SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 76 

The Morphology and Taxonomy of 

Aneilema R. Brown (Commelinaceae) 

Robert B. Faden 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1991 
ABSTRACT 

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 ................. la . 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 hem we would hst hem 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 ornamentals. 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 circumscription 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 Pedunculosa). 

2. Preserved material and living plants only from cultivation: acuminatum, biflorum, neocaledonicum (section Aneilema); gillettii (section Amelina); longicapsa, pusillum subsp. 

gypsophilum, pusillum, and thulinii (section Somaliensia); 

forskalii, woodii, woodii x forskalii (section Lamprodithyros); 

lanceolatum subsp. lanceolatum, pomeridianum (section Brevibarbata). 

3. Preserved material and living plants from the field and 

often also from cultivation: somaliense (section Somaliensia); 

brenanianum, rendlei, taylorii (section Rendlei); aequinoctiale, ephemerum, hockii, johnstonii, longirrhizum, nyasense 

(section Amelina); calceolus, clarkei, indehiscens (all subspecies), lamuense, petersii (both subspecies), recurvatum, sebitense, succulentum, tanaense, zebrinum (section Lamprodithyros); 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% methylene 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 Amelina); 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 available) 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. benadirense) 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 photographed 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 whenever possible. However, the large number of taxa surveyed has 

necessitated an examination of numerous herbarium specimens. 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 recognize 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); longtrailing, 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 

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NUMBER 76 5 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

m 

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NUMBER 76 7 

shoots with terminal and axillary inflorescences, a characteristic 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 ascending 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 decumbent, fibrous-rooted rhizomatous, and tuberous-rooted, ascending 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 specialized, 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, dispermum, dregeanum, silvaticum, umbrosum (section Brevibarbata); leiocaule (scction Pedunculosa). 

Pcrcnnials with spccialized, Subterranean storage organs: 

sclerocarpum, siliculosum (scc Lion Aneilema); gillettii, hockii, 

johnstonii, longirrhizum, plagiocapsa (scction Amelina); brenanianum, new species (section Rendlei); longicapsa, new 

species, obbiadense, pusillum, somaliense (section Somaliensia, new section); homblei, lanceolatum, macrorrhirum, paludosum 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); angolense, arenicola, brunneospermum, mortonii, paludosum 

subsp. paludosum and subsp. pauciflorum, schlechteri (section 

Brevibarbata); chrysopogon, hirtum, minutiflorum, nicholsonii, 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 specimens 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. pallidijlorum, 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, indehiscens, minutiflorum, pedunculosum, spekei, tanaense, termitarium, 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.Sequence 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 exclusively 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 populations 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. aequinoctiale, 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 development 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 (dorsiventral 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 SMITHSOW 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. welwitschii (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 vemation. 

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 subverticillate 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. ’ 

L1 

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 inflorescencebearing 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 ails anywhere along the 

stem, The term secondary inflorescence has sometimes been 

employed for an inflorescence-shoot arising from the inflorescence 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. pedunculosum and A. spekei, it completely encloses and protects the 

young, pre-flowering inflorescence. In A. clarkei, A. succulenrum, 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 

tt 

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 

ped. f* --r( 

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 of 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 somewhat 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. 

T f.l. 

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 characters 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 polygamomonoecious 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 polygamomonoecious 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 

f5 

=Pistillate flowers 

Perfect flowers 

Istaminate 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. 

12 3 4 5 6 7 8 9 10 

Day in Flowering Sequence 

FIGURE lrl.-Idealized flowering sequence in an inflorescence of a polygamomonoecious 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 sometimes 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, Iongicapsa, longirrhizum, minutijlorum, nicholsonii, nyasense var. 

brevisepalum, obbiadense, pedunculosum, petersii (both subspecies), pomeridianum, pusillum (all subspecies but gypsophilum), 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 polygamomonoccious 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 

2051 

12 13 14 

a perfect flowers 

0 staminate flowers 

1 I 

15 16 17 18 

Lateral stamen filament length (mm) 

10 I5 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 inflorescence 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 arrangement 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. Curvature 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 Lamprodithyros 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. neocaledonicum, A. umbrosum (both subspecies), and A. taylorii (frequently). 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; ij, 

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 Lamprodithyros, 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 le,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 subcordate. 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: Coleotrype, Commelina, Floscopa, and Murdannia. In Aneilema 

they are present only in some species of sections Amelina 

(aequinoctiale, ephemerum, johnstonii, nyasense) and Brevibarbata (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 li) and A. ephemerum, they are inconspicuously 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 taxonomically. 

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 staminodes 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 Lamprodithyros, 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 differentiated 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 populations). 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 staminode 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); gi, 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); jl, 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 Aneilem 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, 2b4, 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 occasionally 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 ma. The antherodes of all 

staminodes are generally yellow except in A. acuminatum 

(Plate lab), 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. neocaledonicum (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 li), 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 28~). 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-df,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. 741 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 li,j) they are distributed along most of 

its length, and in A. gillettii (Plate lf) 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 dehiscent (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 

lj) is an exccption, for its lateral filaments are 

minutcly bearded, but its medial anther is reduced. Aneilemu 

gilletfii (Plate le,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 lhJ, 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 (cd): 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 31~9). 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 spinules 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. a945 (from Faden & Faden 74/237): b, A. somalienre C.B. 

Clarke, ~3400 (from Faden & Faden 74/923); c, A. beniniense (P. de Beauvois) Kunth. ~2890 (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, ~3400, b, polar view, ~3400, 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, campylotropous 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 righthanded 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 34.4tyle curvature types in Aneilemo: a, straight; b, arcuatedecurved; 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 (<go"), e.g., A. aequinoctiale and A. hockii, 

or widely (9Oo-18O0), e.g., A. acuminatum, A. johnstonii, and 

A. welwitschii. In some species the dorsal valve is deciduous 

(sometimes tardily or only occasionally so) and serves as a 

disseminule, carrying with it the dorsal locule seed, which is 

usually developed in these species (occasionally lacking 

through abortion). Deciduous dorsal valves have been noted in 

A. sclerocarpum (section Aneilema), A. plagiocapsa (section 

Amelina), A. pusillum and A. somaliense (section Somaliensia), 

all species of section Lamprodithyros to varying extents, except 

A. zebrinum, and A. pedunculosum (tardily, section PeduncuThe capsules are regularly bivalved except in some 

specimens of A. acuminatum. These plants, sometimes segregated as A. papuanum, have trivalved capsules with three equal 

losa). 

valves and up to five seeds in the dorsal locule. The prominent 

terminal ridge on the dorsal locule of A. indehiscens subsp. 

indehiscens is also sometimes partially dehiscent middorsally 

and the capsules thereby partially trivalved. 

Capsule pubescence differs from ovary pubescence in being 

less dense due to the increased size of the developing fruit. 

During the two- to three-week development period of the 

capsule, the hairs remain functional. At maturity, when the 

capsule turns from green to its ripe color, the thin-walled 

glandular hairs collapse. When only glandular microhairs are 

present on the ovary (section Rendlei species and A. johnstonii 

of section Amelina), the mature capsules appear glabrous. 

Seed number per capsule locule is frequently less than the 

ovule number due to the abortion of ovules or failure of some 

of them to set seed. The only species in which regular abortion 

of particular ovules appears to occur is A. plagiocapsa. All four 

ventral ovules abort, while the strongly laterally compressed 

dorsal locule produces a single, large seed (-6 mm long), the 

largest seed known in the genus. 

The shape of the cells of the outer wall of the capsule was 

first used as a taxonomic character in Aneilema by Brenan 

(1952), who separated A. aequinoctiale from A. hockii partly on 

this character. Brenan (1961) also used capsule wall cell shape 

to distinguish A. dregeanum from A. schlechteri. Faden (1984) 

showed that this character could be used to separate three 

closely related species, A. schlechteri, A. brunneospermum, and 

A. arenicola. 

The cells of the capsule wall in Aneilema vary from 

transversely elongate to isodiametric or rarely longitudinally 

elongate. Transversely elongate capsule wall cells are the most 

common type, occurring in all species of sections Aneilema, 

Rendlei, Somaliensia, Lamprodithyros, and Pedunculosa, most 

species of section Brevibarbata, and in A. gillettii, A. hockii 

(Figure 36a), A. johnstonii, and A. longirrhizum of section 

Amelina. The capsule wall cells are isodiametric in A. 

aequinoctiale (Figure 36b), A. ephemerum, A. nyasense, and A. 

plagiocapsa (section Amelina) and in A. mortonii and A. 

paludosum (section Brevibarbata). Only A. brunneospermum 

(section Brevibarbata) has longitudinally elongate capsule wall 

cells (Faden, 1984, fig. 1). Because they frequently vary in 

shape on different parts of the capsule, cells from the same part 

of the capsule must be used when comparing different taxa. For 

standardization, only the cells of the dorsal valve halfway 

between the capsule base and apex and halfway between the 

lateral and middorsal sutures have been described. 

In addition to cell shape, the surface of the capsule wall cells 

shows interspecific differences when viewed with the scanning 

electron microscope. The cells of A. hockii capsules (Figure 

36a) are minutely papillose while those of A. neocaledonicum 

(Figure 36c) are smooth. 

Many capsule characters appear to be useful taxonomically, 

the most important being capsule shape, dimensions, dehiscence, locule number, apex shape, pubescence, seed number 

per locule, and shape and surface features of the capsule wall 
NUMBER 76 37 

C 

FrGuRE 35.4apsules of Aneilem species. A. petersii (Hasskarl) C.B. Clarke subsp. petersii (a.6,): a, capsule 

attached to C~~~MUS, side view; b, dorsal valve, dorsal view (from Faden & Evam 70181 9). A. calceolur Brenan 

(c,d): c, capsule attached to chchnus; d, capsule, side view (from Faden & Faden 74324). A. clarkei Rendle: 

e, inflorescence with capsule (from Faden et al. 71/633). (Bar = 1 mm.) 

cells. The capsules are measured in the same manner as the 

ovaries. 

Seeds: Aneilema seeds have a linear hilum and prominent, 

lateral embryotega (Figures 37-39; Plates 5-7). They are 

usually dorsiventrally compressed, with the ventral surface 

being planar to slightly concave, and the dorsal surface slightly 

to strongly convex. The seeds range in length from about 1 to 

6 mm, and can reach 3 mm in width and 2 mm in thickness. 

Seed shape and size vary according to the species. The shape 

and size of any particular seed are also determined by its 

position in the capsule and by the number of other seeds 

developing in the same locule. The apical end of the apical 

ventral locule seed is often very different in shape from the 

basal end of the basal ventral locule seed (Plates 5a,b,df, 6,7f) 

and thus these seeds are commonly distinguishable from each 

other. When more than two seeds are present in each ventral 

locule, the medial seeds are also distinct in shape from the 

apical and basal seeds (Plates Sa,b,d, f, 6,7b, f ). The seeds from 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 36.Scanning electron micrographs of Aneilema capsule walls: a, A. 

hock2 De Wildeman, x-174 (from Faden & Faden 7411180); b, A. 

aequinoctiale (P. de Beauvois) G. Don. x-170 (from Hall s.n.1 c, A. 

neocaledonicum Schlechter, X-166 (from Bierhorst in Kew Accession No. 

