.. 666,667,1974. [nm Clarke, 19011. Richard, 1850 = A . forskalii Kunth]. Herbae perennes foliis spiraliter dispositis laminis anguste *. , ,? . . I. FIOURE 49.-Aneilema r e c w a f w n Faden, new species: a, habit; b, inflores- cence; c, bractlcteole; d, perfect flower, front view; e, perfect flower, side view;$ lateral petai, 8. medial staminode; h, antherode of lateral staminode; i, androecium, top view, medial staminode omitted, gynoecium removed; j , ovary and base of style, side view; k, capsule attached to cincinnus, side view; I , dorsal capsule valve, dorsal view; m, dorsal locule seed, dorsal view; n, ventral locule seed, dorsal view; 0, ventrallocule seed, ventral view (a, b, m-o, from Faden & Faden 771785; c-l, from Faden 6911300). (Bars = 20 mm for a; 5 mm for b; 2 mm for ccf.k: 1 mm for c. &I: 0.67 mm for i: 0.5 mm for mo.) NUMBER 76 107 108 SMITHSOMAN CONTRIBUTIONS TO BOTANY supramedial (occasionally medial), usually herbaceous, foli- aceous or bract-like. Inflorescence axis puberulous. Cincinni to 4.2 cm long and 15-flowered (to 10 cm long and 33-flowered in cultivation). Cincinnus bracts (rarely caducous) membranous (rarely the lowermost herbaceous), ascending to patent (rarely reflexed), ovate-elliptic or ovate-lanceolate to ovate or lanceo- late (rarely the lowermost linear), 1.5-3(-9.5) mm long, glandular subapically, usually glabrous or subglabrous. Cincin- nus peduncles generally f uniform within the inflorescence and exceeding the cincinnus bracts, (1.1-)3-10.5(-12) mm long, puberulous. Cincinnus axis puberulous. Bracteoles attached (0.8-) 1-4(-5) mm apart, herbaceous, k symmetrically cup- shaped, perfoliate, 1-2 mm long, 0.5-1.25 mm high, promi- nently glandular subapically and with smaller, inconspicuous glands along the margin, glabrous (occasionally a few hairs near the base, very rarely 1-2 uniseriate hairs at the apex or along the fused edge) (Figure 49c). Flowers perfect and staminate, odorless, 10-14 mm wide (Figure 49d,e; Plate 3124. Pedicels (4.4-)5-8(-9) mm long in flower, to 10 mm long in fruit, ascending in flower, in fruit recurved generally -180?, either k uniformly or mostly at base and apex, persistent, puberulous apically, otherwise glabrous (Figure 49k). Sepals prominently glandular near the apex; medial sepal elliptic to ovate-elliptic or ovate, 2.4-3(-3.5) mm long, 1.8-2.3 mm wide, usually glabrous (occasionally with a few hairs at the base), subapical gland distinctly bilobed, rarely smaller marginal glands also present; lateral sepals broadly elliptic to obovate-elliptic, oblong-elliptic, or ovate-elliptic, 2.5-3.5(-3.8) mm long, 1.6-2.4 mm wide, usually sparsely puberulous basally or medially (occasionally glabrous), subapi- cal gland bilobed or unlobed, marginal glands absent. Paired petals 6.5-9(-11) mm long, 4-6.5(-7.5) mm wide (Figure 49f), limb ovate, 4.5-6.5(-7) mm long, lilac (RHS color: 85C-D and D, Fuden & Faden 77/785, cultivated; 8%-D Fuden & Faden 77J786, cultivated; 87D, Fuden et al. [in Fuden] 69/1066, cultivated), apex acute to obtuse, margin crenulate, claw 1.5-3(-4) mm long, whitish, glabrous. Medial petal slipper-shaped, elliptic, broadest below the margin (viewed from the apex) (Figure 23g-z). retaining the lateral stamens when the flower opens (Plate 3c), 5-7.5(-8.5) mm long, 3-4(-4.8) mm wide, 3.2-4.3(-5) mm deep, concolorous with the limbs of the paired petals laterally, whitish medially. Lateral stamen filaments fused basally to medial stamen filament and sometimes to lateral staminode filaments (Figures 26d, 499. Medial staminode with filament 1.4-2.3(-3) mm long, pale lilac basally, shading to yellow near the apex, antherode bilobed, yellow, lobes sessile or subsessile, obovate- cuneate to transversely elliptic, 0.3-0.65 mm long, connective not or scarcely elongate (Figure 49g). Lateral staminodes with filaments 3-4.8 mm long, very pale lilac shading to yellow above the middle, antherodes bilobed, yellow, lobes sessile or subsessile, obovate-cuneate, sometimes slightly decurved, 0.5-1 mm long, connective not elongate (Figure 49h). Lateral stamens with filaments slightly divergent in basal l/3, strongly divergent in middle l/3, less strongly divergent in apical '/3 (Figures 26d, 499, 6.2-8.6 mm long, S-shaped, geniculate, glabrous, anthers lanceolate to elliptic (to ovate), 0.8-1.2 mm long, 0.5-0.9 mm wide, pollen yellow. Medial stamen with filament 4.2-6 mm long, anther ovate-elliptic to oblong- elliptic, saddle-shaped, 1.5-2.2 mm long, 0.95- 1.4 mm wide, pollen concolorous with lateral anther pollen. Ovary substipi- tate, obovate. to obovate-oblong or obovate-subquadrate, 1.3-1.8 mm long, 0.9-1.45 mm wide, densely and uniformly covered with patent, glandular hairs (Figures 4 2 , 49j), apex emarginate, dorsal locule prominent, 1 -ovulate, ventral locules each 2-ovulate; style 7-8.5 mm long, straight to gently arcuate-decurved for most of its length, also curved laterally out of the floral midplane, stigma capitate, held in front of the anthers and about the Same level as the lateral anthers. Capsules (Figure 49k,I) substipitate or subsessile, oblong- elliptic to obovate-oblong (occasionally obovate), dehiscent, bivalved, trilocular, (3-)3.5-5(-5.5) mm long, 2-2.5 mm wide, pale gray to grayish tan, lustrous, puberulous, apex emarginate, valves persistent or dorsal valve tardily deciduous (occasionally both valves deciduous in old capsules), dorsal locule prominent, 1-seeded or, by abortion, empty, ventral locules each 2-(or, by abortion, 1-)seeded, cells of the capsule wall transversely elongate. Dorsal locule seed (Figure 49m; Plate 6c) elliptic to oblong-elliptic, 1.9-2.6 mm long, 1.25-1.45 mm wide, 0.75-09 mm thick, hilum dark brown, usually in a very shallow groove, much shorter than the seed, testa brown, shallowly scrobiculate or with shallow pits radiating from the embryotega on the dorsal surface, white (rarely gray) farinose granules sparse in the depressions and around the embryotega, usually dense around the hilum, hypha-like filaments occasionally present among the farinose granules; ventral locule seeds (Figure 49n,o; Plate 6c) ovate to trapezoidal, apical seed rounded to obtuse apically, basal seed angular (rarely obtuse) basally, 1.4-2.3 mm long, 1.25-1.6 mm wide, 0.7-0.9 mm thick, hilum dark brown, raised in a groove, not extended onto apical and basal surfaces, testa brown, shallowly scrobiculate or with shallow pits radiating from the embryotega on the dorsal surface, white (rarely gray) farinose granules sparse in the depressions (more abundant than in dorsal locule seed depressions), usually sparse around the embryotega, dense around the hilum, hypha-like filaments occasionally present among the farinose granules. HABrrAT-Grassland, bushland, and bushland thickets, sometimes near streams; sandy or clayey soils; partial shade; 700-1300 m. FmwERING.-Flowering specimens have been seen from March, April, and July-November. In cultivation, the flowers of two populations open at the same time as those of three populations of A. clarkei, which is equivalent to a field time of about 0900 hrs. The flowers of one of the A. recurvutum populations fade at an equivalent time of 1345 hrs. CHROMOSOME NUMBER.-n = 13,2n = 26. DISTRIBU-ITON.-SE Ethiopia, N Uganda, W Kenya, and SPECIMENS EEN.-ETHIOPIA. GEMU-GOFA: Rive del NW Tanzania (Map 12). NUMBER 76 109 8 I I I I ,.::;! 0 ......... ........... .... :.:.:.:.:. ,:::;:x::y .......... ..... G;3 ........ . .-, r e C 0- - - - - - - - : 0 - ' .' . \ r-- - ' . I 1 z . . -r a I ' '-1 I 1. . . ' . . . 1 ,;' . I 0 I - - \ . 100 200 MILES \ , - - \ 0 100 200 KILOMETERS MAP 12.- Distribution of Aneilema recwvatwn Faden, new species. Ghizo, Corradi 2159, p.p. F); base of escarpment W of Arba Minch, 6 O O O ' N , 37'352, Gilbert & Gilbert I61 7 (EA, K); Arba Minch, below escarpment E of town -6'05'N, 37"37'E, Gilbert & Thulin 318 (MO); Arba Minch, cultivated in Addis Ababa, Mistry M-IlA (EA) & M-27 @A). UGANDA. BUNYORO: 23 mi N of Butiaba turn off on Butiaba-Murchison Falls Nat. Park road, 6 mi N of Sambiye River crossing, near Kisansya, Faden et al. [in Faden] 69/1066 (BR, EA-2 sheets, K, MHU); same collection, cultivated Missouri Botanical Garden (MO, PRE, US); near Kisansya on Butiaba Flats, 2'10'N, 310242,Lye et al. 3981 (K). TEsO: 1.8 mi W of Wera, Lewis 5999 (K, MO). KENYA. Without locality, Graham 3019 (K). ELGEYO: Kerio Valley, Kabarnet-Tambach road, 2.8 km before junction with Tot road, Oo32'N, 35'352, Faden & Faden 77/785 (DSM, EA, F, K, MO, US); Tambach-Tot road, Toilang, -0?53'N, 110 SMITHSONIAN CONTRIBUTIONS TO BOTANY 35?372, Faden & Faden 771786 (EA, F, K, US); Tambach- Tot road, 10 km N of junction with Kabarnet-Tambach road, 0?37?. 35?36%, Faden et al. 701894 (BR, EA, K, MO, US). KISUMU-LONDIANI: Kisumu, Hindorf806 (EA). SOUTH Urn- RONDO: Lambwe Valley, Riamkanga, Oo42?30?S, 34?12%, Faden 6911300 (BR, EA, FT, K, MO, PRE); same collection, cultivated Missouri Botanical Garden (B, C, DSM, G, P, UPS, S, US, WAG); Lambwe Valley, Ruma thicket, -0?37?S, 34?18?E, Faden 6911608 @A); Lambwe Valley, between Otuok and Ruma thickets, -Oo39?S, 34?172, Faden 6911642 (EA, MO); Lambwe Valley, Got Jope, W slope, Oo34?S, 34?19?E, Faden 6911699 (EA, K); Lambwe Valley, E side of small hill E of Got Rabondo, -Oo41?S, 34?133, Faden 6911802 (EA, K); Wiga Valley, -Oo40?S, 34?123, Faden 6911933 (EA, FT, K, MO); Lambwe Valley near River, Napier & Fox 3431 [= 68211 (EA, K). TANZANIA. MUSOMA: Ushashi Rest House, Ushashi, Tanner 4011 (BR-2 sheets, K). DISCUSSION Corradi 2159, which is a mixed collection with A. rendlei C.B. Clarke, has been doubtfully included in A. recurvatum. It differs from typical specimens in having more densely pubescent leaves, more patent cincinni, more regularly reflexed cincinnus bracts, shorter cincinnus peduncles, and less widely spaced bracteoles. An apparently mature flower bud was dissected on Corradi 2159 and showed larger antherode lobes on the lateral staminodes and much less curled lateral stamen filaments than buds of typical A. recurvatum plants. Corradi 2159 agrees with A. recurvatum in the form, sue, and lack of pubescence of the bracteoles and in the pubescence of the cincinni, pedicels, and sepals. Although aberrant in many characters, Corradi 2159 is closer to A. recurvatum than to any other species of section Lamprodithyros. However, since it lacks flowers and capsules, a final decision about its taxonomic status must await the collection of better material. Both Faden 6911300 and Faden et al. 701894 are mixed collections with A. indehiscens subsp. keniense (6911300A and 701894A respectively). Their distinctions from the more abundant material of A. recurvatum are discussed under A. indehiscens. Aneilema recurvatum approaches A. indehiscens in habit: however, the long, trailing shoots of the former tend to be much more branched than those of the latter. The differences are maintained in cultivation. The stamen-retention mechanism is well developed in A. recurvatum (Plate 3c,d). In cultivated plants the stamens are held by the medial petal for about one to two hours. Aneilema recurvatum is very common in the Lambwe Valley of southwestern Kenya, where it occurs at the edges of moist thickets composed of a variety of trees and shrubs, including Mystroxylon aethiopicum, Euphorbia candelabrum, Scutia myrtina, Erythroxylumfischeri, and species of Cadaba, Euclea, Grewia, Rhus, and Scolopia. In the Kerio Valley of western Kenya, it occurs in drier thickets with Acacia mellifera, A. brevispica, Teclea pilosa, Grewia sp., Sansevieria ehrenbergii, and Cissus rotundifolia. Aneilema recurvatum is a distinctive species because of its nearly symmetric, glabrous bracteoles, narrow, pointed paired petals, and small capsules and seeds. It appeafs to be most closely related to A. forskalii, A. sebitense, and A. benadirense because of the form of its bracteoles and their marginal glands and lack of pubescence. It differs from A. forskalii by its perennial habit, less-widely spaced bracteoles, smaller, less- obovate, more shortly stipitate to subsessile capsules and smaller seeds. From A. sebitense A. recurvatum may be separated by the latter?s glabrous or glabrescent internodes, more recurved fruiting pedicels that are puberulous at the apex, smaller capsules with never more than two seeds per ventral locule, and smaller dorsal locule seeds. Aneilema recurvatum differs from A. benadirense by the former?s more regularly ascending cincinni, which are fewer per thyrse, generally less-recurved fruiting pedicels, well-developed medial stami- node, and less-humpbacked ventral capsule valve. It further differs from all three species in its less dimorphic dorsal and ventral locule seeds. Aneilema recurvaturn can be confused with A. indehiscens and A. petersii but is easily separated by its more symmetric, glabrous bracteoles that are always perfoliate. It can also be distinguished by its glabrous or subglabrous sepals, more pointed paired petals, well-developed stamen-retention habit, more dimorphic antherodes, more divergent lateral stamen filaments, and generally smaller capsules. Its more or less uniformly colored capsules are paler than those of A. indehiscens. Its seeds are much less dimorphic and smaller than those of A. petersii. 