000-69.12064). The views are of the dorsal valve approximately halfway 

between the capsule apex and base. The capsule apex is in the direction of the 

arrow. 

one-seeded ventral locules are always longer and somewhat 

differently shaped from those of two- or more seeded ventral 

locules of capsules from the same population (Plates 5a, 6e,f, 

7d). 

In addition to the relatively minor dimorphism or polymorphism among the seeds within the ventral locules, there is 

sometimes a much greater differentiation between dorsal and 

ventral locule seeds. Because dorsal locule seeds are almost 

always solitary and occur only in species having two or more 

ovules (and generally seeds) in each ventral locule, a difference 

in size and shape between the seeds of these locules is to be 

expected. Only such differences in shape and sometimes length 

between dorsal and ventral locule seeds occur in some species, 

e.g., A. indehiscens, A. recurvatum, A. zebrinum (Plates 6b,c, 

7a), and A. grandibracteolatum. 

A more profound dorsal and ventral locule seed dimorphism 

occurs in other species. These differences may include (in 

addition to seed shape and length) seed width and thickness, 

dorsal surface convexity, hilum prominence, embryotega color, 

testa surface pattern and color, and farinose granule abundance 

and distribution. The most strongly differentiated dorsal and 

ventral locule seeds occur in some species of section 

Lamprodithyros: A. petersii, A. calceolus, A. tanaense (Plate 

Q,fg), and A. forskalii. Moderately dimorphic dorsal and 

ventral locule seeds also occur in A. nicholsonii (section 

Pedunculosa, Plate 76). It is noteworthy that within section 

Lamprodithyros, the most dimorphic dorsal and ventral locule 

seeds occur in those species in which the dorsal capsule valves 

are most regularly deciduous, while the least dimorphic seeds 

are found in the species with generally persistent capsule 

valves. The degree of seed dimorphism is probably related to 

seed dispersal mechanisms in the species of this section. 

The hilum usually lies in a longitudinal groove that is open 

at the apical and basal ends of the seed (Figure 38c). Rarely, 

both ends of the groove are cut off by ridges as in A. 

grandibracteolatum, A. pusillurn, A. richardsiae (Plates 5c,h, 

7e), and A. longicapsa. In a few species, mostly in section 

Amelina, the hilum is flush with the ventral surface and not at 

all in a groove (Figure 37b; Plate 15d,e,j). The hilum may be 

confined to the ventral surface or extend onto the apical and/or 

basal surfaces. 

The embryotega is circular to elliptic in outline and has a 

central prominence. Micropores are visible in scanning electron 

micrographs (Figures 37c, 38d). The embryotega is pushed off 

or to one side by the cotylcdon of the germinating seed. 

The testa is composed of usually clearly discernible, 

polygonal cells, each in contact with five to eight cells (Figure 

37d). Occasionally the cells are less distinct (Figure 39). The 

testa may be completely smooth, e.g., some populations of A. 

hockii (Figure 37a; Plate 5~3, but more commonly it is 

variously pitted, grooved, or ribbed. Very regular patterns 

sometimes occur: alveolate (e.g., some populations of A. 

hockii, Plate 5e); reticulate-foveate (e.g., A. minutiforum and 

A. umbrosum, Plate 7c,g); sulcate (A. pusillurn, Plate 5h); or 

ribbed (e.g., A. paludosum, Morton, 1966:447, fig. 6). 

Generally, however, the pits or furrows are irregular, sometimes discrete, sometimes coalescing. The general term used 

here for such patterns is scrobiculate (e.g., A. taylorii and 
NUMBER 76 

FIGURE 37.4canning electron micrographs of seeds of Aneilem hockii De Wildeman: a, dorsal view, X-38: b, 

ventral view showing hilum (arrow), X-38; c, lateral view showing embryotega (arrow), x-38; d, dorsal surface 

cells, X-197; e, ventral view showing farinose granules (arrow), x-80. All from Faden & Faden 741180. 

39 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 38.-Scanning electron micrographs of seeds of Aneilema calceolus Brenan: a, dorsal view showing 

farinose granules (arrow), X-56; b, farinose granules, x-315; c, ventral view, X-56; d, lateral view showing 

hypha-like filaments (arrow), x-56. All from Faden & Faden 741324. 
NUMBER 76 

FIGURE 39.4canning electron micrographs of seeds of Aneilema neocaledonicwn Schlechter: a, dorsal view, 

x-43; b, ventral view, X-43; c, region of hilum, X-1 15. All from Bierhorst in Kew Accession No. 000-69.12064. 

41 
42 SMITHSONTAN CONTRIBUTIONS TO BOTANY 

ventral locule seeds in A. calceolus, Plates 5a, 6f). 

Various particles are often present on the testa. The most 

common are whitish or light brown (rarely black) granules that 

I have called farinose granules (Figures 37e, 38a,b). These 

occur in all sections of the genus except section Aneilema. The 

farinose granules are often confined to the depressions in the 

testa and to the regions around the hilum and embryotega. 

Occasionally they may be so abundant that they cover most of 

the seed surface, as in A. longirrhizum (Plate 59. In A. 

paludosum and A. dregeanum subsp. mossambicense (Faden, 

1984). a continuous whitish layer, perhaps derived from fused 

farinose granules, sometimes covers the entire seed except for 

the hilum. 

Small filaments resembling fungal hyphae sometimes occur 

among the farinose granules (Figure 38d). These have been 

observed only in species of section Lamprodithyros. Their 

nature has not yet been established. 

As has been shown for Tripogandra by Handlos (1970, 

1975) and for various species of Floscopa by Brenan (1952, 

1961), seeds are often very useful for specific characters in the 

Commelinaceae. This has been confirmed for Aneilema. The 

combination of seed shape, dimensions, dimorphism or 

polymorphism, testa color, and surface pattern is apparently 

sufficient to distinguish nearly all species. Further scanning 

electron microscope studies may provide a basis for recognition 

of all species. Among the three dimensions, seed width is the 

most useful. 

Variation in seed characters between populations of the same 

species is usually not very great. However, the seeds of the two 

subspecies of A. petersii and A. dregeanum, are probably 

distinguishable from each other. Seeds from West African and 

East African collections included in A. dispermum by Brenan 

(1952) and Morton (1966) are very different, and it is clear that 

these geographically isolated populations will have to be 

separated at the subspecific or specific level when they are 

studied in detail. Populations of A. hockii from the Tana River 

District of Kenya have very characteristic seeds (Figure 37; 

Plate 5j) that readily separate them from all other populations 

throughout the extensive range of this species (Plate 5e). 

Further study of A. hockii is required in order to determine the 

taxonomic status of this local variant. 

The terminology used for the surfaces of the seed is as 

follows: ventral, the surface bearing the hilum; dorsal, the 

surface opposite the ventral; apical and basal, the surfaces 

facing the apex and base of the capsule, respectively; lateral, 

the surfaces, one of which bearing the embryotega, directed 

toward the dorsal and ventral surfaces of the capsule. Seed 

length is the distance between the apical and basal surfaces; 

width, the distance between the lateral surfaces; and thickness, 

the distance between the ventral and dorsal surfaces. Seed 

shape is the linear outline viewed from the dorsal surface. 

Anatomy of Aneilema 

INTRODUCTION TO ANATOMY 

Previous anatomical studies that have included Aneilema 

species are Staudermann (1924)-A. aequinoctiale, hairs; 

Bruckner (1926)-A. beniniense, all vegetative parts: Rohweder (1963)-A. beniniense, stems and flowers; Tomlinson 

(l966,1969)-A. acuminatum, A. aequinoctiale, A. beniniense, 

A. hockii, A. lanceolatum (including A. subnudum), A. 

setiferum, A. umbrosum, and A. rebrinum (as Ballya zebrina), 

all vegetative parts; Rohweder (1969)-A. aequinoctiale and A. 

beniniense, ovaries and seeds; and, Walker (1974)-A. beniniense, epidermis. This study is of limited scope, intending to 

survey the variability of certain anatomical characters within 

Aneilema and to determine which attributes, if any, are 

sufficiently diverse to merit more detailed examination in the 

future. Only floral hairs have been investigated more thoroughly. 

STEMS.-Aneilema stems are always succulent. Transverse 

sections of internodes of nine species, representing all sections 

of the genus, show a great similarity to one another. Their 

general structure is like that illustrated for Commelina imberbis 

by Tomlinson (1969:46, fig. 12). An outer cortex, varying in 

thickness according to species, is separated from an inner 

central cylinder by a continuous one-celled endodermis-like 

layer. The cortex lacks vascular bundles. Starting from the 

outside it comprises a thin cuticle, one-celled epidermis 

containing stomata, a zone of numerous alternating patches of 

collenchyma and chlorenchyma, and a narrow band of 

chlorenchyma, continuous with the subepidermal chlorenchyma patches. The central cylinder is composed of ground 

parenchyma with scattered vascular bundles. 

LEAVES.-Transverse sections of the leaves show a thin, 

smooth cuticle, an upper and lower epidermis usually composed of more or less isodiametric cells (cells elongate at right 

angles to the leaf surface occur in A. succulentum and A. 

zebrinum), and a relatively thin mesophyll composed of a 

single, adaxial palisade layer and an abaxial spongy mesophyll 

generally composed of three to four cell layers. A hypodermis 

is present only in the midrib region as two small, adaxial 

patches, one on either side of the midrib. The hypodermis is, in 

part, usually more than one cell layer thick. Collenchyma is 

present between the midrib and the lower epidermis (Tomlinson, 1969:40, fig. 10-1). The form and position of the 

hypodermis and collenchyma result in a more or less W-shaped 

mesophyll in the midrib region. A submarginal vein is present, 

but margin collenchyma and sclerenchyma are lacking. Silica is 

absent. 

RAPH1DEs.-Raphides are present in most parts of the plant, 

including the anthers and root tips. According to Tomlinson 

(1966, 1969). the raphides occur in elongate raphide-sacs 

arranged in longitudinal series to form raphide canals. 

Fresh-cut shoots exude a sap that is sometimes slightly milky 
NUMBER 76 43 

due to the presence of abundant raphides. 

STOMATA.-stomata occur on all green parts of the plant. 

On the leaves they are always more abundant on the abaxial 

surface than the adaxial, on which occasionally they may be 

lacking. The stomata typically have six accessory or subsidiary 

cells, two lateral pairs and one terminal pair (Tomlinson, 

1969:30, fig. 6). The stomata on the sepals are often less regular 

in accessory cell number, two to seven accessory cells having 

been observed. 

HAIRS.-Hairs may occur on all aerial parts of the plant with 

the exceptions of the staminodes and medial stamen. Among 

the vegetative and reproductive organs, only the leaves and 

sepals always have hairs. The following hair types are 

recognizable (terminology partly after Tomlinson, 1966): 

1. Glandular microhairs (Figures 40a-c,i, 42a,b,h): threecelled, appressed hairs, each with small basal and medial cells 

and an elongate, glandular apical cell. These may occur on all 

parts of the plant where hairs are found except the stamen 

filaments. They are particularly dense on the adaxial surface of 

the sepals, where they have been found in all 26 Aneilema 

species that have been checked for this character. Tomlinson 

(1966) reported that glandular microhairs on the vegetative 

organs vary little within each species. In contrast, the glandular 

microhairs, when present on the abaxial sepal surface, are 

always much smaller than those of the adaxial surface. 

Glandular microhairs are probably present in all Aneilema 

species . 2. Hook-hairs (Figures 40d,e,j, 41g, 42j,m): hairs in which 

the terminal cell is hooked. As defined by Tomlinson (1966), 

hook-hairs are always two-celled. In Aneilema that is true of all 

non-floral hook-hairs as well as some floral ones. However, 

some of the latter, which are also termed hook-hairs here, may 

have as many as five cells. Two-celled hook-hairs of two or 

more size classes occur on the vegetative and/or reproductive 

parts in some species of sections Amelina and Brevibarbata and 

some or all species of section Pedunculosa (Figure 40d,e). 

Hook-hairs occur on the leaves of all Aneilema species and in 

the inflorescences of all but a few species. 

3. Prickle hairs (Figure 40g): two-celled hairs with a broad 

basal cell and short, stiff apical cell not exceeding the basal cell 

in length. These occur on the leaf margins of most Aneilema 

species. They are typically apically directed and make the 

margins scabrid. They sometimes also occur on the lamina 

surfaces, but are rare on other parts of the plant. They grade 

imperceptibly into the next hair type. 

4. Uniseriate or acicular hairs (Figure 4Of): two- to 

several-celled hairs with sharply pointed apical cells, when 

two-ccllcd, the apical cell longer than the basal cell. Thcse are 

the cilia on the sheath apex and frequently of the petiole and 

lamina margins of many Aneilema species. Uniseriate hairs are 

also frequent on the sheath and lamina surfaces. They are 

uncommon on the internodes and in the inflorescences and 

flowers. Their occurrence in the inflorescences is sometimes a 

diagnostic specific character. The distinction used here between two-celled uniseriate and prickle hairs is quite arbitrary. 

Further studies may suggest that the two should be separated at 

a different level. 

5. Uniseriate-eglandular hairs (Figure 4 la- c,e,h): two- to 

six-celled hairs differing from uniseriate hairs in having 

rounded apical cells. These are found only on lateral stamen 

filaments, occurring in all species (as far as is known) of 

sections Brevibarbata and Pedunculosa and in A. nyasense of 

section Amelina. 

6. Uniseriate-glandular hairs (Figures 40h, 4 If, 42cg,i,k,l,n,o): three- to six-celled hairs in which the apical cell is 

glandular. The three-celled ones (Figure 424 differ from 

glandular microhairs, from which they are probably derived, in 

having much stronger basal cells and small apical cells. 

Uniseriate-glandular hairs are confined to the flowers, being 

common on the ovaries and rare on the petals (only in A. 

aequinoctiale) and lateral stamen filaments (only in A. gillettii). 

These hairs may be further subdivided on the basis of whether 

or not the apical cell is capitate. 

7. Branched hairs (Figure 416): four- to five-celled, eglandular hairs known only from the stamen filaments of A. 

nyasense. 

The diversity of floral hairs suggested that these might be 

diagnostic at the specific or sectional ranks. The floral hairs of 

19 species, representing all seven sections of the genus, were 

studied in detail. The distribution of the different hair types 

within these species is shown in Tables 3-5. The results 

support the morphological and cytological data indicating that 

sections Aneilema, Rendlei, Lamprodithyros, and Pedunculosa 

are natural, while sections Amelina and Brevibarbata may not 

be natural. Aneilema somaliense of section Somaliensia is 

separable from all of the above by a combination of characters, 

but no light has been shed on the naturalness of section 

Somaliensia as a whole. Floral hairs also appear to be useful at 

the specific level in many instances. 

Vegetative hairs and anatomy seem to be of more limited 

use, but detailed studies of epidermal patterns and of the 

distribution of hairs on the lamina may be worthwhile in certain 

critical species complexes, e.g., A. beniniense, A. umbrosum, 

and A. welwitschii. 

It is not yet clear whcther Aneilema can be distinguished 

anatomically from all other genera of Commelinaceae. It 

appears that at least the closely related genus Rhopalephora is 

probably not separable on anatomical characters alone. 

Cytology of Aneilema 

Chromosome numbers and chromosome morphology of 

Aneilema species were studied extensively by Faden and Suda, 

and their findings, as well as a summary of all previous counts, 

were included in Faden (1975). Several counts from this survey 

were published in Faden (1983b). Along with data accumulated 

since, the complete results will be published separately. 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

b 

.p 

.,?. :" .:.:. . 

..:>.. :.:. ... ... .+'. 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 microhairs): 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 ma. The basic numbers obtained for 

each section are included in the descriptions of the sections. 

Chromosome counts for species of sections Rendlei, Somaliensia, 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 Cglycosides 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- Uniseriatemicrohairs 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 biologically 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; Syrphidae: 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 individuals), 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 combination 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. clarbi 

A. recwvatwn 

A. rebrinwn 

A. beniniense 

A. brunneospermwn 

Section Pedunculosa 

A. hirtum 

A. termitarium 

Section Brevibarbata 

UniseriateGlandular 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 population 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, pedunculosum, and spekei) all have bearded filaments, and they include 

all three species of section Pedunculosa (nicholsonii, pedunculosum, 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. umbrosum) 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 glabrousfilament 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 Aneilemperhaps 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 temperature, 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 selfincompatible; 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 “Acknowledgments’’) 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 discontinuities. 

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, karyotypes, 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 Sea 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 endemics) 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. Morton, 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 SMITHSOW CONTRTBUTIONS TO BOTANY 

hh~ 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 Lamprodithyros 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 Commelinaceae. Only five other genera (Buforrestia, Commelina, Floscopa, 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, suggesting 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) Floscopa 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 synonymous with Commelina by most workers until Kunth (1843) 

recognized it. Between 1843 and 1881 a number of generic 

segregates were described: Dictyospermum Wight and Dichoespermum 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 Aneilema 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 Tricarpelema 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 sometimes 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 Rhopalephora 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 abovementioned 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. StefaniniPaoli 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 [oh. 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.] 

hlinu 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 (inflorescence 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, deliquescent, 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 Aneilema 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 Dicarpellaria 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 rugosetuberculate, 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 distribution 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 aequinoctide (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 cupshaped, 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 generally 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, reticulate-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, uniseriate 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 inconspicuously 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 maroonspotted. 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, bivalved, 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 

hlA~ 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 microhairs) 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 variable 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 occasionally 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 occasionally membranous, generally very reduced, occasionally foliaceous. 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 lanceolateelliptic 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 (sometimes all curvature types within the same inflorescence), 

persistent, glabrous. Sepals with a subapical, moderatcly 

prominent gland, glabrous; medial sepal elliptic to lanceolateelliptic 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 

dM.) 
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, arcuatedecurved 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 lanceolateelliptic 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, whitefarinose 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. 

DISTRIBUTION.~O~S~~~ SE Kenya, coastal and subcoastal 

NE Tanzania (Map 5). 

SPECIMFNS SEEN.- 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 handwriting as “Fimbine.” S.A. Robertson (personal communication) 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 

. . . 70 

\ 

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; j8 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 seminibus 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.) 
70 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, herbaceous, 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 Usambara 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 sufficiently 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 occasionally 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, supramedial or occasionally subapical, herbaceous or less commonly 

membranous, bract-like or occasionally foliaceous. Inflorescence 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 oblongelliptic, 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 lanceolateovate, 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 lp) 4.5-5.5 mm long, question-markshaped, 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 lanceolateelliptic, 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, oblongelliptic, 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 lp), 

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 = 16.h = 32. 

DISTRIBUTION.-SW Ethiopia N and SE Kenya, NE Tanzania (Map 5). 

SPECIMENS SEEN.-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 (US). 

LAIKIPIA~ORTHERN FRONTIER: Near meeting point with 

Samburu District, Nolaleta River, -Oo44'N, 36'572, A. Fuden 

16185 (US). MACHAKOS: Kibwezi, Dummer 5020 (K); Bushwackers, 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 mono [Kwamkono], 

Archbold 713 (K); Kideleko, 7 km S of Handeni, Archbold 

1897 (EA). LUSHOTO: Western Usambara Mts., LushotoMombo 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 decurved, 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 puberulous, 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, persistent, 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 supramedial, 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 47~9; 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 ovateelliptic 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, stramineous, 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 SEEN.-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 transpiration 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. longirrhizum, 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 fewerseeded 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 plerumque 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 cincinni. 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 glandularpuberulous, 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 ma, 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, commonly 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 hit, 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 lanceolate-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 orangeyellow. 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) trapezoidal, 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; hero (Meta-Gafersa), Cufodontis 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 collected 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 significant. 

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 Actiniopteris 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 lanceolatis 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 lanceolate-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 peduncles, becoming reflexed with age, lanceolate to lanceolateelliptic (2.3-)3.5-5.8 mm .long, glandular near the apex, 

glabrous to sparsely puberulous. Cincinnus peduncles relatively 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 SEEN.-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 characters, 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 filaments, 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] 

Ov.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, subterranean, 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 foliaceous. Distance between inflorescence bract and lowest 

cincinnus 1-2 cm. Inflorescence axis puberulous with uniformlength 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, asymmetric, 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 SEEN.-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 subterranean 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 linearolanceolatis (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 plerumque 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, disarticulating 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 lanceolateelliptic), 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 moderately 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, puberulous. 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, ascending to reflexed, lanceolate to lanceolate-oblong or ovateoblong, 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 subapically, 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 spreading 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.<entral Somali Republic. 

SPECIMENSEEN.-SOMALI REPUBLIC. 17 km S of 

Mugakori, 3’57’N, 46’11’E, Gillett & Beckett 23289 (EA, K, 

US); 5 km W of Mukwakori, 4’07’N. 46’05’E, Gillett et al. 

22609 (K, US); Sultanato di Obbia, tra Scermarca, Hassam e 

Tobungab. Puccioni & Stefanini 579 [6341 F); 48 km on 

Maas-Bulo Burti road, -4’06’N, 45’462, Thulin & Warfa 

4614 (K, UPS); 5-10 km along the road to Maas from the 

turning N of Bulo Burti, -3’57”. 45’362. Thulin & Warfa 

5341 (UPS). 

DISCUSSION 

When I first examined Puccioni & Stefanini 579 [634], the 

holotype and then sole specimen of A. obbiadense var. 

angustifolium, I concluded that it could be encompassed within 

A. obbiadense without recognition as a distinct variety (Faden, 

1975). The typical variety of the species was then also known 

from a single specimen. On the basis of more recent collections 

of both varieties, as well as the opinion of Gillett (in litt.), the 

only modern collector to have seen both in the field, I am now 

convinced that they represent distinct species. The differences 

in leaf shape, cincinnus peduncle length, and bracteole spacing 

given in the key appear consistent and give the plants quite a 

different facies. The species also seem to occur at different 

elevations. Aneilema longicapsa generally has shorter and less 

recurved fruiting pedicels than A. obbiadense. In all probability, additional characters will be found to separate these species 

when living material of A. obbiadense can be obtained and 

floral characters determined. 