4. Aneilema sebitense Faden, new species Aneilema rendlei C.B. Clarke. FL Trop. Africa, 8:69, 1901 [pro Donaldron Smith 3461. Herbae perennes foliis spiraliter dispositis laminis lanceo- lato-ellipticis ad lanceolatas, ovato-lanceolatas vel ovatas, 3-13.5 cm longis, (0.7-)1.5-4 cm latis, margine ciliata. Inflorescentiae thyrsi ovoidei ad late ovoideos, 4-10 cm longi, 3.5-10 cm lati, cincinnis (2-)5-9(-12) compositi. Bracteolae 2-5(-6.2) mm semotae, plus minusve herbaceae, plus minusve symmetrice cupulatae perfoliatae, glabrae, glandibus numero- sis marginalibus praeditae. Pedicelli tempore fructigero plerumque ascendentes vel erecti, glabri. Capsulae, stipitatae, dehiscentes, (4-)4.5-7.5 mm longae, (2.3-)2.8-3.5(-3.7) mm latae, loculo dorsali l-seminali, loculis ventralibus uterque plerumque 2-3(-4)-seminalibus. Semen loculi dorsalis 2.15- 2.85 mm longum, 1.5-1.7 mm latum, 1-1.2 mm crassum. Semina loculorum ventralium (1.35-)1.5-2(-2.9) mm longa, 1.55-1.9(-2) mm lata (Figure 50). NUME3ER 76 111 FIGURE 50.-Aneilema sebitense Faden, new species: a, habit; b, inflorescence: c, perfect flower, front view; d, perfect flower, side view: e, lateral staminode;f, medial staminode; g, androecium, top view, medial staminode omitted, gynoecium removed; h, gynoecium, side view; i, capsule attached to cincinnus, side view: j , dorsal capsule valve, dorsal view; k, dorsal l o d e seed, dorsal view; I, dorsal locule seed, ventral view; m, ventral locule seed, dorsal view; n, ventral locule seed, ventral view (all from Faden & Faden 77/803). p a r s = 20 mm for a; 10 mm forb; 2 mm for c,d,i; 1 mm for e-h, j ; 0.4 mm for kn.) 112 SMITHSONIAN CONTRIBUTIONS TO BOTANY TYPE.-Kenya, West Suk District (West Pokot District on label), near Sebit on Sebit-Parua road, along Sebit River, lo24?N, 35022?EE, 15 Mar 1977, Faden & Faden 771803, field collections and specimens pressed from cultivation 27 Sep 1977 and other dates ( U S , holotype; BR, EA, F, K, MO, P, US, WAG, isotypes). Perennial herbs (habit types IB, IC, Figures 1, 5Oa). Roots fibrous. Shoots ascending, sometimes rooting at the lower nodes, to -1 m tall. Internodes 1.5-16.5 cm long, puberulous. Leaves spirally arranged, sheaths 1-2.5 cm long, pilose- puberulous, ciliate at the apex, laminae petiolate (except the uppermost leaves), sometimes reduced towards the inflores- cence, lanceolate-elliptic to lanceolate, ovate-elliptic or ovate, 3-13.5 cm long, (0.7-)1.5-4 cm wide, apex acuminate to acute, margin slightly undulate, scabrid, also ciliate at least below the middle and on the petiole, both surfaces k dull, densely puberulous, sometimes also with scattered uniseriate hairs, those of the adaxial surface longer than those of the abaxial. Inflorescences thyrses, terminal and sometimes axillary from the inflorescence bract, lax to moderately lax, ovoid to broadly ovoid, 4-10 cm long, 3.5-10 cm wide, with (2-)5-9(-12) cincinni, ascending (occasionally the lower becoming decli- nate), alternate or a few subopposite (Figure 50b). Peduncles (3-)5- 13 cm long, puberulous. Inflorescence bract usually medial, occasionally supramedial, membranous and bract-like or herbaceous and foliaceous. Inflorescence axis glabrous to sparsely puberulous (at least basally). Cincinni to 5 cm long and 13-flowered (to 6 cm long in cultivation). Cincinnus bracts membranous, appressed to the cincinnus peduncles or patent, ovate-elliptic to ovate or lanceolate, 2-3(-5) mm long, at least the upper ones glandular subapically but marginal glands lacking, glabrous or occasionally sparsely puberulous. Cincin- nus peduncles relatively uniform within the inflorescence, exceeding the cincinnus bracts, 5-lo(-14) mm long, usually glabrous, occasionally sparsely puberulous. Cincinnus axes usually glabrous, occasionally puberulous. Bracteoles attached 2-5(-6.2) mm apart (to 8 mm apart in cultivation), k herbaceous, nearly symmetrically cup-shaped, perfoliate, 1.8- 2.7 mm long, 1-1.6 mm high, prominently glandular subapi- cally and with numerous smaller glands along the margin, glabrous. Flowers perfect and staminate, odorless, 10-20 mm wide (Figure 50c,d; Plate 3e). Pedicels 6.5-10.5 mm long in flower, to 13 mm long in fruit, ascending in flower, ascending to erect or very slightly further recurved in fruit, persistent, glabrous (Figure 50i). Sepals glandular near the apex, glabrous; medial sepal ovate-elliptic to ovate-lanceolate, 3.5-4.4(-6) mm long, 2.2-2.9 mm wide, gland bilobed; lateral sepals broadly elliptic, obovate-elliptic or oblong-elliptic to obovate-oblong or ovate- lanceolate, 3.5-4.5(-6) mm long, 1.9-3.5 mm wide, gland unlobed or bilobed. Paired petals 9-10.5 mm long, 6-10 mm wide, limb ovate, 7-8 mm long, lilac (RHS colors: 81D, Faden & Faden 77J803, cultivated; 82D, Faden & Faden 77/803A, cultivated), apex obtuse or rounded to acute, claw 2-3 mm long, white basally, glabrous. Medial petal cup-shaped, elliptic to obovate, broadest just below the margin (viewed from the apex), retaining the lateral stamens when the flower opens, 7.5-13 mm long, 5.5-8 mm wide, 5-7 mm deep, concolorous with the limbs of the paired petals or slightly paler (Figure 50c,d). Stamen filaments shortly fused basally, sometimes also fused with the lateral staminodes (Figure 50g). Medial staminode with filament 1.7-3 mm long, white or pale lilac shading to yellow above the middle, antherode bilobed, yellow, lobes sessile, transversely elliptic to reniform, 0.4-1 mm long, connective slightly elongate (Figure 50f). Lateral staminodes with filaments 4-5 mm long, white or pale lilac shading to yellow near the apex, antherodes bilobed, yellow, lobes stipitate or sessile, broadly ovate-reniform to obovate, 0.5- 1 mm long, connective not at all to scarcely elongate (Figure 50e). Lateral stamens with filaments parallel to divergent (Figure 50g), 8.7-11.5 mm long, S-shaped, geniculate and decurved near the middle, glabrous, anthers ovate-elliptic to lanceolate-elliptic, 1.1-1.6 mm long, 0.5-1 mm wide, pollen creamy yellow to dirty white. Medial stamen with filament 6-7.5 mm long, anther ovate to lanceolate-ovate or oblong- elliptic, saddle-shaped, 1.7-2.5 mm long, 0.9-1.5 mm wide, pollen golden yellow. Ovary substipitate, obovate to obovate- oblong, -2 mm long, 1.3- 1.5 mm wide, densely and uniformly covered with patent, glandular hairs, apex truncate, dorsal locule prominent, 1-ovulate, ventral locules each 2-3(-4?)- ovulate; style -10.5 mm long, arcuate-decurved for most of its length, also slightly curved laterally, stigma capitate, usually held in front of the lateral anthers (Plate 3e). Capsules (Figure 50i,j) stipitate, obovate-oblong to obovate, oblong, oblong-elliptic or obovate-elliptic, dehiscent, bivalved, trilocular, (4-)4.5-7.5 mm long, (2.3-)2.8-3.5(-3.7) mm wide, tan or grayish tan, sometimes marked with blue apically or flecked with dark brown spots, lustrous, puberulous to subglabrous, apex emarginate to rarely subtruncate, dorsal valve deciduous, dorsal locule prominent, 1-seeded or, by abortion, empty, ventral locules each 2-3(-4) seeded, some- times (in cultivation), by abortion, 1-seeded; cells of the capsule wall transversely elongate, arranged in regular files. Dorsal locule seed (Figure 50k,I) not or scarcely humpbacked towards the capsule apex, elliptic, 2.15-2.85 mm long, 1.5-1.7 mm wide, 1-1.2 mm thick, hilum dark brown, in a shallow groove, shorter than the seed, testa buff or buff-orange, very shallowly reticulate or scrobiculate to k smooth, brownish farinose granules very sparse in the depressions and more abundant around the hilum or confined to the hilum, hypha-like filaments very sparse in the depressions or absent. Ventral locule seeds (Figure 50m,n) ovate to trapezoidal, apical seed rounded to obtuse apically, basal seed angular basally, (1.35-)1.5-2(-2.9) mm long, 1.55-1.9(-2) mm wide, 1-1.25 mm thick, hilum dark brown, raised in a groove, not or slightly extended onto apical and basal surfaces, testa tan or yellowish brown, shallowly reticulate to reticulate-scrobiculate, brownish NUMBER 76 113 farinose granules sparse in the depressions and around the embryotega, dense around the hilum, hypha-like filaments occasional or frequent in the depressions. HABITAT.-d?acia -Commphora -Combreturn woodland and dry hillsides above river; rocky areas; -1 120-1550 m. FLOwERING.-~OWeritIg specimens have been seen from April, July, and August. In cultivation, the two Faden and Faden collections and Gilbert & Thulin 266 flower early in the morning (before A. recurvaturn), but the exact hours have not been determined. CHROMOSOME.-2n = 52. DISTRIBUTION.-SW Ethiopia and NW Kenya (Map 10). SPECIMENS EEN.-ETHIOPIA. GEMU-GOFA: Arba Minch, escarpment below the town, -6?OSN. 37?3% Gilbert & Thulin 266 (MO). SIDAMO: [?Somali-land? on label], Donuld- son Smith 346 (BM). KENYA. WEST SUK: Near Sebit on Sebit-Parua road, along Sebit River, 1O24?N, 35?22?E, Fuden & Fuden 771803 (BR, EA, F, K, MO, P, US, WAG); same locality, Fuden & Faden 77/803A (F, US); Cherangani Hills, R. Sebit, 1?12? [sic: changed to 1?24? on EA sheet], 35?19?E, Mubberley & McCull88 (EA) and 88a (K) [part of same collection according to Mabberley (in litt.)]. DISCUSSION Donaldson Smith collected his number 346 on 23 April 1895, the same day as he collected the type of Aneilemu rendlei. The locality is in Ethiopia (foot of the Amaro Mountains, Burga Camp) not in the Somali Republic, according to the collector?s account of his journey (Donaldson Smith, 1897). Clarke (1901) treated this specimen as A. rendlei but noted that it could hardly be distinguished from the Australian A. acuminutum. The resemblance to the latter is purely superficial. For distinctions from A. rendlei see discussion under that species. In Faden (1975:289) I treated this as a species apparently distinct from but closely related to A. forskalii. The inadequacy of the four herbarium specimens then available dissuaded me from formally describing this species. The living material subsequently obtained of both A. forskalii and A. sebitense has confirmed my original conclusion. Aneilemu sebitense resembles A. forskalii in its bracteoles relatively widely spaced, more or less symmetrically cup- shaped, glabrous, and with numerous marginal glands, fruiting pedicels mostly erect, and fruits commonly with more than two seeds per ventral locule. It differs from A. forskulii by its perennial habit, ciliate leaf margins, generally less widely spaced bracteoles, narrower, more shortly stipitate capsules with up to four seeds per ventral locule, and smaller seeds. In addition, the petals of A. sebirense seem to be darker in color and the antherodes of the lateral staminodes different in shape than those of A. forskalii. Aneilemu sebitense also resembles A. recurvaturn but can be distinguished by its glabrous pedicels that are generally less reflexed in fruit, less-divergent lateral stamen filaments, and larger capsules with larger seeds that are more dimorphic. Living plants of Fuden & Faden 77/803 are much larger, more robust, and have much larger flowers than those of Gilbert & Thulin 266. Their flowers also differ in details of color and the shape of some parts. I initially thought that these populations might be worth separating at some taxonomic level, but they agree in all diagnostic characters, and both populations proved to be tetraploid. 5. Anei lem forskalii Kunth Aneilema forskalii Kunth [?Anilema Fonkalei?], Enum. Pl., 4:71. 1843.4.B. Clarke in De Candolle, Monogr. Phan., 3:231. 1881.Schweinfurth, Bull. Herb. Boissier, 1, append.. 2:58, 1894. [Type as for Commelina tuberosa Forsska.] Commelina tuberosa Forsska, Fl. Aegypt.-Arab. Descript.. 12, 1775 [nom. illeg., non L. (1753). Syntypes: Arabia felix [Yemen], Montes Hadienses ad Bulgliose, Mar 1763, Forsska in Herb. Forskalii 33 (C, lectotype; photo K, MO, US), in Herb. Forskalii 31 (C, photo US) and in Herb. Forskalii 32 (C; photo K, MO, US)]. Commelina panicdata Vahl, Enum. PI., 2:179, 1805-1806 [nan. illeg., non Hill (1773)].-Roemer and Schultes, Syst. Veg., 1538, 1817. [Type as for Commelina fuberosa Forssk&l.] Aneilema tacozzeanum Hochstetter ex A. Richard, Tent. Fl. Abyssinicae, 3(5):343, 1850 [?tacamana?].-Walpen, Ann. Bot. Syst., 3:658, 1852.- Hua, Bull. Mus. Hist. Nat. (Paris), 1:120, 1895 [in adnot. sub Aneilema aequinoctiale].-Schumann in Engler. Pflanzenwelt Ost-Afrikas C, 136, 1895.-Andrews. Fl. P1. Sudan, 3:239, 1956.-Brenan, Kew Bull., 15:215, 1961.4ufodontis. Bull. Jard. Bot. Natl. Belgique, 41, suppl., 1518, 1971. [Syntypes: Ethiopia, bords du Tacazze [undated]), Quartin Dillon & Petit s.n. (P-2 sheets; photo EA, K lectotype: P sheet ex Herbarium Richard); Valle fluvii Tacaze, prope Dscheladscheranne [Djeladjeranne; cited by Richard as ?Tchelatchekannte?] Schimper 1660 (G-2 sheets, K-2 sheets, Lomprodithyrm tacazzeanus (Hochstetter ex A. Richard) Hasskad. Flora, 46:390, 1863; in Peten, Natunviss. Reise Mossambique, Bot., 531. 1864 [nan. tant., desc. et Peters s.n. excl.]. Lomprodithyros ehrenbergii Hasskarl in Schweinfurth, Beitr. Fl. Aethiop., 210, 1867. [Type: Ethiopia, Togodele, 1822, Ehrenberg 413 (B; photo K, fragment L).] Commelina ebracteata Ehrenberg, in sched. (Ehrenberg 413).-Hasskad in Schweinfurth, Beitr. Fl. Aethiop., 210,1867 [pro syn.]