The recently collected Thulin & Warfa 5341 has much 

broader leaves (0.8-1.9 cm wide) than the other collections 

(0.35-0.8 cm wide) and also differs from them in a few other 

details (some leaf bases rounded vs. all cuneate; lamina 

lanceolate to lanceolate-elliptic vs. lamina linear-lanceolate; 

capsules 12-14 mm vs. 14-17 mm long). In inflorescence 

characters, however, it fully agrees with the other specimens, 

and it has therefore been included in A. longicapsa. 

Living material of Gillett & Beckett 23289 was obtained by 

removing and planting seeds from the EA sheet. Although all 

four seeds that were planted germinated shortly after sowing in 

1983, none of the plants survived dormancy, which is a 

common problem in cultivating species of section Somaliensia. 

The dorsal locule in one of the three ovaries dissected in 

preserved flowers of Gillett & Beckett 23289 contained an 

ovule. Therefore it is likely that capsules will be found that also 

contain a seed in this locule, although none has been 

encountered yet. 

Another collection from the Somali Republic, Thulin 5667 

(US) (121 km SW of Dusa Mareb along road to Belet Huen), is 

similar to Thulin & Warfa 5341, but its leaves are still broader 

(to 2.5 cm wide) and its minimum dimension for cincinnus 

peduncle length (6 mm) falls outside the range for A. 

longicapsa, as circumscribed here. In both of these characters 

the specimen approaches A. grandibracteolatum. 

Plants of Thulin 5667 have been grown from seeds removed 

from the specimen, and they have been found to agree with 

previously cultivated plants of A. longicapsa (Gillett & Beckett 

23289) in a number of significant characters of floral 

morphology (paired petal limbs hooded at apex; medial 

staminode filament broadened apically; lateral stamen filaments longer in staminate than in perfect flowers; pollen in 

lateral and medial anthers discolorous; ovary covered by 

relatively long, eglandular hairs) and in chromosome number 

(2n = 64). Because none of these characters is known in A. 

grandibracteolatum, the relationship of Thulin 5667 to that 

species has not been clarified, whereas its closeness to A. 

longicapsa has been confirmed. 

I believe that Thulin 5667 will have to be included in A. 

longicapsa in the long run and that it likely will have to be 

separated at the subspecific level. I prefer to await further 

collections before taking that step. For this account, I have 

mapped it as A. longicapsa with a question mark (Map 7), but 
NUMBER 76 87 

I have omitted it from the exsiccatae above. If the specimen 

were formally included in A. longicapsa, some of the 

dimensions that would have to be changed in the species 

description, in addition to leaf width and cincinnus peduncle 

length cited above, are flowers to 20 mm wide; sepals to 5 mm 

long; paired petals to 12 mm long and 9.5 mm wide; medial 

petal to 8 mm long and 5 mm wide; lateral staminode filaments 

to 7 mm long; style to 13 mm long. Whether these dimensions 

will prove to fall within the normal range for A. longicapsa or 

perhaps will be useful is separating subspecies remains to be 

determined. It should be noted that the altitude range for A. 

longicapsa would have to be extended to 450 m if Thulin 5667 

were included in the species. 

5. Aneilema pusillurn Chiovenda 

Aneilema pusillurn Chiovenda, Resultati Scientifici Missione Stefanini-Paoli 

Somalia Italiana, 1:167, 1916.-Cufodontis, Bull. Jard. Bot. Natl. Belgique. 

41, suppl., 1518, 1971. [Type: Somali Republic (Somalia Italiana on label), 

Praterie del Dafet fra Uanle Uein e Ilduc Uein, 19 Nov 1913, Puoli 1277 

F).1 

Perennial herbs (habit types IVB2, IVB3, IVC2, Figures 1,3; 

Plate 2i). Roots tuberous, fusiform, or uniformly thickened 

(subsp. gypsophilurn), to 5 cm long and 0.7 cm thick, not 

stipitate. Rhizome absent, very short or somewhat elongate, to 

2 cm long, old basal sheaths persistent or not. Flowering shoots 

annual, erect, unbranched or sparsely branched, 5-15(-18) cm 

tall. Internodes 1.3-4.5 cm long, glabrous or sparsely 

puberulous. Leaves spirally arranged or distichous, all cauline 

or mostly basal (subsp. thulinii), sheaths often split to the base, 

to -0.8(-2) cm long, glabrous or sparsely puberulous, glabrous 

or sparsely ciliolate with hook-hairs at the apex, laminae 

moderately to very succulent when fresh, flat or conduplicate 

when drying, linear to semicircular in transverse section when 

fresh, usually ascending, sessile, linear or linear-oblong to 

linear-lanceolate, lanceolate-elliptic or ovate-elliptic, 2-9 cm 

long, 0.1-2 cm wide, apex acute to acuminate, base broadly 

cuneate to rounded, margin scabrid or not, usually with 

prickle-hairs towards the apex, glabrous or with hook-hairs 

below, sometimes ciliate basally, adaxial surface glabrous or 

nearly so, abaxial glabrous to puberulous with hook-hairs, 

uniseriate hairs lacking. 

Inflorescences thyrses, terminal on main shoot and sometimes also on lateral shoots, moderately lax to moderately 

dense, narrowly ovoid to pyramidal, 1.5-6.5 cm long, 

(0.7-)1-3.5 cm wide, with (2-)6-lo(-13) cincinni, alternate 

or occasionally subopposite, ascending to erect, the lowermost 

sometimes becoming patent or declinate. Peduncles 2-7(-8.5) 

cm long, glabrescent to puberulous. Inflorescence bract usually 

medial, generally foliaceous, rarely bract-like. Distance between inflorescence bract and lowest cincinnus 0.5-4 cm. 

Inflorescence axis often zigzag, sparsely to densely puberulous 

with uniform hook-hairs. Cincinni to 3 cm long and 8-flowered 

(or more in cultivation). Cincinnus bracts patent to reflexed or 

ascending, scarious (rarely the lowermost foliaceous), lanceolate to ovate, 1.2-7.5(-25) mm long, sometimes glandular near 

the apex, puberulous. Cincinnus peduncles relatively uniform 

within the inflorescences or the upper ones somewhat reduced 

(occasionally lowermost greatly exceeding the others), usually 

exceeding the cincinnus bracts, (1 -2-)1.5-53-83 mm long, 

puberulous or glabrous. Cincinnus axes puberulous. Bracteoles 

attached (0.6-)1-3.6(-6) mm apart, scarious, eccentrically 

cup-shaped, perfoliate, 1.1-3 mm long, 0.3-1.3 mm high (very 

raiely split to base), glandular near the apex, puberulous. 

Flowers perfect and staminate, 6-9 mm wide (Plate 2j,k). 

Pedicels (1.5-)3-5 mm long in flower, to 6 mm long in fruit, 

aligned with the cincinnus axis in flower, erect or recurved 

120'-270' in fruit, persistent, sparsely to densely puberulous. 

Sepals reflexed or not, green, sometimes tinged with purple or 

maroon, puberulous; medial sepal lanceolate to lanceolateelliptic or ovate, 2-4 mm long, 1.3-2 mm wide, glandular or 

eglandular subapically; lateral sepals lanceolate-elliptic to 

ovate-elliptic, 2-4 mm long, 1.5-1.9 mm wide, not distinctly 

glandular. Paired petals 3-5.4 mm long, 2.5-4.5 mm wide, 

limb reflexed or not, ovate or ovate-elliptic to suborbicular (or 

obovate), -3-4.4 mm long, pinkish red (RHS color: 51B-C, 

Gillett et al. 22.593, pale lilac or blue (Thulin & Warfa 4659), 

apex rounded to obtuse, claw, (0.3-)0.7-1 mm long, yellowgreen or whitish, glabrous. Medial petal narrowly lanceolate to 

Iancmlate-ovate, ovate-elliptic or ovate, 2.5-4.5 mm long, 

1.5-2 mm wide, greenish, tinged with pinkish red and pinkish 

red at apex, or flesh pink or white (sometimes tinged with 

pink), apex acute, sometimes recurved. All filament bases free. 

Medial staminode yellow, very variable in development, 

filament 0.45-1.4 mm long, antherode bilobed, lobes sessile to 

shortly stipitate, obovate-cuneate, transversely elliptic, transversely lanceolate or reniform, -0.1-0.8 mm in diameter, 

connective slightly or not at all elongate. Lateral staminodes 

yellow, filaments 2-3.2 mm long, sharply decurved near the 

apex, antherodes bilobed, lobes stipitate, transversely ellipsoid 

or transversely lanceolate to ovoid, reniform or obovoid, 

0.4-1.4 mm in diameter, connective not elongate. Lateral 

stamens with filaments strongly divergent at least above the 

middle, 2.8-6.5 mm long, S-shaped, glabrous, anthers elliptic 

to broadly elliptic, 0.8-1.5 mm long, connective pale gray or 

gray-green, pollen yellow or dirty yellow. Medial stamen with 

filament 2-5.2 mm long, anther shield-shaped to saddleshaped, broadly elliptic to ovate, 0.5-1.4 mm long, connective 

yellow or white, pollen yellow or orange-yellow, discolorous 

from lateral anther pollen. Ovary sessile, elliptic or oblongelliptic, 0.85-1.3 mm long, 0.65-1 mm wide, densely and 

uniformly puberulous, apex rounded to truncate, dorsal locule 

l-ovulate, ventral locules each 2-ovulate; style 3.5-6.5 mm 

long, arcuate-decurved for most of its length, recurved near the 

apex, yellow in basal two-thirds, shading to lavender, lilac or 

maroon above, glabrous, stigma small or slightly enlarged. 

Capsules sessile, elliptic to oblong-elliptic, dehiscent, bivalved, trilocular, 2.4-5 mm long, 1.5-2.5 mm wide, 
88 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

stramineous (sometimes transversely streaked with brown) or 

brown, lustrous, puberulous with rather long hairs (to -0.5 mm) 

with the apical cell sometimes becoming somewhat hooked 

(but true hook-hairs absent), apex acute to obtuse, truncate or 

emarginate, dorsal valve deciduous, dorsal locule prominent, 

1-seeded or empty, ventral locules each 2-seeded (or less by 

abortion), cells of the capsule wall transversely elongate. Seeds 

(Plate 5h) transversely elliptic (dorsal locule seed) or subdeltate to ovate (ventral locule seeds), apical ventral rounded (or 

rarely acute) apically, basal ventral rounded basally, 1- 1.9 mm 

long, 0.75-1.3 mm wide, hilum dark brown, much shorter than 

the seed, prominently raised within a groove, groove f cut off 

apically and basally by ridges, with apical, basal and lateral 

edges crenate, testa tan or orange-tan, dorsal surface sulcate, 

(sulci radiating from the embryotega and becoming deeper 

towards the margins) or smooth with crenate margins, 

tan-farinose granules present or absent, if present, then 

confined to the embryotega and hilum. 

Key to the Subspecies of Aneilema pusillurn 

1. Leaves all or mostly cauline at flowering time, iamina linear to linear-lanceolate, 

0.1-0.8 cm wide. 

2. Plants not rhizomatous (or, if so, then other characters not as below); roots 

fusiform; plants growing on various substrates but usually not on gypsum. 

3. Lamina 0.1-0.3 cm wide; flowers coral red ....... A. p. subsp. pusillurn 

3. Lamina 0.3-0.8 cm wide; flowers pale lilac ................... .................... A. p. subsp. variabile, new subspecies 

2. Plants rhizomatous; roots f uniformly thickened, not fusiform; plants growing on 

gypsum .............. A. p. subsp. gyspsophilurn, new subspecies 

1. Leaves often mainly basal at flowering time, lamina lanceolate to ovate-lanceolate, 

the broadest 1-2 cm wide ......... A. p. subsp. thulinii, new subspecies 

5a. Aneilema pusillurn Chiovenda subsp. pusillurn 

HABlTAT.-open Acacia-Commiphora bushland on level 

red sand over limestone (Gillett et al. 22597); 135 m. 

FLOWERING.- Flowering specimens have been seen from 

June and November. In greenhouse cultivation, flowers of 

Gillett et al. 22597 opened between 1230-1400 hrs and 

remained open for about 4 hours. 

CHROMOSOME NUMBER.-n = 14,2n = 28. 

DIsTRIBunoN.-~oma~i Republic (Map 8). 

SPECIMENSEEN.-SOMALI REPUBLIC. 8 km NE of 

Budbud, 4’15’N,46O3 l’E, Gillett et al. 22597 (US); Praterie del 

Dafet fra Uanle Uein Wen] e Ilduc Uein Wen], Paoli 1277 

0. 

5b. Aneilerna pusillurn Chiovenda subsp. variabile Faden, 

new subspecies 

Ab subspecie typica foliis latioribus (0.3-0.8 cm latis, nec 

0.1-0.3 cm) et flonbus pallide lilacinis (nec roseis); ab subsp. 

thulinii foliis angustioribus semper omnibus caulinus tempore 

florendi differt. 

TYPE.-Kenya, Mandera District, 6 km S of El Wak, 

-2’46’N, 4Oo56‘E, 30 Apr 1978, Gilbert & Thulin 1251 (US, 

holotype; F, isotype). 

HABITAT.-Acacia-Commiphora bushland and woodland 

and Dichrostachys-Acacia bushland, on limestone, among 

limestone rocks. or on level gypsum plain; 160-420 m. 

April to June. 

FLowERING.-Flowering specimens have been seen from 

CHROMOSOME NUMBER.-Unknown. 

DISTRIBU~ON.-E Ethiopia, Somali Republic, and NE 

Kenya (Map 8). 

SPECIMENS SEEN.-ETHIOPIA. HARAR: Harrodigit, communicated Apr 1885, James & Thrupp s.n. (K). 

SOMALI REPUBLIC. 43 km E of Bahdo, 5O5O’NN, 47’08’E, 

Gillett et al. 22399 (US); 12 km S of El Bur along road towards 

El Dere, -4’37’N, 46’37’E, Thulin & Warfa 4659 (UPS). 

KENYA. NORTHERN FRONTIER: 6 km S of El Wak, -2’467% 

40°56’E, Gilbert & Thulin 4659 (F, US); Accampamento di 

Bela [Beila] sul Daua, Ruspoli & Riva 457 (844) (438) (FT). 

5c. Aneilema pusillurn Chiovenda subsp. gypsophilurn 

Faden, new subspecies 

Ab subspeciebus omnibus ceteris habitu rhizomifero radicibus plus minusve uniformiter incrassatis haud fusiformibus 

differt. 

TYPE.-Somali Republic [Somaliland], Bihen, 2 Jun 1945, 

Glover & Gilliland 1031 (K, holotype; EA, isotype). 

HABITAT.-Gypsum hills and pediment; limestone pediment 

(Beckett 1022); 570-710 m. 

FLOWERING.-FlOWering specimens have been seen from 

April and June. Plants collected with old inflorescences in 
NUMBER 76 89 

November indicate flowering occurs in that month too. 

CHROMOSOME NUMBER.-^^ = 28. 

DISTRIBUTION.-N Somali Republic (Map 8). 

SPECIMENSEEN.-SOMALI REPUBLIC. E of Gardo, 

9'33'N,49'07'E,Beckett 1022 @A); Bihen, Glover & Gilliland 

1031 (EA, K); 5 km SW of Halin along road to Gardo, 9'05'N, 

48'402, Thulin & Warfa 5423 (UPS, US); 8 km SW of Halin 

along road to Garoe, -9'03'N. 48'462, Thulin & Warfa 5395A 

(US). 

5d. Aneilema pusillurn Chiovenda subsp. thulinii Faden, 

new subspecies 

Ab subspecie typica foliis multo latioribus (ad 1-2 cm lam, 

nec 0.1-0.3 cm) saepe praecipue basalibus tempore florendi et 

floribus pallide lilacinis (nec roseis); ab subspecie variabili 

foliis latioribus (ad 1-2 cm lata, nec 0.3-0.8 cm) saepe 

praecipue basalibus tempore florendi differt. 

TYPE.-Somali Republic, 4 km N of Bulo Burti, Gaala 

Karor, -3'52'N, 45'342, 19 Nov 1985, Thulin & Warfa 5320 

(US, holotype; UPS, isotype). 

HABITAT.-LimeStone hills with Acacia -Commiphora bushland; 150-250 m. 

FLOWERING.-Flowering specimens have been seen from 

May and November. Flowers of Thulin & Warfa 5320 were 

observed in cultivation to open between 1130 and 1200 hrs. 

CHROMOSOME Numm.-Unknown. 

DISTRIBUTION.-€ Somali Republic (Map 8). 

SPECIMENS SEEN.-SOMALI REPUBLIC. 4 km N of Bulo 

Burti, Gaala Karor, -3'52'N, 45'342, Thulin & Warfa 4570 

(UPS); same locality, Thulin & Warfu 5320 (UPS, US); 62 km 

N of Bulo Burti along road to Belet Huen, Thulin 5673 (US). 

DISCUSSION 

Recent field work, especially by Thulin and Warfa, has 

increased the total number of collections of A. pusillum from 

four to 14 since I first studied the species (Faden, 1975). At that 

time I suggested that three of those collections might require 

taxonomic segregation from the fourth (the type). The data 

obtained from the newer gatherings fully support that proposal, 

but some problems remain, largely due to the still incomplete 

information about these taxa. 

Four subspecies are recognized here, but the number and 

taxonomic ranks of these infraspecific tam may have to be 

reconsidered when more data have been obtained, particularly 

from living plants. One of the most distinctive subspecies is 

subsp. gypsophilum, which is characterized by a rhizomatous 

habit and non-fusiform, thick roots. It is also allopatric from the 

other subspecies. However, even subsp. gypsophilum is 

inadequately known. Capsules and seeds have not been 

collected, and the flowers have been studied only from pressed 

specimens. Specimens lacking bases and ecological notes and 

occurring outside the recorded range of subsp. gypsophilum, 

e.g., James & Thrupp s.n., cannot be separated from subsp. 

variabile at present. They have been treated as the latter. Living 

plants of subsp. gypsophilum obtained for me by Mats Thulin 

yielded a chromosome number, but they died before flowering. 

Plants of the typical subspecies are characterized by their 

gracile habit and very narrow leaves that are semicircular in 

cross-section. Living material was obtained from a still 

succulent base on one of the dried plants of Gillett et al. 22597 

that was received as a gift for determination. The red flower 

color reported by these collectors and confirmed from the 

living plant is unique in the genus, but whether it occurs in 

other populations of subsp. pusillum remains to be determined. 

When I first received a specimen of subsp. thulinii (Thulin & 

Warfa 4570), I was convinced that it represented a new species 

related to A. pusillum. Its very broad leaves which were mainly 

basal gave the plant a very distinctive facies. At my request, 

Mats Thulin later collected for me living plants from the same 

population as his earlier gathering and from another population. 

Unlike the first collection, the new ones had capsules and seeds. 

Some plants grown from seeds exhibited the rosette habit of the 

original specimen. Others produced mainly caulescent leaves, 

as in the other collections of A. pusillum. The capsules and 

seeds also firmly placed the new collections in A. pusillum, but 

the very broad leaves and the tendency for them to be basal 

indicated that these plants needed to be segregated taxonomically. 

Subspecies thulinii is named in honor of Dr. Mats Thulin, 

whose collections of this and other Aneilema taxa from 

Somalia, Ethiopia, and Kenya have greatly aided my research 

on the genus. 

The most widespread subspecies-the only one not confined 

to the Somali Republic-and also the most poorly defined is 

subsp. variabile. Its leaves are always broader than those of 

subsp. pusillurn but narrower than those of subsp. thulinii, with 

both of which it shares the character of fusiform roots. Its 

leaves vary from flat, as in subsp. thulinii, to conduplicate, as in 

at least some plants of subsp. gypsophilum. It is not certain 

whether subspecies variabile exhibits any unique characters; it 

is presently defined by the absence of the distinct features that 

define the other subspecies. Thulin & Warfa 4659 has a distinct 

rhizome, but it is more elongate than that of subsp. gypsophilum and the roots are fusiform, so the collection has been 

placed in subsp. variabile. 

Despite all of the variation, Aneilema pusillum is a very 

distinctive species because of the plants' small size, very 

succulent leaves, few-branched inflorescences, small, lateopening flowers, deciduous dorsal capsule valves, seeds with a 

short hilum completely surrounded by a crenate-margined 

ridge, and unique chromosome number for the genus. 
90 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Section 5: Lumprodithyros (Hasskarl) C.B. Clarke 

Aneilema section Lampraiithyros (Hasskarl) C.B. Clarke in De Candolle, 

Monogr. man., 3:196, 1881. p.p.; in Thiselton-Dyer, FI. Trop. Africa, 8:62, 

1901, p.p.-Morton, J. Linn. Soc., Bot., 59:439, 1966, pap. 

Lamprodithyrar Hasskarl, Flora, 46388, 1863, pro gen.; in Peters, Naturwiss. 

Reise Mossambique, Bot., 529, 1864. [Lectotype: Lamprcdiihyros petersii 

Hasskarl (= Aneilema petersii (Hasskarl) C.B. Clarke).] 

Aneilema section Pseudo-axillares C.B. Clarke in Thiselton-Dyer, Fl. Trop. 

Africa, 8:63. 1901, in clad-Morton. J. Linn. Soc., Bot., 59:439,1966, pro 

syn. [Lectotype: Aneilema clarkei Rendle.] 

Ballyo Brenan, Kew Bull., 19:63, 1964, pro. gen. [Type: Ballyo zebrino 

(Chiovenda) Brenan (= Aneilema zebrinum Chiovenda).] 