. Aneilema ehrenbergii (Hasskarl) C.B. Clarke in De Candolle. Monogr. Phan., 3:229. 1881 [Welwirsch 6610 excl.]; in Thiselton-Dyer, Fl. Twp. Africa, 8:74. 1901.-Durand and Schinz, Conspect. Fl. Africae, 5430, 1895.- Cufodontis, Bull. Jard. Bot. Natl. Belgique, 41. suppl., 1517, 1971. Aneilema tacazzeanm Hochstetter ex C.B. Clarke in De Candolle, Monogr. Phan., 3:222, 1881; in Thiselton-Dyer, Fl. Trop. Africa, 8:66, 1901.- Durand and Schinz. Conspect. Fl. Africae, 5:132,1895.-Tacaolm and Drar, H. Egypt, 2:433, 1950. [Syntypes: Ethiopia, Schhper 1660, p.p. maj. [fide Clarke, 18811 (G, P); Schimper 1729 (G, K. P); Oudgerade. PefL s.n. (P, not seen); Mozambique, Peters s.n. (B, destroyed, fragment L).] Aneilema aequinoctiale (P. de Beauvois) G. Don p minor C.B. Clarke in De Candolle, Monogr. Phan., 3:221, 1881.-Durand and Schinz, Conspect. Fl. Africae. 5429,1895. [Syntypes: Ethiopia, Schimper 1660, p.p. [fide Clarke] (P); Quartin Dillon [& Perit] s.n. (P); Yemen, Botta s.n. (P).] Annual herbs (habit types IB?, IC, IIIA, IIIB, Figure 1). Roots fibrous. Shoots erect to ascending or shortly decumbent, MO, P-3 sheets, photo EA).] 114 SMITHSONIAN CONTRIBUTIONS TO BOTANY unbranched or sparsely branched, 20-50(-70) cm tall. Inter- nodes (0.5-)2.5-12(-19) cm long, sparsely to densely puberulous. Leaves spirally arranged, sheaths 0.5-1 3 - 2 ) cm long, pilose-puberulous (occasionally puberulous), ciliate at the apex, laminae petiolate, sometimes gradually reduced towards the inflorescences, mostly lanceolate-elliptic to ovate-elliptic (occasionally lanceolate or ovate), 2.5-12 cm long, (0.5) 1.3-4.5 cm wide, apex acute to acuminate, margin usually planar (occasionally slightly undulate), scabrid, occasionally also sparsely ciliolate, ciliate on the petiole, both surfaces puberulous, adaxial also generally with scattered, long, uniseriate hairs, abaxial often with numerous, shorter uniseriate hairs on the veins. Inflorescences thyrses, terminal on the main shoots and frequently also on long or short axillary shoots, lax to moderately lax, ovoid to broadly ovoid, 2.5-6(-12.8) cm long, 2.5-6(-12) cm wide, with (1-)3-lo(-12) cincinni, alternate or subopposite (rarely some subverticillate), ascending (occa- sionally the lower becoming patent or declinate). Peduncles 4-15(-22.5) cm long, puberulous. Inflorescence bract supra- medial (rarely medial), herbaceous, bract-like or foliaceous. Inflorescence axis puberulous to subglabrous (rarely glabrous). Cincinni to 9.5 cm long and 21-flowered. Cincinnus bracts herbaceous to membranous, usually ascending and closely appressed to the cincinnus peduncles, ovate to ovate-elliptic or lanceolate-ovate (rarely narrowly lanceolate to lanceolate- elliptic), (1.2-)1.5-3(-5.5) mm long (the lowermost always longest and commonly perfoliate), prominently glandular subapically, also usually with smaller glands along the margin basally (marginal glands rarely absent), glabrous (rarely the lowermost sparsely puberulous). Cincinnus peduncles gener- ally k uniform within the inflorescence, exceeding the cincinnus bracts, (2.3-)4-12(-15.5) mm long, glabrous or sparsely puberulous. Cincinnus axes glabrous (rarely sparsely puberulous). Bracteoles attached (1-)3-7(-8.5) mm apart, herbaceous, f symmetrically cup-shaped, perfoliate, (1.15-)1.3-2.2(-2.95) mm long, (0.5-)0.75-1.3(-1.75) mm high, prominently glandular subapically, with smaller glands along the margin, glabrous. Flowers perfect and staminate, odorless, 11-18 mm wide (Plate 3f). Pedicels 4-8(-11.5) mm long in flower, to 12 mm long in fruit, ascending in flower, erect or recurved to -150" in fruit, persistent, glabrous or very sparsely puberulous at the apex. Sepals prominently glandular near the apex, glabrous (very rarely with a few hairs at the base); medial sepal lanceolate-elliptic to ovate-elliptic, 2.1-4.2 mm long, -1.7-2.3 mm wide, subapical gland bilobed (one sepal in Defers 968 also with a marginal gland); lateral sepals ovate-elliptic to oblong-elliptic (rarely suborbicular), 2.6-4.5 mm long, -2 mm wide, subapical gland unlobed or bilobed. Paired petals white to pale shades of lilac, lavender, mauve or blue, limb ovate, apex k obtuse, claw white or whitish, glabrous. Medial petal cup-shaped or slipper-shaped, 5.5-8(-lo?) mm long, concol- orous with paired petal limbs or paler than them. Lateral stamen filaments fused basally to medial stamen filament and to lateral staminode filaments (Figure 26c). Medial staminode with filament 1.5-1.8 mm long, antherode bilobed, lobes sessile, usually * transversely elliptic, occasionally obovate-cuneate, connective elongate or not. Lateral staminodes with filaments 2-3.5 mm long, antherode bilobed, lobes sessile or subsessile, obovate-cuneate, 0.3-0.8 mm long, connective not elongate (rarely scarcely elongate). Lateral stamens with filaments 5.4-9.5 mm long, S-shaped, glabrous, anthers lanceolate- elliptic to elliptic or ovate-elliptic, 0.85-1.3(-2) mm long, pollen whitish, yellowish or golden yellow. Medial stamen with filament 4-6 mm long, anther ovate to ovate-elliptic (rarely lanceolate), saddle-shaped, 1.1-1.5(-2.5) mm long, pollen golden yellow, concolorous with lateral anther pollen or strongly discolorous. Ovary stipitate, obovate-subquadrate, 1.5-2(-3) mm long, densely covered with capitate, glandular hairs, apex emarginate, dorsal locule prominent (always?), l-ovulate or empty, ventral locules (1-)2-3-ovulate; style (3-)6-9.5 mm long, arcuate-decurved, sometimes recurved towards the apex, sometimes curving out of the floral midplane, stigma capitate, held near or in contact with the lateral and/or medial anthers. Capsules stipitate, obovate to obovate-oblong (rarely obo- vate-elliptic), dehiscent, bivalved, usually uilocular, (3.5-) 4.5-7.5(-8) mm long, (2.25-)3.3-4.5(-4.9) mm wide, buff to pale grayish tan or tan, frequently with small, scattered, brown spots, lustrous, glabrous or subglabrous, apex emarginate (rarely truncate), dorsal valve deciduous (rarely both valves deciduous in old capsules), dorsal locule prominent or not, when prominent, 1-seeded or, by abortion, empty, ventral locules each (1-)2-3-seeded, cells of the capsule wall transversely elongate or irregular. Dorsal locule seed convexo- planar, slightly humpbacked towards the capsule apex, elliptic to ovate-elliptic or oblong-elliptic, (2.4-)2.7-3.6 mm long, (1.5-)1.95-2.1 (-2.4) mm wide, 1.25-1.5 mm thick, hilum dark brown, in a shallow groove, shorter than the seed, testa buff-orange or buff, f smooth, farinose granules and hypha-like filaments absent; ventral locule seeds (Plate 6 4 ovate to triangular or trapezoidal, apical ventral seeds acute to obtuse or rounded apically, basal ventral seeds angular basally, (1.5-)1.8-2.6(-3.1) mm long, (1.4-)1.7-2.2(-2.5) mm wide, (1.1-)1.2-1,45(-1.55) mm thick, hilum blackish or dark brown, raised in a groove, usually slightly extended onto apical or basal surface, testa brown or gray-brown (occasionally buff-orange or orange-brown), shallowly scrobiculate or scrobiculate-reticulate (rarely shallowly foveolate), white or brownish farinose granules sparse in the depressions and around the embryotega, denser around the hilum, hypha-like filaments present among the farinose granules. HABITAT.-Thickets, scrub, and dry forest, often near streams or seepage areas, commonly among rocks; partial or dense shade; (300-)600-1500(-1700?) m. RowERINa-Flowering specimens have been seen from January-April, June, August-October, and December. From NUMBER 76 115 MAP 13.-Distribution of Aneilema forskalii Kunth. collection dates and condition of the specimens, July should be added. In cultivation the flowers are open in the morning, but exact opening and fading times have not been determined. CHROMOSOME NUMBER.-n = 26,2n = 52. DISTRIBUTION.-R~~ Sea Hills of Sudan, N and central Ethiopia, Djibouti, E Yemen Arab Republic, S Southern Yemen, and S Oman (Map 13). SPECIMENS SEEN.-SUDAN. KASSALA: Erkowit (Jebel Seila), Andrews A3589 (K); Erkowit, Red Sea Hill, Aylmer 182 6): Gebl Elba, Wadi Rabdit, Drur 114 ( S ) ; Gebl Elba, Drar 287 ( S ) ; Gebl Elba, Wadi Idaib, Drar 300B ( S ) ; Red Sea Hills, Hetatera, Jackson 2804 (K); Red Sea Hills, below Erkowit, Mafley 51 (K); Gebel Elba, Wady Ekwal, Shabetai F1507 bis (K); Gebel Elba, 23-27 Jan 1929, Tackholm s.n. ( S ) . ETHIOPIA. ERITREA: Ghinda, Baldrati 933 0; Filfil- Marrao, Baldrati 1387 0; Sul fiume Mederreba, Ghinda, Baldrati 1388 (FT); Togodele, Ehrenberg 413 (B, fragment L); Hamasen, Ghinda, Fiori 874 0; Hamasen, Embatcalla, Fiori 875 m); Amasen, Dongollo press0 Ghinda, Pappi 4511 (EA, G-2 sheets, MO, US); Au-dessous de Gheleb, Schweinfurth 1416 (G-2 sheets); Gorge d?Ailet pres Saati, Schweinfurth & Riva 427 (G-2 sheets, K); Mont Bosco press Saati, Schwein- furth & Riva 565 (G-2 sheets); Haichello Kobob au nord d? Acrour, Schweinfurth & Riva 1672 (G-2 sheets). HARAR: Rock valley 40 km from Harar, cultivated at Addis Ababa, Mistry M - 7 (EA). SHOA: Blue Nile Gorge, 3 km S of bridge on Addis Ababa-Debre Marcos [road], Gilbert & Gilbert 2186 (K); Blue Nile Gorge, S side, Gilbert & Thulin 969 (MO); -40 km E of Nazareth, de Wilde & de Wilde-Duyfes 10529 (WAG). TIGRE: In valle fluvii Tacaze p r o p Dscheladscheranne [Djeladjeranne], Schimper 1660 (B, G-2 sheets, K-2 sheets, M, MO, P-3 sheets, S; photo EA) and 1729 (G, K, P, S ) . TIGRE/BAGEMDER:Chire, Quartin Dillon 449 (P); without locality or date (ex Herb. de Franqueville & Herb. Cosson 18), Quartin Dillon s.n. (P); Bords du TacazzE, Quartin Dillon & Petit 449 (K) & Quartin Dillon s.n. (P-2 sheets; photo EA, K). DJIBOUTI. District uncertain: Matoki, Saboureau 259 (P). TADJOURA: Goda Mts., steep gorge leading down to Wadi Toha, Lavranos 11466 (K, MO). YEMEN ARAB REPUBLIC. Without locality, 1837, Botta 116 SMITHSONIAN CONTRIBUTIONS TO BOTANY s.n. (P); Montes Hadienses ad Bulgliose, Forsskal in Herb. Forskalii 31 (C, photo US) and 32 and 33 (both C; photo K, MO); wadi above Suq Ar Ribat, Jebel Raymah, Wood 1892 0; between Suq Ar Ribat and Beni Danan on the Nagila path, Jebel Raymah, Wood 1902 0; Wadi Dur, Udayn, Wood 1997 0; above Suq As Sabt, Jebel Bura, Wood 21 73 0; above the Wadi Yure between Jebels Hufash and Melhan, Wood 2395 0; above the Wadi Hijan, Usil in the Haraz, Wood 2580 0; N of Udayn Suq, Wood 2895 0; by the Wadi Sughna, Haimai al Dakhalia, Wood 2959 0; Gebel Bura, Hille, Schweinfurth 289, p.p. (G-2 sheets); Regma, Hodjela, Schweinfurth 944, SOUTHERN YEMEN. Bilad Fodhli, montis el?Areys, prope Serrya, Deflers 968 (G-3 sheets, P). OMAN. Anat Road, 17?09?N, 54?142, Mandaville 6860 (BM); Salalah-Thamarit road, 17?09?N, 54?082, Mandaville 6886 (BM); Jariz Dhofar, foothills, Vesey -FitzGerald 1236/8 P.P. (G-2 sheets). (BM). DISCUSSION The nomenclatural history of A. forskalii is one of the most complex in the genus. Forsskal?s illegitimate name for the plants that he collected in the Yemen, Commelina tuberosa, was published posthumously (Forsskal, 1775). The basis for his specific name is unclear: the three specimens in the ForsskAl Herbarium in Copenhagen lack bases, and ForsskAl?s descrip- tion does not mention the roots. All collections of this species with bases and the living material observed in cultivation show only fibrous roots. Vahl(l805-1806) appears to be the only botanist (with the possible exception of Schweinfurth) who studied ForsskAl?s collections of this species. He expanded ForsskAl?s description and changed the name to Commelina paniculata, which, unfortunately, was also illegitimate. Kunth (1843) properly placed this species in Aneilema and changed the specific epithet to honor the collector. Richard (1850) described Aneilemu tacazreanum from two Ethiopian collections (Quartin Dillon & Petit s.n. and Schimper 1660). These collections clearly belong to one species, the only notable differences being the longer sepals and glabrous cincinnus axes in Schimper 1660. A comparison of these specimens with the syntypes of A. forskalii revealed no significant differences. Accordingly, A. tacazreanum Hochstet- ter ex A. Richard has been placed in synonymy. It should be noted that Richard (1850) omitted Petit?s name as a collector, apparently inadvertently. The specimen from the Richard Herbarium, which is here chosen as the lectotype of A. tacarzeanum, has both Quartin Dillon?s and Petit?s names on the label. However, there is also a sheet in the Paris Herbarium with only Quartin Dillon?s name on it, which is almost certainly part of the same collection, but it is not clear whether this specimen was seen by Richard. Quartin Dillon 449 and Quartin Dillon & Petit 449 are also part of the type collection. The only previous worker to suggest that A. tacazzeanum is a synonym of A. forskalii was Schweinfurth (1894:59), who had collected plants on both sides of the Red Sea, including some from near the type locality of A. forskalii. Clarke, who had not seen ForsskAl?s collections, stated that such synonymy was highly unlikely and that, based on the description of Vahl (1805-1806) of Commelina paniculata, A. forskalii ?was almost certainly a species of Commelina? (Clarke, 1901:66). Richard?s species has been much confused. Hasskarl(1864a) made the combination Lamprodithyros tacazzeanus, based on Richard?s name, and he provided a very detailed description, but the latter was based on a collection from Mozambique (Peters s.n.), where the species does not occur. Brenan (1961) stated that the Mozambique collection was most likely A. nyasense, and a fragment that I found in the Leiden Herbarium (L), which came from Hasskarl?s personal herbarium, has confirmed that. Similarly, A. tacazzeanum of Baker (1875) is A. spekei; that of Chiovenda (1939) is partly A. somaliense and partly A. petersii (not at all A. tacazzeanum); and that of Lewis (1964) is A. recurvatum. Clarke (1881a) concluded that Schimper 1660 was a mixed collection and that Richard (1850) had not selected the specimens that Hochstetter had intended to bear the name Aneilemu tacazzeanum. Clarke (1 88 la) therefore distinguished A. tacazzeanum Hochstetter ex C.B. Clarke from A. tacazzeanum A. Richard, which he then described as A. uequinoctiale var. minor. Clarke?s interpretation of Richard?s A. tacarreanum was based largely on the latter?s description of the capsule locules being about six-seeded, which, as Brenan (1961) later pointed out, was evidently an error by Richard. Clarke?s distinction of A. aequinoctiale var. minor from A. tacazzeanum was challenged by Hua (1895: 121), who noted that the plants annotated by Clarke as his var. minor were totally different from other collections of A. aequinoctiale. He further stated that the specimens of var. minor could not be distinguished from A. tacazreanum. The response of Clarke to Hua?s evidence was, ?Hua appears to draw the line between the two species at a different point? (Clarke, 1901:66). Brenan (1961), however, examined the specimens upon which Richard (1850) based his A. tacazzeanum, including those used by Clarke for the description of A. aequinoctiale var. minor, and found them all conspecific and distinct from A. aequinoctiale. I fully concur with this conclusion. Both A. aequinoctiale var. minor and Clarke?s A. tacazzeanum are therefore synonyms of A. tacarzeanum Hochstetter ex A. Richard. Neither Hasskarl (1867) nor Schweinfurth (1894) mentions a collection number for the type of Lamprodithyros ehrenber- gii, but Ehrenberg 413 bears both Hasskarl?s and Schwein- furth?s handwriting on it, and it is clearly the type. The holotype consists of a sheet with two clearly annual plants mounted on it. The few small inflorescences lack flowers and have only two immature capsules, These plants differ from all other speci- mens of A. forskalii in having more densely puberulous cincinnus peduncles and axes, smaller bracteoles, and shorter NUMBER 76 117 capsules. They also lack marginal glands on the cincinnus bracts, which are present in nearly all specimens of A. forskalii. Despite these differences, Ehrenberg 413 agrees with A. forskalii in its annual habit, glabrous bracteoles with prominent marginal glands, more or less erect, moderately long fruiting pedicels that are puberulous only at the apex, and glabrous sepals. Even in the characters in which it departs from A. forskulii, Ehrenberg 413 is connected to more typical speci- mens by a series of intermediates. Although there is no direct evidence that A. forskalii can ever be a perennial, several particularly robust specimens make one wonder. Schweinfurth 1416 and Schweinfurth & Riva 1672 have particularly large bracteoles (2-2.75 mm long, 1-1.75 mm high) and frequent lateral inflorescence-shoots. They have the longest cincinnus bracts observed in the species, the bracts also being unusual in nearly always lacking marginal glands. Like these two collections, De Wilde & De Wilde-Duyfjes 10529, the southernmost collection of A. forskalii, has unusually long bracteoles and cincinnus bracts. These three collections otherwise agree with A. forskalii, and they probably represent no more than plants growing under very favorable conditions. Mistry M-7 is known to me from a single specimen of a plant cultivated in Addis Ababa. On the basis of vegetative and inflorescence characters, it cannot be separated from A. forskulii. Its flowers, which have been very carefully pressed, appear to conform with specimens of this species except for the smaller size of some parts, e.g., medial petal 5.5 mm long and medial stamen filament 4 mm long. The style is particularly short, 3-4 mm long, and it is not recurved at the apex. The position of the stigma relative to the lateral anthers suggests self pollination. Whether the flowers of Mistry M-7 are truly unusual for A. forskulii is difficult to ascertain, because so few specimens have measurable floral parts. The most unusual characters of Mistry M-7 are those of the capsules and seeds. The capsules are often obovate-elliptic, a shape rare in A. forskulii. Their width (2.25-3.25 mm) falls completely outside the range of other specimens of this species. The ventral locules are almost all one-seeded-a single two-seeded example has been noted-a character that, although known, is uncommon in A. forskalii. The ventral locule seeds are narrower and thinner than comparable seeds from other material of A. forskalii. Their elliptic shape and shallowly foveolate testa have not been found elsewhere in the species. Mistry M-7 could readily be segregated from A. forskalii either at the specific or subspecific level on the basis of capsule and seed characters. I feel that such separation should await the availability of specimens pressed in the field in order to determine whether some of the peculiarities observed in Mistry M-7 may have been due to cultivation conditions. The pedicels of A. forskalii are less recurved than those of any other species of section Lumprodithyros with thyrsiform inflorescences except A. sebitense. They are sometimes erect, but more commonly the apex is recurved somewhat further. Great variation in fruiting pedicel length is found within inflorescences of some specimens, e.g., DefIers 968. Overall, variation in pedicel length is considerable in A. forskalii, but no clear geographic pattern is recognizable, although most specimens with particularly short pedicels are from Eritrea (but so are some very longly pedicellate ones!). Petal color, rarely noted by collectors, appears to be white or paler shades of bluish or pinkish purple. The possible yellow color, described by Andrews (1956) for Sudanese plants and perhaps based only on the collector's notes on Aylmer 182, is most likely erroneous. Staminodes have been seen in many flowers and buds. The differentiation between the medial and lateral antherodes- lobes wider than long in the medial antherode and longer than wide in the lateral antherodes-is probably constant in this species. The antherodes are all yellow. The number of seeds per ventral locule varies from one to three. No collection definitely has solely one-seeded nor solely three-seeded ventral locules, although several show only two-seeded ventral locules. The one-seeded locules do not appear to be the result of abortion of ovules in many cases, but rather to the presence of solitary ovules in the ovary. Similarly, the absence of a dorsal locule seed in the capsules of Gilbert 3941 was probably due to the absence of an ovule in that locule of the ovary (noted in the one ovary which could be dissected), not to abortion. A decrease in the number of seeds per ventral locule within the inflorescence has been noted in Defers 968. The first capsules produced within the inflorescences are the largest and have two-seeded ventral locules. They also have the longest pedicels. Later capsules are smaller and have only one-seeded locules. Similar variation has not been noted in other collections. An altitude of 1700 m is recorded on both sheets of Schweinfurth 1416; however, the same collection is listed as 1500 m by Schweinfurth (1894). Several collections made by Wood are also from about 1500 m, and that is the highest confirmed altitude for A. forskalii. The records of A. forskalii (as A. tucazreanum) from Egypt ('Rickholm and Drar, 1950) appear to be based on old boundaries between that country and Sudan. All of the specimens are from the Gebel Elba area, which is shown in modem maps to be part of Sudan. Aneilema forskalii should be expected to occur in the mountains of northern Somali Republic (Lavranos, personal communication), but it has not yet been collected in that country. In addition to the specimens seen for this study, further collections are cited by Schweinfurth (1894) and Tackholm and Drar (1950). It is uncertain whether any of the Schweinfurth collections cited by him but not seen by the writer still exist. None are in the Berlin Herbarium. Despite great variation, Aneilema forskalii is a very distinctive species because of its annual habit, small cincinnus bracts with generally glandular margins, usually glabrous or 118 SMITHSOMAN CONTRIBUTIONS TO BOTANY subglabrous cincinni, relatively thick-textured, widely spaced, nearly symmetric bracteoles with conspicuous marginal glands, generally erect or only slightly further recurved, glabrous or subglabrous fruiting pedicels, glabrous sepals, and large, longly stipitate, glabrous, humpbacked capsules sometimes with three seeds in each ventral locule. Aneilemu forskulii appears to be most closely related to A. recurvutum, A. sebitense, and A. woodii. From the first species it differs by its generally glandular cincinnus bract margins, more widely spaced bracteoles, generally more erect fruiting pedicels, paler flowers, larger, more longly stipitate, f glabrous capsules, and larger seeds. For differences from A. sebitense and A. woodii, see the key and discussions under those species. 6. Aneilema woodii Faden, new species Herbae perennes foliis plerumque spiraliter dispositis, laminis lanceolatis ad lanceolatoellipticas, lanceolato-ovatas vel ovato-ellipticas, 5-lo(-13) cm longis, (1.5-)2-4.5 cm latis, supra pilis longis uniseriatis plerumque carentibus. Inflorescentiae thyrsi ovoidei, ad 5 cm longas et 7 cm latas, cincinnis 2-9 compositi. Bracteolae 0.8-3 mm semotae, membranaceae, cupulatae (symmetrice vel asymmetrice), plerumque perfoliatae, 1.5-2 mm longae, ad 1 mm altas, puberulae, glandibus marginalibus carentibus. Pedicelli 4.5- 7.5 mm longi, tempore fructigero recurvati plerumque -180?, puberuli. Sepala purberula. Capsulae stipitatae, dehiscentes, triloculares, 5-7 mm longae, 4-5 mm latae, loculo dorsali plerumque l-seminali, loculis ventralibus uterque plerumque 2-seminalibus. Semen loculi dorsalis 2.7-3.2 mm longum, 2.1-2.5 mm latum, 1.4-1.8 mm crassum, testa laevi. Semina loculorum ventralium 1.9-2.9 mm longa, 1.9-2.4 mm lata, testis reticulato-foveolatis non profunde (Figure 5 1). TYPE.-Yemen [Yemen Arab Republic; Yemen (Sana'a)], just above Hilla, Jebel Bura, 15 Feb 1980, Wood 2616B (living only); pressed from cultivation 4 Nov 1980 and other dates (US, holotype; EA, F, K, MO, P, isotypes). Perennial herb (habit types IB, IC, Figure 1). Roots fibrous. Shoots erect to ascending (sometimes decumbent?), to more than 1 m tall in cultivation. Internodes (2-)5-7(-15) cm long, green, puberulous at least below the nodes. Leaves spirally arranged (rarely distichous in part), sheaths 1-2.5 cm long, puberulous, also sparsely pilose (particularly at their bases), ciliate at the apex, laminae petiolate, lanceolate to lanceolate- elliptic, lanceolate-ovate or ovate-elliptic, 5-lo(-13) cm long, (1.5-)2-4.5 cm wide, apex acuminate, margin undulate, scabrid, sparsely ciliate basally, adaxial surface puberulous, typically lacking long, uniseriate hairs, pale-veined, abaxial surface pilose-puberulous. Inflorescences thyrses (Figure 51b), terminal on the main flowering shoots and on axillary inflorescence-shoots from the inflorescence bract and upper leaves (the lowermost of these inflorescence-shoots perforating the subtending leaf sheath, Figure Slu), moderately lax, ovoid, to 5 cm long and 7 cm wide, with 2-9 cincinni, alternate to subopposite, ascending to (in the lowermost cincinni) patent. Peduncles 3-12 cm long, puberulous. Inflorescence bract usually supramedial, bract-like or foliaceous. Inflorescence axis puberulous. Cincinni to 3 3 - 5 3 cm long and 15(-17)-flowered. Cincinnus bracts membranous, appressed to the cincinnus peduncles or patent, lanceolate-oblong to ovate, 1.5-4(-6) mm long, glandular subapically (apparent only in young inflorescences), puberu- lous. Cincinnus peduncles relatively uniform within the inflorescence, exceeding the cincinnus bracts, 4-9(-11) mm long, puberulous. Cincinnus axes puberulous. Bracteoles attached 0.8-3 mm apart, membranous, cup-shaped (symmetri- cally or asymmetrically), usually perfoliate, 1.5-2 mm long, to 1 mm high, glandular subapically but not along the margins, puberulous. Flowers perfect and staminate, odorless, 10.5-17 mm wide (Figure 51c,d; Plate 3g). Pedicels 4.5-6 mm long in flower, to 7.5 mm long in fruit, ascending in flower, recurved generally -180' in fruit (Figure 5121, persistent, puberulous in the apical 1/2-3/4. Sepals glandular subapically with unlobed glands, puberulous; medial sepal elliptic to ovate-elliptic, 3-4.5 mm long, -3 mm wide, subapical gland prominent; lateral sepals not reflexed, broadly elliptic to obovate-elliptic, 4-5 mm long, 3.5-4 mm wide, subapical gland not prominent. Paired petals 8-11 mm long, 