Perennial or annual herbs with fibrous roots and spirally 

arranged or distichous leaves. 

Inflorescences terminal (and often axillary) thyrses, rarely all 

axillary, sometimes all reduced to single cincinni. Inflorescence 

axis (when present) puberulous with hook-hairs of uniform 

length (rarely glabrous). Cincinni elongate or contracted. 

Bracteoles membranous to herbaceous, usually cup-shaped, 

perfoliate, glandular, or eglandular subapically , occasionally 

also with small glands along the margin. 

Flowers perfect and staminate (rarely all perfect) (Plates 

h-p, 3, 4&), odorless. Pedicels erect in fruit or, more 

commonly, recurved 120°-270°(-360'). persistent, usually 

puberulous at least at the apex (occasionally glabrous). Sepals 

glandular subapically or occasionally apparently eglandular; 

hook-hairs, when present, of uniform length. Petals subequal, 

usually concolorous, white to lilac or lavender, glabrous; 

medial petal cup-, boat-, or slipper-shaped (Figures 23, 24), 

often retaining the lateral stamens when the flower opens. 

Filament bases of the stamens fused, frequently also fused to 

the lateral staminode bases for a shorter distance (Figures 

26-28). Staminodes with yellow, bilobed antherodes; medial 

staminode absent or much shorter than the lateral staminodes, 

antherode, when present, generally equal to or smaller than 

those of the lateral staminodes. Lateral staminodes with 

filaments nearly straight, then usually sharply decurved 

(occasionally recurved) near the apex. Lateral stamens with 

filaments f S-shaped, commonly sharply recurved above the 

middle, glabrous. Medial stamen with filament generally 

yellow apically, anther larger than the lateral anthers, saddleshaped, connective and pollen usually yellow (rarely white). 

Ovary sessile to stipitate, f densely and uniformly covered with 

patent, glandular hairs, sometimes mixed with few to many 

hook-hairs, dorsal locule prominent, usually 1-ovulate, ventral 

locules each 2-(rarely 1- or 3-4-)ovulate; style arcuatedecurved, usually recurved towards the apex, also commonly 

curved laterally out of the floral midplane, stigma capitate. 

Capsules usually stipitate or substipitate (Figure 35). usually 

dehiscent (rarely indehiscent), bivalved (rarely partially trivalved), usually trilocular, lustrous, generally puberulous, apex 

commonly emarginate (rarely obtuse, rounded, or truncate), 

valves persistent or dorsal valve deciduous, dorsal locule 

usually prominent, commonly 1-seeded, ventral locules each 

2-(or, by abortion, 0- 1-)seeded (rarely 3-4-seeded), cells of 

the capsule wall usually transversely elongate (occasionally 

irregular). Seeds of the dorsal and ventral locules commonly 

dimorphic, that of the ventral locule often differing in shape, 

size (particularly length and thickness), testa color and 

markings, presence or absence or abundance of farinose 

granules and color of the embryotega (Plate Q,f,g); dorsal 

locule seeds flat or convexo-planar, usually elliptic, oblongelliptic or ovate-elliptic in outline; ventral locule seeds usually 

ovate to triangular or trapezoidal, apical seed rounded to obtuse 

or acute apically, basal seed usually angular basally, hilum 

usually in a groove, testa generally shallowly scrobiculate or 

reticulate, farinose granules usually sparse (except around the 

hilum), occasionally absent, hypha-like filaments sometimes 

present among the farinose granules. 

BASIC CHROMOSOME NUMBER.-x = 13. 

DISTRIBUTION.-R~~ Sea Hills of NE Sudan and S Arabian 

Peninsula S to Zimbabwe, Swaziland, and South Africa (N 

Transvaal, N Natal) (Map 9). 

Key to the Species of Section Lamprodithyros 

1. Some or all inflorescences terminal thyrses with a distinct inflorescence axis; 

2. Pedicels glabrous, or with a few hairs at apex, mostly erect in fruit; bracteoles 

cup-shaped, prominently glandular along the margin, glabrous; sepals glabrous; 

capsules sometimes with more than 2 (up to 4) seeds per ventral locule. 

3. Plants annual; leaf margins usually only scabrid, occasionally also sparsely 

ciliolate; capsules (2.25-)3.3-4.5(-4.9) mm wide; dorsal locule seed 

2.7-3.45 mm long, 1.95-2.1 mm wide, 1.25-1.5 mm thick; ventral locule 

seeds (1.4-)1.7-2.2(-2.5) mm wide; (1.1-)1.2-1.45(-1.55) mm thick . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . , . . . 5. A. forskalii 

3. Plants perennial; leaf margins scabrid and ciliate; capsules (2.3-) 2.8-3.6 

(-3.7) mm wide; dorsal locule seed 2.15-2.85 mm long, 1.5-1.7 mm wide, 

1-1.2 mm thick; ventral locule seeds 1.55-1.9(-2) mm wide, 1-1.25 mm 

thick . , . , . . . . . . , , . , . . . . . . . . .4. A. sebitense, new species 

cincinni elongate. 
NUMBER 76 91 

2. Pedicels puberulous above the middle or occasionally only at apex, mostly 

recurved 180° or more in fruit; bracteoles various; sepals usually puberulous, 

rarely nearly glabrous; capsules with never more than 2 seeds per ventral locule. 

4. Medial sepal with marginal (as well as subterminal) glands; fruiting pedicels 

recurved 18O0-27Oo(-36O0); bracteoles with marginal glands; medial 

staminode lacking or vestigial and lacking an antherode. 

5. Inflorescences with 10-21 cincinni; bracteoles attached (1.5-)2-3(-3.3) 

mm apart, glabrous; sepals glabrous or subglabrous; capsules 3.5-4 mm 

5. Inflorescences with up to 8 cincinni; bracteoles attached 1-2(-2.5) mm 

apart, puberulous at least basally; sepals puberulous; capsules (2.4-)2.7- 

3(-3.4) mm long ...................... 8. A. tanaense 

4. Medial sepal with only a subterminal gland; fruiting pedicels mostly recurved 

-180"; bracteoles lacking marginal glands (except A. recurvaturn); medial 

staminode with an antherode. 

6. Pedicels puberulous only at apex; upper sepal glabrous; bracteoles glabrous, 

with small, inconspicuous marginal glands; capsules mostly 3.5-5 mm 

long x 2-2.5 mm wide .......... 3. A. recurvaturn, new species 

6. Pedicels usually puberulous in upper half, rarely only at apex; all sepals 

puberulous; bracteoles puberulous (rarely subglabrous), lacking marginal 

glands (margin sometimes thickened in A. indehiscens but not distinctly 

glandular); capsules mostly 4.5-7.5 mm long x 2.3-5 mm wide. 

7. Lamina lacking long, uniseriate hairs on the adaxial surface; bracteoles 

symmetrically to asymmetrically cup-shaped, lacking long, uniseriate 

hairs; Arabian Peninsula ........... 6. A. woodii, new species 

7. Lamina with long, uniseriate hairs on both surfaces; bracteoles 

asymmetrically cup-shaped or not cup-shaped, commonly with 1- 

several long, uniseriate hairs towards the base; Africa. 

8. Bracteoles not cup-shaped, although sometimes shortly perfoliate 

basally; lateral stamen filaments crossing; capsules (2.4-)3.3-5 

(-5.5) mm wide, pale gray or brown, peppered with numerous, small, 

dark brown and/or black spots, dorsal valve deciduous; dorsal and 

ventral locule seeds strongly dimorphic, differing in shape, size, testa 

surface and color, and embryotega color; dorsal locule seed 

hemispherical, humpbacked towards the capsule apex, 2-2.6(-3.1) 

mm wide, 2.3-3.4 mm long, 1.6-2.1 mm thick, testa smooth . , . . 

8. Bracteoles cup-shaped, usually some or all perfoliate; lateral stamen 

filaments not crossing; capsules (1.9-)2.3-3(-3.4) mm wide, 

chestnut brown or mottled dark and light brown or gray-brown, dorsal 

valve usually persistent; dorsal and ventral locule seeds scarcely 

dimorphic, differing only in shape and length, dorsal locule seed k 

flat, 1.35-1.65(-1.9) mm wide, 2-2.9 mm long, 0.65-1 mm thick, 

testa shallowly scrobiculate ............ 2. A. indehiscens 

1. All inflorescences composed of solitary or clustered, usually abbreviated cincinni, 

9. Some or all inflorescences terminal on the main and major lateral shoots; flowers 

(9-)lo-15(-20) mm wide; pedicels (4-p.5-11(-13) mm long, recurved in 

fruit 180°-2700(-3600);capsules dchiscent, apex truncate to emarginate, dorsal 

locule very prominent; ventral locule seeds 0.8-1.2 mm thick, testa usually 

shallowly scrobiculate. 

10. Inflorescences consisting of 1-8 fasciculate cincinni, not enclosed in a pair of 

leafy bracts; fruiting pedicels recurved k uniformly for their entire length; 

medial staminode absent or vestigial and lacking an antherode. 

long ........................... 7. A. benadirense 

.............................. 1. A. petersii 

always lacking an inflorescence axis or all inflorescences axillary. 
92 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

11. Bracteoles attached 1-2(-2.5) mm apart, with small marginal glands: 

medial sepal generally with marginal glands; lateral stamen pollen yellow 

............................... ,&A. tanaense 

11. Bracteoles attached 0.2-1(-1.4) mm apart, lacking marginal glands: medial 

sepal lacking marginal glands: lateral stamen pollen white ........ ............................... 9. A. calceolus 

10. Inflorescences consisting of solitary cincinni partially enclosed in a pair of 

leafy bracts; fruiting pedicels recurved only at the apex: medial staminode 

generally with an antherode. 

12. Laminae petiolate, bases symmetric; internodes not swollen; capsules 

3.6-4.5(-5) mm long; dorsal locule seed 1.65-2.15 mm long, 1.35-1.55 

mm wide. 

13. Flowers (12.5-)15-20 mm wide; pedicels puberulous apically; sepals 

glandular subapically; medial anther yellow, with yellow pollen: 

capsules 2.3-3 mm wide: testa of dorsal locule seeds light brown . . , 

.............................. .lo. A. clarkei 

13. Flowers 10.5- 14(-15.5) mm wide; pedicels pilose-puberulous apically; 

sepals apparently eglandular; medial anther white with white pollen; 

capsules 1.95-2.1(-2.55) mm wide; testa of dorsal locule seeds tan, 

orange-buff or orange-brown, usually heavily spotted and striped with 

dark brown ................ 11. A. lamuense, new species 

12. Laminae sessile, bases asymmetric; internodes commonly swollen; capsules 

(2.8-)3.2-3.7(-4.3) mm long; dorsal locule seed 1.35-1.55 mm long, 

1.2-1.25 mm wide. .......... 12. A. succulentum, new species 

9. All inflorescences axillary; flowers 7-10 mm wide: pedicels (0.5-)1.5-5 mm 

long, erect or slightly further recurved in fruit: capsules indehiscent or partially 

dehiscent, apex obtuse to rounded or truncate, dorsal locule a low hump: ventral 

locule seeds 0.5-0.6 mm thick, testa shallowly reticulate . . 13. A. zebrinum 

The lectotypification of Lamprodithyros Hasskarl by L. 

petersii requires some explanation because Hasskarl’s circumscription (Hasskarl, 1863, 1864a) of the genus is equivalent to 

Aneilema as a whole in the sense used here, rather than to a 

section of it. Along with his description of the genus, Hasskarl 

(1 863) published ten combinations in Lamprodithyros, two of 

which, L. radicans and L. stenorrhachys, are based on 

manuscript names and are nomina nuda. Of the remaining 

eight, five (L. schomburgkianus (Kunth) Hasskarl, L. petersii 

Hasskarl, L. dregeanus (Kunth) Hasskarl, L. tacazzeanus 

(Hochstetter ex A. Richard) Hasskarl, and L. beniniensis (P. de 

Beauvois) Hasskarl) belong to Aneilema in the present sense. 