8-10 mm wide, limb broadly ovate to ovate-reniform, 6-8 mm long, white faintly tinged with pink, apex rounded; claw concolorous, 2-3 mm long, glabrous. Medial petal cup-shaped, obovate to broadly elliptic-obovate (Figure 51c,d), broadest at the margin (viewed from the petal apex), apparently not retaining the lateral stamens when the flower opens, 7-8 mm long, 5-6.5 mm wide, 3.5-4.5 mm deep, concolorous with paired petals. Stamen filament bases shortly fused, staminodes f free (Figure 51s). Medial stami- node with filament -3-3.5 mm long, white shading to yellow at the apex, antherode bilobed, yellow, lobes f sessile, curving downward, f obliquely transversely elliptic, -0.7 mm long, connective not or scarcely elongate (Figure 51e). Lateral staminodes with filaments -5.5-6.5 mm long, white shading to yellow apically, antherodes bilobed, f bowtie-shaped, yellow, lobes f sessile, obovate, -0.9 mm long, connective not elongate (Figure 5 If). Lateral stamens with filaments k parallel and held close together, S-shaped, 9-10.5 mm long, glabrous, anthers elliptic to ovate-elliptic, -1 mm long, pollen golden yellow. Medial stamen with filament -7 mm long, anther ovate to ovate-elliptic, saddle-shaped, -1.5 mm long, pollen * concol- orous with lateral anther pollen. Ovary stipitate (Figure 51h), obovate, -2 mm long x 1.5-1.8 mm wide, densely and f uniformly covered with patent (but apically pointing) glandular hairs, apex truncate, dorsal locule prominent, l-ovulate, ventral locules 2-ovulate; style 8-8.5 mm long, arcuate-decurved after the ovary, slightly recurved near the apex, also strongly curved out of the floral midplane, stigma slightly capitate, held at the same level as the lateral anthers and frequently making contact with them. Capsules stipitate (Figure 51q, obovate to obovate-elliptic, 1 20 SMITHSONIAN CONTRIBUTIONS TO BOTANY dehiscent, bivalved, trilocular, 5-7 mm long, 4-5 mm wide, carinate middorsally, stramineus to gray-brown, flecked with dark brown, lustrous, puberulous, apex emarginate, dorsal valve deciduous, dorsal locule very prominent, humpbacked (Figure 51j), 1-seeded or, by abortion, empty, ventral locules each 2-(or, by abortion, 1-)seeded, cells of the capsule wall mainly transversely elongate, occasionally isodiametric to longitudinally elongate. Dorsal locule seed (Figure 51k, l ) slightly humpbacked towards the capsule apex, broadly oblong-elliptic, 2.7-3.2 mm long, 2.1-2.5 mm wide, 1.4-1.8 mm thick, hilum dark brown, in a shallow groove, shorter than the seed, testa buff, smooth, lacking farinose granules and hypha-like filaments. Ventral locule seeds (Figure 51m,n) ovate to deltate or trapezoidal, apical seed thickened and k obtuse apically, basal seed somewhat angular basally, 1.9-2.9 mm long, 1.9-2.4 mm wide, 1.2-1.8 mm thick, hilum dark brown, raised in a groove, slightly extended onto apical and basal surfaces, testa buff, shallowly reticulate-foveolate, with dark brown or grayish farinose granules sparse in the depressions and around the embryotega, denser around the hilum, with hypha-like filaments sparse among the farinose granules around the hilum. HABITAT.-Scrub, track sides, rocky slopes and cliffs in rainfall areas of -400-1000 mm per annum (Wood, in litt.); -600 m. FLOWERING.-Wood (in litt.) reports the plant flowering in mid February, ?the driest time of the year.? Because he collecried mature capsules and seeds, the plants must also have been flowering in January. CHROMOSOME NUMBER.-^ = 26. DISTRIBUTION.-Rare, local endemic in the Yemen Arab Republic (Map 14). SPECIMENS SEEN.-YEMEN ARAB REPUBLIC. Jebel Bura, Hilla [= Hilla], Schweinfurth 289, pap. (G) (mixture with A. forskalii); Regma, Hodjela [= Hojeilah], Schweinfurth 944, p.p. (G) (mixture with A. forskalii); Jebel Bura, just above Hilla, Wood 2616B (EA, F, K, MO, P, US). DISCUSSION This species is named for J.R.I. Wood, whose invaluable recent collections and notes have enabled me to establish the identity of the nineteenth-century gatherings made by Schwein- fruit. Aneilema woodii differs from A. forskalii, its closest relative, in its perennial habit, general absence of long, uniseriate hairs from the adaxial leaf surface, more densely puberulous cincinnus peduncles and axes, more shortly spaced, thinner taxtured, pubescent bracteoles that lack marginal glands, shorter, more pubescent pedicels that are generally more recurved in fruit, puberulous sepals, broader medial petal, larger, more circular antherode lobes, concolorous pollen in all three anthers, and thicker, proportionally broader dorsal locule seeds. My suggestion (Faden, 1975:25 1) that Schweinfurth?s plants, then the sole collections of this taxon, might be closely related to A. petersii (despite the noted disjunct distributions) has proven to be unfounded. From that species, A. woodii differs most conspicuously in the general absence of long, uniseriate hairs from the adaxial leaf surface, more symmetric and generally perfoliate bracteoles that lack long, uniseriate hairs, differently shaped petals, and parallel lateral stamen filaments. In addition, A. woodii is tetraploid, whereas A. petersii is consistently diploid. The recognition of A. woodii as a distinct species was prevented initially by the incomplete condition of the then available material. This problem was compounded by Wood?s collections 2126,2128, and 2616A, which at first I considered conspecific with Schweinfurth?s collections (which were themselves parts of mixed collections with A-forskalii), despite their intermediate morphology between Schweinfurth?s plants and typical A. forskalii. The living plant grown from a fleshy base of the specimen Wood 261 6A failed to produce capsules in spite of repeated, induced self-pollinations, and thus a description of this taxon could not be finished. This problem was finally resolved when Mr. Wood collected yet another living plant for me (as well as dried infructescences, capsules, and seeds) in February 1980 (Wood 2616B). This second plant produccd flowers that differed in some details from those of Wood 2616A, e.g., considerably longer stamens, but, more significantly, it much better matched the original Schweinfurth specimens than did Wood?s first three collec- tions. The new plant was self-compatible and autogamous, and thus it soon produced capsules and seeds, enabling me to complete the description. The problem remaining was the interpretation of Wood?s earlier collections (see below). Aneilema woodii appears to be genuinely rare rather than just inadequately collected. Wood (in litt.) states that although he had an eye open for the plant for two years he found it at only one locality. 6a. Aneilema woodii x forskalii PLATE 3g HABITAT.-In scrubby vegetation; -500-700 m. FLOWERING.-November. CHROMOSOME NUMBER.-h = 52. DISTRIBUTION.-The Yemen Arab Republic (Jebel Bura) SPECIMENS SEEN.-YEMEN ARAB REPUBLIC. Jebel Bura, head of Wadi Hille [= Hilla], Wood 2126 0; Jebel Bura, just above the suq, Hille, Wood 2128 0; Jebel Bura, just above Hilla, Wood 261 6A 0. (Map 14). DISCUSSION The above cited specimens are morphologically intermediate between A. woodii and A. forskalii. The living plant of Wood 2616A is apparently sterile. Its buds frequently fail to open, and NUMBER 76 121 I I I I I I I c 20 15 \ \ 10 0 ?00 200 KIL 1 T RS MAP 14.--Distribution of Aneilemcl woodii Faden. new species (solid and open dots), and A. woodii x forskalii (open dot). self-pollination never leads to fruit set. The pollen was found to be 100% abortive, based on their mostly shrunken appearance and poor staining with cotton blue (in marked contrast to the pollen of A. woodii). The absence of fruits on the field collected specimens of all three collections, despite the presence of large inflorescences on Wood 2128 and 2616A (a combination of characters that is unusual in Aneilemu section Lamprodirhyros), further suggests that these plants are sterile hybrids. The hybrids may be distinguished from A. woodii by the presence of long, uniseriate hairs on the adaxial leaf surface, generally more widely spaced (up to 5 mm apart), glabrous (or with a few hairs basally) bracteoles that have some marginal glands, and less pubescent pedicels (hairy in the apical l/4-l/3 of their length) and sepals. From A. forskalii they differ in their perennial habit and more densely pubescent cincinnus pedun- cles, cincinnus axes, pedicels and sepals. 122 SMITHSONIAN CONTRIBUTIONS TO BOTANY The hybrids may be of relatively recent origin, for Schweinfurth collected only pure A. woo&' and A. forskalii. Wood (in litt.) states that although the two species are identical in their ecology they do not occur together. On Jebel Bura Aneilem woodii and the hybrids are restricted to the valley above Hilla. Aneilem forskalii is apparently absent from that valley, but it occurs in the adjacent valley above Suq As Sabt (Wood, in litt.). Hybridization may have occurred as a result of moderatzly long distance pollen transfer by an insect vector. Alternatively, the annual A. forskalii may formerly have occurred in the same valley as A. woodii and may have died out there, leaving behind A. woodii and the perennial hybrids. Some reproductive isolation between A. woodii and A. forskalii may occur as a result of partially discrete flowering seasons. Wood 26168 was collected in February, during the dry season, when A. forskalii would not be expected to be in flower (unless there were unseasonal rains) (Wood, in litt.). However, at the height of the rains, both species should be expected to be flowering concurrently, which the presence of hybrids ips0 facto demonstrates. 7. Aneilerna benadirense Chiovenda Aneilema benadirenre Chiovenda YBenadirensis'q, Aui 1st. Bot. UNv. Pavia, ser. 4,7:154, pl. 12, 1936.-Cufodontis, Bull. lard. BOL Natl. Belgique, 41, suppl.. 1517. 1971. [Type: Somali Republic, Basso Uebi Scebeli, sud di Merca, 15 Oct 1934, Ciferri 73 (PAV, holotype: Fl', K. is0type.s) (Figure 5 2 ~ Herbs, probably perennial (habit type unknown). Roots unknown. Flowering shoots apparently erect to ascending, to 22 cm long (including the inflorescence) in the specimens. Internodes 2-5 cm long, densely puberulous in longitudinal bands below the fused edges of the sheaths, otherwise sparsely puberulous to glabrous. Leaves spirally arranged, sometimes gradually reduced in size towards the apices of the flowering shoots, sheaths (5-)7-15 mm long, pilose-puberulous, ciliate at the apex, laminae sessile or occasionally very shortly petiolate, narrowly lanceolate-elliptic to lanceolate or ovate- lanceolate, 3.5-8 cm long, 1.5-2.3 cm wide, apex acuminate to acute, margin planar or slightly undulate, scabrid, sparsely ciliate towards the base and on the petiole (when present), upper leaves ciliate to a greater extent than the lower, adaxial surface pilose-puberulous, abaxial surface puberulous. Inflorescences thyrses, terminal and occasionally axillary from the inflorescence bract and uppermost leaves, moderately dense, ovoid to ovoid-ellipsoid, (2.5-)3-5.5 cm long, 2-4 (-5.5) cm wide, with 10-21 cincinni (axillary inflorescences with perhaps fewer), alternate, subopposite or, commonly, subverticillate, ascending to patent (the lower becoming declinate with age). Peduncles 2-4 cm long, puberulous. Inflorescence bract usually supramedial (rarely submedial, subapical or, apical), herbaceous and foliaceous (or rarely, membranous and very reduced). Inflorescence axis puberulous. Cincinni to 2.5 cm long and 5-flowered. Cincinnus bracts membranous (rarely the lowermost herbaceous and f foli- aceous), appressed to the cincinnus peduncles or patent, ovate to ovate-elliptic, 1.5-2(-2.3) mm long, usually glandular near the apex, glabrous. Cincinnus peduncles relatively uniform within the inflorescence, exceeding the cincinnus bracts, (3-)4-7(-8) mm long, puberulous. Cincinnus axes puberu- lous. Bracteoles attached (1.5-)2-3(-3.3) mm apart, f herbaceous, slightly eccentrically to nearly symmetrically cup-shaped, perfoliate, 1.5-1.7 mm long, 0.9-1.2 mm high, with a very prominent subapical gland and smaller glands along the margin, glabrous except for a few hook-hairs occasionally present at the base. Flowers perfect and staminate. Pedicels (5-)6-7.5(-9) mm long, ascending and straight or arcuate in flower, in fruit, recurved (uniformly or predominantly above the middle) 180'-270?, persistent, glabrous below the middle, puberulous above. Sepals prominently glandular near the apex, with smaller glands along the margins; medial sepal apparently lanceolate or lanceolate-elliptic, 2.6-3.5(-4.3) mm long, glabrous, subapical gland deeply bilobed; lateral sepals apparently f elliptic, 2.7-4(-4.5) mm long, glabrous or with a few hook-hairs towards the base, subapical gland unlobed or slightly bilobed. Paired petals -6-8 mm long, limb 4.5-6.5 mm long, to 6 mm wide (Chiovenda, 1936); claw 1-1.5 mm long. Medial petal apparently slipper-shaped, 4.5-6(-7) mm long, -2-2.5 mm deep. Lateral stamen filaments fused basally to medial stamen filament and to the lateral staminode filaments, medial staminode free. Medial staminode vestigial (always?), represented by a filament less than 1 mm long, antherode not developed. Lateral staminodes with filaments 2.5-3 mm long, yellow apically, antherodes bilobed, yellow, lobes shortly stipitate, obovate-cuneate to transversely elliptic, 0.6-0.8 mm long, connective not elongate. Lateral stamens with filaments 6-7 mm long, f S-shaped, anthers lanceolate- elliptic, (0.9-)1-1.2(-1.4) mm long, -0.5 mm wide. Medial stamen with filament 3.5-4(-4.3) mm long, anther ovate- oblong, saddle-shaped, 1.3-1.8 mm long, 0.8-1.2 mm wide. Ovary substipitate, -1 mm long, densely covered with minute glandular hairs (hook-hairs apparently absent), dorsal locule very prominent, l-ovulate, ventral locules each 2-ovulate; style 7-8 mm long, stigma slightly capitate. Capsules substipitate, probably obovate-elliptic to obovate, dehiscent, bivalved, trilocular, 3.4-4 mm long, -2 mm wide, grayish tan with dark brown spots on the dorsal locule, lustrous, sparsely puberulous, apex emarginate, dorsal valve deciduous (tardily?), dorsal locule very prominent, 1 -seeded, ventral locules each 2-(or, by abortion, l-)seeded, cells of the capsule wall transversely elongate. Dorsal locule seed (Plate 6e) hemispherical, 1.8-2 mm long, 1.3-1.4 mm wide, 1-1.25 mm thick, hilum dark brown, arising from a shallow groove in the flat, ventral surface, much shorter than the seed, testa yellowish buff or orange-buff, f smooth, lacking white-farinose granules and hypha-like filaments except around the hilum where both are abundant. Ventral locule seeds (Plate 6e) ovate to subtriangular, apical seed rounded apically, basal seed angular NUMBER 76 123 FIOURE S2.