The description of Lamprodithyros and accompanying notes 

do not necessitate the selection of any particular species as the 

lectotype. However, Hasskarl(l863) included diagnoses for L. 

petersii and L. tacazzeanus-these were later amplified into 

lengthy descriptions (Hasskarl, 1864a)-but not for the others. 

It is reasonable to choose one of the two species that Hasskarl 

studied in detail as the type species of Lamprodithyros. 

Lamprodithyros petersii has been selected because (1) the type 

still exists in the Berlin Herbarium, and this species was not 

confused with any other by Hasskarl, and (2) Hasskarl’s 

diagnosis and description of L. tacazzeanus were based on a 

Mozambique collection, but true Aneilema tacazzeanum 

Hochstetter ex A. Richard (= A. forskalii Kunth), does not 

occur south of Ethiopia, and the specimen that Hasskarl studied 

(fragment seen at L) was A. nyasense C.B. Clarke. 

Clarke’s section Pseudo-axillares includes three species, A. 

pedunculosum C.B. Clarke, A. clarkei Rendle, and A. taylorii 

C.B. Clarke, which are here placed in three different sections. 

Section Pseudo-axillares was based on the presence of axillary 

inflorescence-shoots, the “cyme-branches” of Clarke (1901). 

Because these occur in all sections of the genus, section 

Pseudo -axillares, as construed by Clarke, is clearly unnatural. 

Clarke’s description does not favor typification of the section 

by any particular species. Aneilema clarkei conforms with the 

description and is here chosen as the lectotype. The resultant 

synonymy of section Pseudo -axillares with section Lampro - 

dithyros agrees with the treatment of Morton (1966). 

The genus Ballya was based on the single species A. 

rebrinum (Brenan, 1964). Supposed anatomical differences 

between Ballya and Aneilema were reported by Tomlinson 

(1966). Brenan’s chief character for recognizing Ballya was an 

artifact of cultivation, and Tomlinson did not have available 

other species of section Lamprodithyros that are anatomically 

similar to A. zebrinum. Further details are given in the 

discussion under A. zebrinum. 

Section Lamprodithyros is very natural as evidenced by the 
NUMBER 76 93 

I I I I I I I 

60 40 20 40 60 

khp 9.--Distribution of Aneilema section Lamprodithyros. 

great similarity in floral and capsular morphology among the 

species, the difficulty in separating many of the taxa from 

incomplete dried specimens, and the single basic chromosome 

number. The large, cup-, boat-, or slipper-shaped medial petal, 

which is usually the same (or nearly the same) color as the 

paired petals and, in many species, retains the lateral stamens 

for some time after the flower opens (see Faden, 1983b), is the 

most distinctive feature. Also very characteristic are the fused 

stamen filament bases and the trilocular capsule in which the 

dorsal and ventral locule seeds are commonly strongly 

dimorphic. When the seeds are very dimorphic, the dorsal valve 

of the capsule is usually deciduous. 

Within section Lamprodithyros there are several closely 

related groups of species: A. sebitense, A. forskalii, and A. 

woodii; A. benadirense, A. tanaense, and A. calceolus; and A. 

clarkei, A. lamuense, and A. succulentum. Aneilema petersii 

and A. recurvatum also seem to be closely related to each other, 

and A. indehiscens may have arisen as an allotetraploid hybrid 

between them. Aneilema zebrinum seems to have been an 

offshoot of the A. clarkei species group. The relationships 

among the various species groups to one another remains to be 

determined. 

1. Aneilema petersii (Hasskarl) C.B. Clarke 

Aneilema petersii (Hasskarl) C.B. Clarke in De Candolle, Monogr. Phan., 

3:225, 1881, p.p.; in Thiselton-Dyer, Fl. Trop. Africa, 8:70, 1901, 

p.p.-Durand and Schinz, Conspect. Fl. Africae, 5131, 1895, p.p.- 

Schumann in Engler, Pflanzenwelt Ost-Afrikas C, 136, 1895, p.p.-Brenan, 

Kew Bull., 7:195, 1952.-Faden in Agnew, Upland Kenya Wild Fl., 666, 

1974. 

Lamprodithyros pefersii Hasskarl, Flora, 46:389, 1863; in Peters, Naturwiss. 

Reise Mossambique, Bot., 529, 1864. [Type: Mozambique (without 

locality), Festland, an feuchten Orten (undated1)Peters s.n. (B, fragment L).] 

Aneilema fetraspermum K. Schumann in Engler, Pflanzenwelt Ost-Afrikas C, 

136, 1895. [Syntypes: Tanzania, Tanga, Feb 1893, Volkens 175 (B, 

destroyed; K, lectotype; BM, G, isolectotypes); Stuhlmann 6062 and 6458 

(both B, both destroyed).] 
94 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

Aneilema leptospermwn K. Schumann [“leptasperma”], in sched. [Volkenr 

1751. 

Aneilema sacleurii Hua, Bull. Mus. Hist. Nat. (Paris), 1:121, 1895.4arke in 

Thiselton-Dyer, Fl. Trop. Africa, 8:74, 1901. [Type: Zanzibar. bord de la 

mer, Apr 1890. Sacleur 1142 (P; photo K) (cited as “Sacleux 1192” by 

Brenan. Kew Bull., 7195,1952).] 

Aneilema chtysanthwn K. Schumann, Bot. Jahrb. Syst., 33:1903. [Type: 

Tanzania, by Mengwa, Stuhlmann 8611 (B, destroyed).] 

Aneilema tacazzeanum sensu Chiovenda. Missione Biol. Paese Borana. 4:305, 

1939 [pro Cufodontis 705, non Hochstetter ex A. Richard (ISSO)]. 

Perennial or annual herbs (habit types IA, IB, IC, IIA4, and 

IIIA, Figures 1,2). Roots fibrous. Shoots erect or ascending to 

decumbent, sparsely to densely branched, sometimes thickened 

at the base, 20-60(-100) cm tall. Internodes (0.7-)2-15.5 

(-19.5) cm long, puberulous at least below the nodes (rarely 

completely glabrous). Leaves spirally arranged (rarely distichous), sheaths 0.5-2.5 cm long, puberulous or pilosepuberulous, ciliate at the apex, laminae petiolate (the upper 

several on the flowering shoot sessile), often gradually reduced 

towards the terminal inflorescence, narrowly lanceolate or 

lanceolate-elliptic to lanceolate-ovate, ovate-elliptic or ovate, 

(2.5-)3.5-1 l(-13.5) cm long, (0.6-)1-3.5(-4.5) cm wide, 

apex acute to acuminate (rarely obtuse), base cuneate to 

rounded, in the upper leaves sometimes subcordate- or 

cordate-amplexicaul, margin often undulate, scabrid and 

ciliate, both surfaces usually lustrous (sometimes the adaxial 

dull), adaxial usually pilose-puberulous (occasionally puberulous), abaxial puberulous or pilose-puberulous, the uniseriate 

hairs, when present, typically shorter than those on the adaxial 

surface; veins sometimes pale on the adaxial surface. 

Inflorescences thyrses (Plate 2m,o), terminal on the main 

flowering shoots and frequently on axillary inflorescenceshoots from the inflorescence bract and upper leaves (also 

frequently terminal on longer, perforating or non-perforating, 

axillary shoots), moderately lax, ovoid, to 7(-9.5) cm long and 

6.5(-9.5) cm wide, with 1-16(-19) cincinni, alternate (or a 

few sub-opposite), patent to ascending. Peduncles (1 -7-)3- 

183-21.5) cm long, puberulous. Inflorescence bract supramedial to medial, usually herbaceous, foliaceous or bract-like 

(Plate 20). Inflorescence axis puberulous. Cincinni to 4(-7) cm 

long and 16(-33)-flowered. Cincinnus bracts membranous, 

appressed to the cincinnus peduncles or patent, occasionally 

reflexed, lanceolate or lanceolate-elliptic to ovate-elliptic or 

ovate, (1.7-)2-4.5(-5.5) mm long, not evidently glandular or 

with subapical glands apparent only in the youngest inflorescences, glabrous to puberulous. Cincinnus peduncles relatively 

uniform within the inflorescence or decreasing in the upper 

cincinni, exceeding the cincinnus bracts, (3.3-)4- 10.5(-14) 

mm long, puberulous. Cincinnus axes puberulous. Bracteoles 

attached 1-4(-5) mm apart, membranous, ovate to ovatelanceolate, sometimes slightly perfoliate at the base, (1.1-)1.3 

-2.6 mm long, to 0.5(-0.7) mm high, sometimes distinctly 

glandular near the apex, sometimes apparently eglandular, 

puberulous (rarely subglabrous), also with 1 -several long 

uniseriate hairs towards the base or on the margin. 

Flowers perfect and staminate, odorless, (6.5-)11-19 mm 

wide (Plate 2~). Pedicels (2.2-)2.5-5(-5.8) mm long in 

flower, to 7 mm long in fruit, horizontal to erect in flower, k 

uniformly recurved in fruit 90’-220” (usually -180”) (Figure 

35a), persistent, puberulous at least apically. Sepals glandular 

near the apex with unlobed glands, puberulous and frequently 

with a few uniseriate hairs; medial sepal lanceolate or 

lanceolate-elliptic to ovate or ovate-elliptic, 2-5.2 mm long, 

(1.7-)2-3 mm wide, subapical gland prominent; lateral sepals 

ovate-elliptic to ovate or ovate-orbicular, 2.2-5.1 mm long, 

(2-)2.2-3 mm wide, subapical gland not prominent. Paired 

petals 4.5-9.5 mm long, (3.5-)4.5-11 mm wide, limb broadly 

ovate to ovate-deltate (occasionally ovate, rarely ovatereniform), 3-6.5 mm long, white to lilac or pinkish purple 

(RHS colors: 76A-B, Faden et al. 741326; 76B-C, Andrews s.n. 

cultivated; 76C, Faden & Faden 741291, cultivated; 87D, 

Faden & Faden 74/1183), apex rounded to truncate (rarely 

slightly emarginate), margin crenulate to subentire; claw 

(1-)lS-3(-4) mm long, concolorous with limb or paler 

basally, glabrous. Medial petal cup-shaped, ovate to obovate, 

broadest at the margin (viewed from the apex), which is slightly 

revolute, not retaining the lateral stamens when the flower 

opens, 3.5-7.5 mm long, 3-8 mm wide, 1.5-3.7 mm deep, 

concolorous with the limbs of the paired petals (Figure 23a-c). 