-Holotype of Aneilema benadirenre Chiovenda (Ciferri 73-PAV). basally, 1.3-1.6(-1.7) mm long, 1.35-1.4 mm wide, 0.9-1 mm thick, hilum dark brown, raised within a groove, not extended onto apical and basal surfaces, testa orange-buff, shallowly scrobiculate on all surfaces, sparsely white-farinose in most of the depressions, densely so around the hilum and embryotega, hypha-like filaments common among the farinose granules. HABITAT.-DuneS and littoral (Ciferri 73). probably just above sea level. FLOWERING.-The single collection is from October and has flowering and fruiting specimens. Unless the capsules and seeds mature unusually rapidly in this species, the largest inflorescences must have been flowering in September. CHROMOSOME Nvmm.-Unknown. DISTRIBUTION.-^ Somali Republic (Map 15). SPECIMENS SEEN.-SOMALI REPUBLIC. BENADIR: Basso Uebi Scebeli [River], sud di Merca, Ciferri 73 (lT?, K, PAV). DISCUSSION The holotype consists of the ten shoots illustrated in the plate accompanying the type description. They are, however, somewhat differently arranged than in the figure (Figure 52). All of the shoots lack bases and none has roots. 124 SMITHSONIAN CONTRIBUTIONS TO BOTANY MAP 1s.-Distribution of Aneilema benudireme Chiovenda (open circle), A. calceolus Brenan (solid squares), and A. tanueme Faden (solid circles). The measurements given above frequently differ from those of Chiovenda (1936). In most cases the discrepancies are minor; in several they are significant and appear to be the result of Chiovenda?s either not having noted the full range of variation shown in the material (distance between bracteoles, sepal length) or perhaps not having measured the best pressed flowers (lateral staminode filament length). For several characters (cincinnus peduncle length, medial anther length and width, dorsal locule seed length and width) Chiovenda?s dimensions are quite wrong and should be ignored. Nearly all of the shoots of the specimens have their longest leaves at the base, suggesting that even longer leaves may have been present on the lower, uncollected portions. Such leaves could well have been longly petiolate, for regularly petiolate lower leaves and sessile or subsessile upper leaves are common in other Aneilema species. NUMBER 76 125 In the larger inflorescences of the specimens, usually all of the cincinni are subverticillate (or a few subopposite). In the smaller ones, however, the arrangement of the cincinni is less regular, generally some of them being alternate. The abaxial leaf surfaces, inflorescence axes, cincinnus peduncles and axes, and occasionally the pedicels all appear to be covered with a mixture of hook-hairs and patent, glandular hairs of similar size. The swollen tips of approximately 15 of the seemingly glandular hairs were gently rubbed with a moistened camel?s-hair brush and found to really be hook-hairs in which the terminal hook had become covered with adherent foreign matter, giving it the appearance of a clavate gland. It is likely that all of the apparent gland-tipped hairs on the above structures can be accounted for in this manner. Some of Chiovenda?s interpretations of floral characters are inaccurate. His description of the flowers as sometimes having two large-anthered stamens, in addition to the two smaller- anthered ones, was evidently based on two flowers pressed one upon the other (visible under a dissecting microscope in the top, right hand shoot in Chiovenda (1936, pl. 12) and Figure 52. Only one such anther is ever present. Chiovenda?s designation of the stamens as five to six is likewise based on this erroneous observation of the medial stamen. The three stamens and lateral staminodes (his ?five stamens?) are always well developed. The medial staminode, apparently always vestigial, seemingly was overlooked by Chiovenda. The flower color, ?greenish- white or yellowish-white,? as cited from the collector by Chiovenda, is almost certainly incorrect. Several important characters cannot be determined with certainty from the specimens. The rather long lateral stamen filaments and their apparent strong curvature suggest a well-developed stamen-retention mechanism. The dorsal cap- sule valve appears to become detached at the base even before the capsules are fully mature. While this is certainly an artifact of specimen preparation, it may indicate that this valve is normally deciduous in mature capsules. The floral and capsular structure of A. benudirense clearly demonstrate that it belongs to section Lumprodithyros. Within the section, its regularly thyrsiform inflorescences suggest affinities with A. forskalii, A . sebitense, A. recurvatum, A. petersii, A. indehiscens, and A. woodii. It differs from A. forskalii and A. sebitense in its less widely spaced bracteoles and much smaller capsules and seeds, from A. recurvatum in its more strongly dimorphic seeds, and from all three in its more recurved fruiting pedicels. It can be separated from A. petersii and A. indehiscens by its more symmetric bracteoles, from A. woodii by its smaller seeds, and from all three by its glabrous or subglabrous bracteoles with marginal glands, glabrous or subglabrous sepals and smaller capsules. It further differs from all six of these species in its generally more numerous cincinni per inflorescence, usually glandular medial sepal margins (rarely present in A. recurvatum and A. forskalii) and vestigial medial staminode. The closest relationship of A. benadirense is with A. tanaense. Both species have cincinni predominantly whorled, bracteoles and sepals with marginal glands, fruiting pedicels often recurved more than 180?, capsules with deciduous or possibly deciduous dorsal valves, and seeds strongly dimor- phic. The general absence or rare vestigial presence of a medial staminode in A . tanuense also suggests an affinity with A. benadirense. The latter may be distinguished from A. tanaense by its generally longer leaves, larger, always thyrsiform inflorescences with more numerous cincinni, longer cincinnus peduncles, more widely spaced bracteoles, glabrous or sub- glabrous sepals, lack of hook-hairs on the ovaries and capsules, and larger capsules. The single collection of Aneilemu benadirense and a recent gathering of A. petersii subsp. pullidiflorum are the sole collections of section Lamprodithyros from the Somali Republic. In view of the diversity of m a in this section recorded from the Kenya coast south of Lamu, further exploration in the poorly collected region north of that town and in the coastal areas of southern Somali Republic may well yield additional taxa. 8. Aneilema tanaense Faden Aneilemu lanuensc Faden, Bothalia, 15:98, 1984. [?Lpe: Kenya, Tana River District, Garissa-Malindi road, 16 km N of junction for Ganen, -2O085, 40?04?E, 15 Jan 1972, Gillerr 19528 (US. holotype; B. BR, EA. FT. K. MO, PRE, isotypes).] Aneilemu clurkei Rendle, J. Linn. Soc., Bot., 30, pl. 34: fig. 8 tantum, figs. 7, 9-12 et descr. excl. 1895.1 Aneilemu culceolus Brenan, Kew Bull., 15:223,1%1 [pro Gregory s.n]. Annual (rarely perennial) herbs (habit types IB, IC, Figure 1; Plate 39. Roots fibrous, produced only at the base and lower nodes. Primary shoot erect or ascending, much branched at the base, 15-35 cm tall, lateral shoots decumbent, or prosaate initially and then ascending. Internodes (1-)2-5(-7) cm long, green, puberulous. Leaves spirally arranged on main shoot, distichous, at least initially, on lateral shoots, sometimes becoming spirally arranged again towards the inflorescences, sheaths (1-)3-7 mm long, puberulous or pilose-puberulous, apex ciliate, laminae sessile or shortly petiolate, gradually reduced towards the terminal inflorescence on the main shoot, lanceolate or lanceolate-elliptic to ovate, 1-6.5 cm long, 0.8-2.5 (-3) cm wide, apex acute to obtuse or rounded (in smaller leaves), margin usually planar, scabrid with patent to apically directed prickle hairs, sparsely ciliate basally or on the petiole (when present), both surfaces lustrous, puberulous, adaxial with longer uniseriate hairs than the abaxial, sometimes mottled with maroon. Inflorescences terminal on the main and major lateral shoots and on very reduced lateral shoots, ultimately produced from nearly all nodes; reduced lateral shoots frequently perforating the sheaths; larger inflorescences thyrses (Figure 536; Plate 3j) , moderately dense, broadly ovoid, 1-2(-3) cm long, 1.5-3(-5) cm wide, with up to 8 cincinni, subopposite or subverticillate 126 SMITHSONIAN CONTRIBUTIONS TO BOTANY 1 I b d e f FIGURE 53.-Aneilema tunaenre Faden: u, habit; b, inflorescence; c, bracteole; d, perfect flower, front view; e, perfect flower, side view;f, medial sepal, showing small marginal glands; g, lateral staminode; h, andcecium, top view, medial staminode omitted, gynoecium removed; i, ovary and base of style, side view; j . capsule attached to cincinnus, side view; k, dorsal capsule valve, dorsal view; I, dorsallocule seed, ventralview; m, dorsal locule seed, dorsal view; n, ventral locule seed, ventral view; 0, ventral locule seed, dorsal view (u, from Fuden & Fuden 7411053; b-e,g,i, from Fuden & Fuden 77/582;f,h,+, from Gillett 19973). (Bars = 20 mm for a; 5 mm for b; 2 mm for d,e; 1 mm for c,f-k; 0.5 mm for 1-0.) NUMBER 76 127 (occasionally some alternate), ascending: smaller inflores- cences consisting of l-several, clustered cincinni, lacking a distinct axis and not clearly thyrses. Peduncles of larger inflorescences (1-)2-5(-8.5) cm long, puberulous. Inflores- cence bract supramedial to subapical, herbaceous and foli- aceous. Inflorescence axis, when developed, puberulous. Cincinni to 2.2 cm long and 10-flowered (to 3.5 cm long and 17-flowered in cultivation). Cincinnus bracts herbaceous or membranous, patent or appressed to the cincinni, lanceolate to lanceolate-elliptic, ovate or oblanceolate-elliptic, 1-10 mm long, lower ones often puberulous, upper glabrous or sub- glabrous. Cincinnus peduncles, except the lowermost, gener- ally exceeding the cincinnus bracts, 2-5 mm long, puberulous. Cincinnus axes puberulous. Bracteoles attached 1-2(-2.5) mm apart, f herbaceous, symmetrically or eccentrically cup- shaped, perfoliate, 1.4-1.8 mm long, 0.8-1 mm high, prominently glandular near the apex and with smaller glands along the margin, puberulous in the basal l/2 or, more commonly, only at the base (Figure 53c). Flowers perfect and staminate, odorless, (9-)lo-14.5 mm wide (Figure 53d,e; Plate 3k,l). Pedicels (4-)5.5-lo(-11) mm long, erect to slightly arcuate in flower, k uniformly recurved in fruit for their entire length 180'-270"(-360") (Figure 53j), often spirally twisted as well, persistent, puberulous. Sepals prominently glandular near the apex, puberulous except for glabrous margins: medial sepal lanceolate-ovate to ovate, 2.5-3 mm long, 2-2.2 mm wide, subapical gland distinctly bilobed, smaller glands also generally present along the margin near the base (Figure 53f); lateral sepals elliptic to ovate- elliptic or ovate-orbicular, 2.8-3 mm long, 2-2.5 mm wide, subapical gland unlobed, marginal glands absent. Paired petals (4.2-)6.5-8 mm long, 4.8-7 mm wide, limb ovate, (2.7-)5-6 mm long, pink or pale lilac (RHS colors: 77D-78D, Fuden & Fuden 7411053; 84B-C, 84C, Fuden & Fuden 7411185), apex acute to obtuse or rounded, claw 1.5-2 mm long, white, glabrous. Medial petal slipper-shaped, obovate-elliptic to suborbicular or subquadrate, broadest below the margin (viewed from the apex), retaining the lateral stamens when the flower opens, 4.7-6 mm long, 3-4.7 mm wide, 3-3.6 mm deep, concolorous with the limbs of the paired petals (Figure 23j-I; Plate 3k,l). Stamen filaments fused basally and also shortly fused to lateral staminode filaments (Figures 27u, 53h). Medial staminode usually absent, rarely vestigial and repre- sented by a short filament. Lateral staminodes with filaments 3.3-4 mm long, pink or white, shading to yellow above the middle, antherode bilobed, yellow, lobes shortly stipitate to sub-sessile, obovate-cuneate to transversely elliptic or broadly reniform, 0.3-0.8 mm long, connective not at all to scarcely elongate (Figure 53g). Lateral stamens with filaments k parallel in the basal l/2, then sharply divergent (Figures 27u, 53h), 5.5-6.5 mm long, S-shaped, geniculate and decurved in the middle, glabrous, anthers elliptic to ovate, 0.7-1.2 mm long, 0.5-0.9 mm wide, pollen yellow or orange-yellow. Medial stamen with filament 3.5-4 mm long, anther ovate, saddle- shaped, 1-1.5 mm long, 1.1-1.3 mm wide, entirely yellow, pollen yellow or orange-yellow (concolorous with lateral anther pollen). Ovary substipitate (Figure 53i), obovate, 1-1.2 mm long, 0.8-1 mm wide, densely and uniformly covered with patent, glandular hairs (mixed with hook-hairs along the lateral sutures), apex slightly emarginate to truncate, dorsal locule very prominent, 1-ovulate, ventral locules each 2-ovulate: style 5.5-6.5 mm long, arcuate-decurved for most of its length, also gently curving out of the floral midplane (rarely not), stigma capitate, usually held below the anthers and in front of them (Plate 3k,l). Capsules (Figure 53jk) substipitate to shortly stipitate, obovate (to ovate), dehiscent, bivalved, trilocular, (2.4-)2.7- 3(-3.4) mm long, (1.1-)1.5-2.1 mm wide, gray to tan, sometimes with a dark gray band around the base of the dorsal locule, frequently with irregular brown spots and stripes, lustrous, puberulous, apex emarginate, dorsal valve deciduous, dorsal locule very prominent, often with a seed, ventral locules each 2-(or, by abortion, l-)seeded, cells of the capsule wall transversely elongate. Dorsal locule seed (Figure 531,m; Plate 6g) hemispherical, 1.1-1.6 mm long, 1-1.4 mm wide, 0.85-1 mm thick, hilum dark brown, arising from a shallow groove and flush with the flat to slightly concave ventral surface, much shorter than the seed, testa tan, smooth, lacking farinose granules and hypha-like filaments except around the hilum. Ventral locule seeds (Figure 53n,o; Plate 6g) subtriangular, apical seed rounded apically, basal seed angular basally, 1.2-1.4(-1.7) mm long, 1.2-1.3(-1.5) mm wide, 0.8-0.95 mm thick, hilum dark brown, raised within a groove, not extended onto apical and basal surfaces, testa gray or grayish tan, shallowly scrobiculate on all surfaces, sparsely white- farinose in many of the depressions and around the embryo- tega, densely so around the hilum, frequently some hypha-like filaments present among the farinose granules. HABITAT.-Deciduous or semi-evergreen bushland and thickets: seasonally water-logged, gray-brown, clayey alluvium with patches of sand (Tana River District), or better drained soil (Kwale District): 10-250 m. FLowERING.-~owering specimens have been Seen from January, February, July, and August. The size of the largest inflorescences in Gillett 19528 indicates that the species must have been flowering in December. Field observations show that the flowers open 0830-0900 hrs and fade 1300-1330 hrs (Figure 25). CHROMOSOME NUMBER.-" = 13,2n = 26. DISTRIBUTION.-Lower reaches of the Tana River and Kwale District, Kenya (Map 15). SPECIMENS EEN.-KENYA. KWALE: Mombasa-Nairobi road, 2.5 km towards Mombasa from turnoff of Maji ya Chumvi Railway Station, 3'493, 39'20'E, Fuden & Fuden 771582 (BR, EA, F, FT, K, MO, P, PRE, US, WAG): Lungalunga-Ramisi road, -6.5 km from Vanga, 4"37'S, 39"10'E, Fuden & Fuden 771738 (EA, F, K, US): [Kilifi District on label], 5 km Maji ya Chumvi-Mackinnon Road, 128 SMITHSONIAN CONTRIBUTIONS TO BOTANY 3'48'S,39'20%, Gilbert & Rankin 4834 (EA). TANA RIVER: 105 km N of Malindi on Garsen road, Andrews in EA15070 (EA): Galole-Garsen road, 8 km towards Garsen from turnoff to Wenje, l052'S,4Oo05'E, Faden & Faden 74110.53 (C, EA, F, K, MO, US); Garsen, 2?16'S,40007'E, Faden & Faden 7411064 (EA, MO); Malindi-Garsen road, 0.8 km towards Garsen from turnoff to Kibusu, 2"21'S, 4Oo07'E, Faden & Faden 7411171 (BR, EA, K, MO, PRE, US); Garsen-Malindi road, 1.6 km towards Malindi from crossing of Lugga Buna, 2'23'S, 4Oo07'E, Faden & Faden 7411179 F A , K, MO); Garsen- Malindi road, 1.5 km towards Malindi from turnoff to Oda, 2"32'S, 4Oo07'E, Faden & Faden 7411185 (B, BR, EA, FT, K, MO, US); Garissa-Malindi road, 16 km N of junction for Garsen, -2'O8'S,4O004'E, Gillett 19528 (B, BR, EA, K, MO, P, PRE, US); 105 km N of Malindi on Garsen road [Kilifi District on label], Gillett 19532 (EA, K); Galole-Malindi road, 16 km S of spot height 106, 2'14'S, Gillett 19973, cultivated at Missouri Botanical Garden (BM, EA, K, MO); Lake Dumi, 13 Feb 1893, Gregory s.n. (BM). DISCUSSION The taxonomic confusion of A. tanaense with A. clurkei is explained in Faden (1984). Of the six figures that accompany the type description of A. clarkei (Rendle, 1895, pl. 34: figs. 7-12), Rendle's figure 8 clearly does not belong to that species. There are two separate, unmixed, Gregory collections with the same data at the British Museum (Natural History). One of them is the type of A. clarkei and, significantly, it has on it all of the drawings published by Rendle except his figure 8. The second Gregory specimen was cited by Brenan (1961) as A. calceolus but it is here treated as A. tanaense. The first inflorescences produced by plants of A. tanaense are invariably distinct thyrses with a short but clear inflores- cence axis. Later inflorescences consist of fewer cincinni (sometimes only one) and may lack an axis, giving them a very strong resemblance to inflorescences of A. calceolus. They differ from the latter chiefly in having more elongate cincinni with more widely spaced bracteoles that have glandular margins. In Tana River District, A. tanaense is restricted to a peculiar type of alluvium found along the lower reaches of the Tana River. This soil type is referred to as "black cotton soil with sand" by local agricultural officers (Riyamy, personal commu- nication). The trees and shrubs most commonly associated with A. tanaense are Acacia bussei, other Acacia spp., Commiphora campestris, Dobera sp., Combretum hereroense, Grewia tenax, and Sansevieria powellii. In Kwale District, A. tanaense occurs in two habitats: (1) termite mounds and thicket edge, the thickets containing trees and shrubs such as Euclea sp., Diospyros consolatae, Manilkara sp., Carissa bispinosa, Haplocoelum inoploeum, and Erythroxylum emarginatum as well as succulents; and (2) bushland and thickets dominated by Acacia spp. and succulents. The only two perennial populations are from the last habitat near Maji ya Chumvi. Aneilema tanaense is transitional between the species of section Lamprodithyros that regularly have thyrsiform inflores- cences (species 1-7) and those that never have them (species 9-13). More precisely, it is intermediate between A. benadi- rense and A. calceolus in lamina length, inflorescence form, cincinnus number per inflorescence, distance between bracte- oles, and degree of development of marginal glands on the sepals. It agrees with both of them in fruiting pedicel curvature, medial staminode development, capsule form, and seed dimorphism. Its differences from A. benudirense are discussed under that species. From A. calceolus it differs in habit and in the characters given in the key. The close relationship between A. tanaense and A. calceolus is supported by preliminary crosses made in cultivation that suggested a high interfertility between these species. 9. Aneilema calceolus Brenan Aneilema calceolw Brenan, Kew Bull., 15223, 1961 [Gregory s.n. excl.]. [Type: Tanzania (Tanganyika on label), near Kange Limestone Gorge, 13 Nov 1956, Milne-Redhead & Taylor 7285 (K. holotype; BR, EA. isotypes).] Annual (rarely perennial) herbs (habit type IIA2, Figure 2; Plate 3m). Roots fibrous. Vegetative shoots repent, much branched, forming mats to 1 m or more in diameter, flowering shoots ascending, to 15(-20) cm tall. Internodes (1)2-7(-9.5) cm long, puberulous. Leaves distichous, sheaths 2-6 mm long, puberulous and generally with a few long, uniseriate hairs, particularly towards the base, ciliate at the apex, laminae sessile or shortly petiolate, ovate to ovateelliptic or lanceolateelliptic, 1-3.5(-5) cm long, 0.5-1.5 (-2) cm wide, apex acute or occasionally obtuse, often mucronulate, margin usually planar, scabrid with spreading to apically directed prickle hairs or short uniseriate hairs, sparsely ciliate basally or on the petiole (when present), both surfaces lustrous, puberulous, also pilose with uniseriate hairs, those on the adaxial surface longer than those on the abaxial. Inflorescences terminal on main and lateral shoots, consist- ing of 1-4(-6) cincinni that appear fasciculate in the a i l s of 1 or 2, alternate or subopposite, sessile, foliaceous bracts or reduced leaves: inflorescences actually reduced thyrses (see discussion below), to -2 cm long and wide. Peduncles to 7.5 cm long, puberulous. Inflorescence bract apical or subapical, herbaceous, foliaceous, often only slightly smaller than the leaf at the node below. Inflorescence axis f not developed. Cincinni (when more than 1) subopposite or subverticillate, ascending or becoming arcuate with age, to 7 mm long and 14-flowered (to 11 mm long and 25-flowered in cultivation). Cincinnus bracts herbaceous and green (the lowermost generally so) or membranous and whitish or brownish, lower ones patent, others appressed to the cincinni, lanceolate-elliptic to ovate, up to 13 mm long (much smaller in the upper cincinni), not evidently glandular (at least the lower ones), puberulous to glabrous. Cincinnus peduncles not developed. Cincinnus axes NUMBER 76 129 very short, generally completely covered by overlapping bracteoles, glabrous. Bracteoles attached 0.2-1(-1.4) mm apart, membranous, slightly to strongly eccentrically cup- shaped, perfoliate, 1.1-1.9 mm long, 0.4-1 mm high, inconspicuously glandular near the apex, glabrous or sub- glabrous. Flowers perfect and staminate, odorless, (10-)12-15 mm wide (Plate 3n,o). Pedicels 7-11 mm long in flower, to 13(-16?) mm in fruit, +_ erect in flower, sometimes slightly arcuate near the apex, k uniformly recurved in fruit for their entire length generally -270?, frequently somewhat spirally twisted as well, persistent, puberulous except at the base (Figure 35c). Sepals glandular near the apex, puberulous; medial sepal lanceolate to ovate-elliptic, 3-4 mm long, 2-2.5 mm wide, gland inconspicuously bilobed, lateral sepals ovateelliptic to ovate, 3-4 mm long, 2.5-2.7 mm wide, gland inconspicuous (sometimes absent?), unlobed. Paired petals 5-9 mm long, 3.5-7.5 mm wide, limb ovate, lilac (RRHS colors: 76B, Organ in EA14970, cultivated, 76C, Faden & Faden 741324, cultivated, 84C, Faden & Fuden 74/1069), apex rounded or obtuse to subacute, claw 1-3 mm long, whitish, glabrous. Medial petal slipper-shaped, elliptic to obovate- elliptic, somewhat keeled medially, broadest below the involute margin (viewed from the apex), retaining the lateral stamens when the flower opens, 5-7.5 mm long, 2.5-4 mm wide, -2.5-3.7 mm deep, concolorous with the limbs of the paired petals (Figure 23m-o; Plate 3n,o). Lateral stamen filaments fused basally to medial stamen filament and sometimes to lateral staminode filaments (Figure 27b). Medial staminode absent, or rarely represented by a short filament. Lateral staminodes with filaments 2.5-3.5 mm long, white basally, shading to yellow above the middle, antherodes bilobed, yellow, lobes stipitate, transversely elliptic to broadly ovate, subreniform or obovate-cuneate, 0.4-0.7 mm long, connective not elongate. Lateral stamens with filaments parallel or divergent in basal l/3, more strongly divergent in middle l/3, less divergent to almost parallel in apical l/3 (Figure 27b), 4-8 mm long, S-shaped, geniculate and decurved near the middle, glabrous, anthers elliptic to ovate, 0.7-1.25 mm long, 0.5-0.75 mm wide, pollen white. Medial stamen with filament 3-5 mm long, anther lanceolate-ovate to ovate, saddle-shaped, 1.4-2 mm long, 1.2-1.5 mm wide, pollen yellow. Ovary substipitate, obovate, 0.9-1 mm long, 0.9-1.1 mm wide, densely and uniformly covered with patent, glandular hairs (mixed with hook-hairs along the lateral sutures), apex truncate to slightly emarginate, dorsal locule very prominent, l-ovulate, ventral locules each 2-ovulate; style 5.5-7 mm long, very gently arcuate-decurved at least l/2 of its length, then recurved towards the apex, also gently curved laterally out of the floral midplane, stigma slightly capitate, held in front of the anthers and vertically between the levels of the medial and lateral anthers. Capsules (Figure 35c,d) substipitate, obovate, dehiscent, bivalved, trilocular, (2.3-)2.5-3.6 mm long, (1.4-)1.9-2.5 (-2.8) mm wide, f carinate middorsally, light brown or pale grayish brown, spotted and striped with dark brown, lustrous, puberulous, apex emarginate, dorsal valve deciduous, dorsal locule very prominent, usually with a seed, ventral locules each 2-(or, by abortion, 1-)seeded, cells of the capsule wall transversely elongate. Dorsal locule seed (Plate 6f) hemispheri- cal, 1.2-1.6 mm long, 1-1.4 mm wide, 0.8-1.1 mm thick, hilum dark brown, arising from a shallow groove, much shorter than the seed, testa tan, smooth, lacking farinose granules or with farinose granules sparse around the embryotega and f dense around the hilum where sometimes mixed with hypha- like filaments. Ventral locule seeds (Figure 38; Plate 6f) subtriangular, apical seed obtuse to rounded apically, basal seed angular basally, 1.2-1.6(-2) mm long, 1.1-1.5 mm wide, 0.7-1 mm thick, hilum dark brown, raised within a groove, not extended onto apical and basal surfaces, testa tan, pinkish tan or grayish tan, shallowly scrobiculate on all surfaces, usually white-farinose in most of the depressions and around the embryotega and hilum, frequently some hypha-like filaments present among the farinose granules (Figure 38d). HABITAT.-D~~ or moist, lowland evergreen forest or thicket; growing on Kambe limestone, often in shallow soil, in the Tanga area of Tanzania; partial or dense shade; 10-450 m. FI.OwERING.-FlOWering specimens have been seen from June, July, and September-November. Field observations of Fuden & Faden 7411069 indicate that the flowers open 0900-1OOO hrs and fade 1300-1330 hrs (Figure 25). CHROMOSOME NUMBER.2n = 26. D I S T R I B U T I O N . ~ O ~ S ~ ~ Kenya and Tanzania (Map 15). SPECIMENS SEEN.-KENYA. KWALE: Lungalunga-Ramisi road, 1 km before turnoff to Kinango, 4'32'30"s. 39'05'30"E, Faden & Fuden 771378 (BR, EA, F, K, MO, PRE, US, WAG). TANA RIVER: Garsen-Witu road, near Nyangoro Bridge, 2'18's. 40'20'E, Faden & Faden 74/1069 (EA, K, MO, US); same locality, Fuden & Faden 771565 (EA, F, K, US). TANZANIA. TANGA: Amboni Cave near Tanga, 5'04'S, 39'03'E. Botany Students in DSM13.51 (EA); 5 mi SE of Ngomeni, Drummond & Hemsley 3561 (B, BR, EA, K, LISC, S); Amboni Caves -2 km E of Tanga, -5"06'S, 39003'E, Faden & Faden 741324 (EA, K, MO, US); Kange Gorge, -5 km E of Tanga, -5'06'S, 39'01'E, Faden & Faden 741333 (EA, F, K, PRE, US, WAG): Usambara foot-hills, Maramba Forest, Faulkner 4045 (K); 2 mi N of Tanga, Harris & Procter 1018 @A); near Kange Limestone Gorge, Milne-Redhead & Taylor 7285 (BR, EA, K): Kiomoni Quarries near Tanga, cultivated in Nairobi, Organ in EA14970 (EA). DISCUSSION Gregory s.n. cited by Brenan (1961) as this species differs from the Tanzanian specimens in being more erect and in having shorter pedicels, more elongate cincinnus axes that are puberulous where exposed, and especially in its bracteoles with small glands along the margin and generally sparsely puberu- 130 SMITHSONIAN CONTRIBUTIONS TO BOTANY lous. It also has small marginal glands on the medial sepal. This specimen is treated above as A. tanaense. Aneilema calceolus is generally annual. Only completely or nearly completely dead plants of Faden & Faden 741324, 741333, and 771378 were found in the field, and 741324 and 771378, when grown from seed, behaved as annuals, i.e., after flowering and fruiting they ceased vegetative growth and were unable to root from cuttings. However, the northernmost, isolated population in the Tam River District of Kenya (Faden & Faden 7411 069 and 771565) is definitely perennial and has been cultivated for more than ten years, The inflorescence of A. calceolus is easily interpreted as a reduced thyrse in which the inflorescence axis and cincinnus peduncles are suppressed and the cincinnus axes are very contracted. When the lowermost cyme appears to be separated from the others, a short inflorescence axis thus seemingly present, careful observations of the sequence of flowering usually indicates that this cyme does not flower until after the one above it. This ?lowermost? cincinnus is therefore an entire inflorescence consisting of a single cincinnus axillary in the inflorescence bract, while the cincinnus above that one is morphologically the true lowermost one of the terminal inflorescence. In inflorescences that consist of a subopposite pair of terminal, leaf-like bracts, each subtending a cincinnus, again the cincinnus of the larger, lower bract does not flower until after that of the upper one. The lower cincinnus in this case must also be considered an entire inflorescence axillary in the inflorescence bract. The true terminal inflorescence also consists of a single cincinnus. In A. calceolus the stamens are retained by the medial petal longer than in any other species of section Lamprodithyros (Figure 25). Field observations of Faden & Faden 7411069 indicate that the stamens are held for an average of two and one-third hours or approximately half of the flowering period. The habitat requirements of A. calceolus appear to fall within the range of those for A. clarkei, with which it is sympatric. The two have never been found growing together, although each has been collected with or near A. taylorii on very similar outcrops of Kambe limestone. There is no obvious reason for their apparent mutual exclusion; competition for space should be considered as a possible factor. Aneilemu calceolus differs from A. clarkei, with which it was contrasted by Brenan (1961), by the former?s having usually an annual habit, blunter laminae, often several cincinni per inflorescence, fruiting pedicels uniformly recurved for their entire length and puberulous for most of their length (versus recurved and puberulous only at the apex), much smaller, lilac petals, usual absence of a medial staminode, smaller capsules with a deciduous dorsal valve, and smaller, more dimorphic seeds. Aneilema calceolus also differs from A. lamuense and A. succulentum in inflorescence structure, pedicel curvature and pubescence, medial petal shape (Figures 23m-0, 24&) and lack of a medial staminode. It is clearly less closely related to these species than they are to each other or to A. clarkei. The most closely related species to A. calceolus is A. runaense. They agree in flower and capsule structure, pedicel curvature and pubescence, and seed dimorphism. Reduced inflorescences of A. tanuense are very similar to inflorescences of A. calceolus. The two species differ in habitat, habit, and in the characters given in the key. 10. Aneilema clarkei Rendle Aneilema clarkei Rendle. J. Linn. Soc., Bot., 30430, pl. 3 4 figs. 7,9-12 [non fig. 81, 1895.Schumann in Fhgler, Pflanzenwelt Ost-Afrikas C, 136, 1895.4arke in Thiselton-Dyer. FI. Trop. Africa, 8:73, 1901.-Brenan, Kew Bull., 15224, 1961 [in adnot.]. [?Lpe: Kenya, Tana [River District], Lake Dumi, 13 Feb 1893, Greg0rys.n. (BM)] Perennial herbs (habit type IIA2, Figure 2). Roots fibrous. Vegetative shoots procumbent to repent, much branched; flowering shoots erect to ascending, generally unbranched, 15-30(-40) cm tall. Internodes (1-)lS-7(-13) cm long, glabrous to puberulous. Leaves distichous (the 2 ranks at an obtuse angle to each other), usually gradually reduced apically on the flowering shoots, sheaths 2.5-10 mm long, puberulous and generally sparsely pilose with uniseriate hairs, particularly at their bases, apex ciliate, laminae petiolate, lanceolate to linear-lanceolate, lanceolate-elliptic or ovate, 1.5-5(-7.8) cm long, (0.4-)0.8-1.5(-2.5) cm wide, apex acuminate to acute, margin usually planar (occasionally somewhat undulate), scabrid, sparsely ciliate basally or on the petiole (when present), both surfaces lustrous, pilose-puberulous, the uniseri- ate hairs of the adaxial surface longer and generally fewer than those of the abaxial surface. Inflorescences terminal on the flowering shoots and also on short, non-perforating shoots produced from the uppermost 1-3 nodes on the flowering shoots, each inflorescence consisting of a short, solitary cincinnus subtended by and partially enclosed in a subopposite pair of herbaceous (or occasionally the upper or both membranous) bracts, the larger, lower one the inflorescence bract, the upper the cincinnus bract (Figure 9; Plate 3p). Peduncles -0-2.5 cm long, puberulous. Inflorescence bract terminal, herbaceous or rarely membra- nous. Cincinni to -5 mm long and 10-flowered. Cincinnus bracts herbaceous or rarely membranous, lanceolate, often falcate, ascending, -5.5-15 mm long (including the sheath) x 3-5 mm wide, eglandular, both surfaces puberulous, adaxial also densely pilose, margins longly ciliate, at least towards the apex, Cincinnus peduncles not developed. Cincinnus axes very short, completely covered by overlapping bracteoles, glabrous (Figure 3%). Bracteoles attached up to 0.2 mm apart, membranous, slightly to strongly eccentrically cup-shaped, perfoliate, -3-5 mm long x 2 mm high, apparently eglandular, lower (outer) ones sparsely puberulous with uniseriate hairs and often hook-hairs, upper (inner) ones apparently glabrous. Flowers perfect and staminate, odorless, (12.5-)15-20 mm wide (Plate 4d). Pedicels 7-11.5 mm long, amate-erect to NUMBER 76 gently S-shaped in flower, in fruit, strongly recurved (-180'- 270') only near the apex and becoming much thickened in that region (Figure 35e), persistent, greenish white, puberulous generally only near the apex, rarely extending to below the middle. Sepals glandular near the apex, glands unlobed; medial sepal lanceolate-ovate to ovate-deltate, 3.2-5 mm long, 2.5-4.5 mm wide, glabrous, or sparsely puberulous basally; lateral sepals ovate-elliptic to broadly ovate or ovate-orbicular, 3.3-5.5 mm long, 2.7-4.5 mm wide, sparsely puberulous medially and basally. Paired petals (7.5-)9-12 mm long, (5.5-)7-11 mm wide, limb ovate to ovate-deltate or ovate- subreniform, -5-8 mm long, lilac to lavender (RHS colors: 87D, Faden & Faden 7411215, field & cultivated, & Faden & Faden 72/226A, cultivated; 85C, 87C-D, Evans & Muikweki 51, cultivated; 88D, Faden et al. 711633, cultivated), apex obtuse to rounded, margin crenulate to subentire, claw 2.5-4 mm long, white or concolorous with limb (always paler at the base), glabrous. Medial petal cup-shaped, ovate, broadest at the margin (viewed from the apex), which is frequently slightly revolute, nearly always retaining the lateral stamens when the flower opens, 6-9 mm long, 3-7 mm wide, 2.5-5 mm deep, concolorous with the limbs of the paired petals, paler basally, (Figure 24ec; Plate 4d). Lateral stamen filaments fused basally to medial stamen filament and to the lateral staminode filaments (Figure 284. Medial staminode (rarely absent) filament 0.7-2 mm long, entirely yellow or white, or pinkish purple basally, antherode bilobed, yellow, lobes sessile, ovate to suborbicular, reniform or transversely elliptic, 0.4-0.8 mm long, connective slightly elongate. Lateral staminodes with filaments 3-5 mm long, lilac to lavender basally, shading to yellow in the apical l/3-%, antherodes bilobed, yellow, lobes subsessile to shortly stipitate (rarely sessile), transversely elliptic to broadly ovate-cuneate or obovate-cuneate, 0.7-0.8 (-1) mm long, connective slightly to not at all elongate. Lateral stamens with filaments divergent (Figure 28a), (5.5-)6.5-8.5 mm long, S-shaped, geniculate and decurved near the middle, glabrous, anthers lanceolateelliptic to broadly elliptic or ovate, 1-1.6 mm long, 0.5-1.1 mm wide, pollen white. Medial stamen with filament 3.5-5.5 mm long, anther ovate-elliptic to oblong, saddle-shaped, 1.4-2.5 mm long, 1-1.6 mm wide, pollen yellow. Ovary substipitate, obovate to obovate-oblong or obovateelliptic, (1-)1.5-1.9 mm long, (1-)1.1-1.3 mm wide, densely and uniformly covered with patent, glandular hairs, usually mixed with few to many hook-hairs (best seen in fruit) (Figure 42iJ), apex truncate to rounded or slightly emarginate, dorsal locule very prominent, 1-ovulate, ventral locules each 2-ovulate; style 7.5-11 mm long, nearly straight or gently arcuate-decurved for most of its length, recurved near the apex, also gently curved out of the floral midplane, stigma capitate, violet, held in front of the anthers and generally below them. Capsules (Figure 35e) substipitate, obovate to oblong- elliptic, dehiscent, bivalved, trilocular, 3.6-4.5 mm long, 2.3-3 mm wide, tan or grayish tan, with small, dark brown 131 spots, particularly around the base of the dorsal locule, lustrous, puberulous, apex truncate to emarginate, valves persistent or dorsal valve tardily deciduous, dorsal locule very prominent, 1-seeded (or occasionally empty), ventral locules each 2-(or, by abortion, l-)seeded, cells of the capsule wall transvarsely elongate. Seeds (Plate 6h) broadly elliptic, 1.8-2 mm long, 1.4-1.5 mm wide (dorsal locule seed) or ovate to subhiangular, with apical seeds rounded apically and basal seeds angular basally, 1.5-1.7(-2.1) mm long, 1.5-1.6 mm wide (ventral locule seeds), 0.9-1.1 mm thick, hilum dark brown, promi- nently raised within a groove, not extended onto apical and basal surfaces, testa light brown, shallowly scmbiculate on the dorsal surface, sparsely white-farinose in some of the depres- sions, densely so around the embryotega and hilum. HABITAT.--hwland evergreen forest (sometimes on Kambe limestone), Brachystegia forest or woodland on tan sand, river slopes, Hyphaene-Albiria scrub on sand, and moist thickets; dense or partial shade; 0-220 m. FLowEruNG.-Flowering specimens have been Seen from January to March, May, and July-November. Field observa- tions of three populations indicate that the flowers open 0845-0930 hrs and fade 1200-1300 hrs. CHROMOSOME NUMBER.-2n = 78. DISTRIBUTION.