Lateral stamen filaments fused basally to medial stamen 

filament and occasionally to lateral staminode filaments 

(Figure 26b), medial staminode free. Medial staminode 

filament (1.2-)1.5-3.5 mm long, white to lilac basally, 

shading to yellow above (occasionally entirely yellow), 

antherode bilobed, yellow, lobes sessile or subsessile, obovatecuneate, orbicular, transversely elliptic or broadly ovate, 

(0.25-)0.3-1.1 mm long, connective not or scarcely elongate, 

Lateral staminodes with filaments 2-4.5(-5.2) mm long, white 

to lilac below the middle, shading to yellow above, antherodes 

yellow, bilobed, similar in size and form to that of the medial 

staminode. Lateral stamens with filaments parallel (occasionally slightly divergent) near the base, then convergent and 

crossing about the middle to near the apex (rarely not crossing) 

(Figure 26b; Plate 2n,p), (3.8-)4.5-10 mm long, S-shaped, 

decurved after -basal l/3 of their length, anthers ovate to 

ovate-elliptic (rarely lanceolate-elliptic to elliptic), 0.55-1.3 

mm long, 0.45-1.1 mm wide, pollen yellow (rarely whitish). 

Medial stamen with filament 3-7 mm long, anther ovateelliptic to ovate or ovate-lanceolate, saddle-shaped, 0.9-2.7 

mm long, 0.6-1.3 mm wide, pollen yellow (occasionally 

orange-yellow), usually concolorous with the pollen of the 

lateral anthers. Ovary stipitate, obovate to obovatesubquadrate, 1.25-2.1 mm long, 1-1.5 mm wide, densely and 

uniformly covered with patent, glandular hairs, apex truncate, 

dorsal locule very prominent, 1 -ovulate, ventral locules each 

2-ovulate; style 2.5-9.8 mm long, decurved after the ovary and 

straight or gently arcuate-decurved for most of its length, also 

generally strongly curved out of the floral midplane (Plate 

2n,p), stigma capitate, held at the same level as the lateral 
NUMBER 76 95 

anthers and in front of them, lateral to them or in contact with 

one of them. 

Capsules (Figure 35a,b) stipitate, obovate to subquadrate 

(occasionally obovate-oblong), dehiscent, bivalved, trilocular, 

(3.4-)4.5-73-9) mm long, (2.4-)3.3-5(-5.5) mm wide, 

carinate middorsally, pale gray or brown, peppered with 

numerous, small, dark brown and/or black spots, lustrous, 

puberulous, apex emarginate (occasionally truncate), dorsal 

valve deciduous, dorsal locule very prominent, humpbacked, 

1-seeded or, by abortion, empty, ventral locules each 2-(or, by 

abortion, 1-)seeded, cells of the capsule wall transversely 

elongate or irregular. Dorsal locule seed (Plate 6a) f 

hemispherical, humpbacked towards the capsule apex, 2.3-3.4 

mm long, 2-2.6 (-3.1) mm wide, 1.6-2.1 mm thick, hilum 

dark brown, in a shallow groove, f equal to the seed length, 

testa orange-buff (occasionally orange-brown or flesh pink), 

smooth, lacking farinose granules. Ventral locule seeds (Plate 

6a) ovate to oblong or rectangular, apical seeds acute to obtuse 

or rounded apically, basal seeds rounded to truncate basally, 

(1.3-)1.5-2.6(-2.9) mm long, 1.6-2.3(-2.6) mm wide, 1-1.8 

mm thick, hilum dark brown, usually not in a groove 

(occasionally in a very shallow groove), slightly extended onto 

apical or basal surface, testa orange-buff (occasionally buff), 

shallowly scrobiculate, not farinose or sparsely blackish 

farinose in the depressions and sometimes around the hilum 

and embryotega. 

Key to the Subspecies of Aneilema petersii 

Perennials; short, perforating, inflorescence-bearing shoots absent from the lower 

nodes of the main shoots; flowers 15-19 mm wide; sepals (3.3-)3.5-5.2 mm 

long; paired petals lilac to pinkish purple, 7-9.5 mm long, with claws 2-3(-4) 

mm long; styles (6.5-)7.5-9.8 mm long , . . . . . . . . . A. p. subsp. petersii 

Annuals; short, perforating, inflorescence-bearing shoots usually produced from the 

middle and lower nodes of the main shoots; flowers (6.5-)11-15 mm wide; sepals 

2-3.4(-3.8) mm long; paired petals white to very pale lilac, 4.5-7(-7.8) mm 

long, with claws (1-)1.5-2 mm long; styles 2.5-6.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. p. subsp. pallidiflorum, new subspecies 

la. Aneilema pefersii (Hasskarl) C.B. Clarke subsp. pefersii 

HABmAT.4rassland with scattered shrubs, bushland, 

thicket margins, wooded grassland, woodland, lowland forest 

margins, roadsides, cultivation, occasionally in damp situations, rarely in rocky places; sandy soil; full sun to dense 

shade; 0-500(-1lOO) m. 

FLOwERING.-~OWering specimens have been seen from all 

months except December. In the field the flowers of one 

population opened from 0600-0630 hrs and faded between 

1100-1130 hrs (Figure 25). Two other populations were noted 

as fading about 1030 hrs. 

CHROMOSOME NUMBER.-n = 13,2n = 26. 

 DISTRIBUTION.-^ Ethiopia to N Kenya and coastal and 

subcoastal Kenya to Mozambique (Map 10). 

SPECIMENSEEN.-ETHIOPIA. SIDAMO: -10 km verso 

Mega [from] Moyale, Cufodontis 705 F). 

KENYA. District unknown: 5 mi W [of] Kisuvi jet, 

Moomaw 951 (EA); Kenya Coast, Rawlins 239 (EA). KILIFI: 

Sokoke Forest, -3 km on track S of Gede to Jilore Forest 

Station, -3"18'S,39'57'E, Faden & Faden 7411237 (MO); 2.5 

km Kakoneni-Jilore Forest Station road, 3'1 l'S, 39'52.52, 

Faden et al. 72/38 (EA, K); Arabuko, Graham 1848 [= K18481 

(EA-2 sheets, K, MO); Sokoke Forest, Jefery K80 (EA, G 

(data different on this sheet), K); Sokoke Forest, Saunders in 

EA11211 (EA); Kakoneni, -20 km W of Malindi, Spjut 3812 

(EA); Mida, Tweedie 3183 (K). KILIFILAMU: N of Mombassa 

[sic] to Lamu and Witu, 1902, Whyte s.n. (BM). KWALE: Near 

Kivukoni [Kivumoni] entrance to Shimba Hills National 

Reserve, Bully & Smith B14357 (EA, K); Shimba Hills 

National Reserve, Campsite No. 1, Faden & Evans 701819, 

cultivated Missouri Botanical Garden (EA, MO); Buda Forest, 

4"27'S,39"23'E, Faden & Faden 741291 (EA, MO); -7 km past 

crossing of Ramisi River on Ramisi-Mrima Hill road, -4'323, 

39'19'E, Faden & Faden 741308 (MO); Kwale, Graham 1903 

[= K1903 & Q5221 (EA-3 sheets, K-2 sheets, MO); between 

Umba and Mwena Rivers, Lucas et al. 256 (EA, K); Shimba 

Hills, Tanga road near Marere Water Works, Magogo & Glover 

906 (EA, K); Kwale, McKeag [or McCraig] in CM9187 

(EA-2 sheets); S of Mombassa [sic] to Shimoni, 1902, Whyte 

s.n. (BM); Shimoni, S of Mombasa, 1902, Whyte s.n. (K). 

LAMU: Mambosasa Game Forest Post along Ziwa la Mkuni, 

2'233, 40'32'E. Faden & Faden 7411142 (BR, EA, FT, K, 

MO, PRE, US); Witu, Rawlins in EA11280 (EA, K). 

NORTHERN FRONTIER: Mt. Burrole just SW of Sololo, Bully & 

Smith B14873 (EA, K); Moyale, 3'32'N, 39'03'E, Gillett 14091 

(B, BR, EA, K). TANA RIVER: 105 km N of Malindi on Garsen 

road, cultivated Missouri Botanical Garden, Andrews sen. (EA, 

K. MO); Garsen-Malindi road, 1.5 km towards Malindi from 

turnoff to Oda, Faden & Faden 7411183 (EA, G, K, LISC, MO, 

PRE, US); Garsen-Malindi road, -8 km from Garsen, -2'22'S, 

40'07'E, Faden et al. 721228 (C, EA, K, MO, US). TEITA: 10 
96 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

40 I 

8 J I I 

hhP 10.-Distribution of Aneilema peiersii (Hasskarl) C.B. Clarke subsp. peiersii (solid circles), A. peiersii 

subsp. pallidij7orwn Faden, new subspecies (open circles), and A. sebiiense Faden, new species (solid squares). 
NUMBER 76 97 

km Taru-Mackinnon Road on Nairobi-Mombasa road, 

3'443, 39'043, Faden & Faden 72188 (EA, K, US, WAG); 

Mt. Kasigau, Faden et al. 691453 (EA, K); Taru Desert near 

Maungu, Tweedie 3365 (K). 

TANZANIA. BAGAMOYO: Bana Forest Reserve, Mgara 781 

(EA, K); Kerege, Dar es Salaam, Sector 30 of Tsetse Research 

Unit, D.C. Robertson in TRU518 (EA, K); Chambezi Agricultural Station, 6"35'S, 38'553, SA. Robertson 529 (EA, K); 2 

mi E of Ruvu River, Tweedie 1624 (K). KILWA: Selous [Game 

Reserve], Madaba, 8'353, 37'45'E, Vollesen 3350 (EA). 

LINDI: 40 km W [of] Lindi, Lutambasee, Schlieben 5942 (B, 

BM, BR, G-2 sheets, M, S). MOROGORO: 3 mi N [of] 

Tununguo, 30 mi SE [of] Morogoro, Boaler 626 (BR, EA, K, 

NY, PRE); Boma, Morogoro Town, Harris BJH5266 (MO); 

No. 1 Fly Round, 10 mi NE of Kingolwira Station, Morogoro, 

Welch 243 (EA, K-2 sheets). PANGANI: Bushiri, Faulkner in 

K712 (K-2 sheets); Mkwaja Ranch, -5'40'-5'47'S, Kloetrli 

et al. 144 (EA); Mwera-Mganbo Flgambol road, Richards 

21988 (K). RUFIJI: Mohoro, Braun 3624 (EA); Mafia Island, 

Kilindoni, Greenway 5236 (EA, K, PRE); Mafia Island, 

Wallace 810 (K). RuFUI/KILWA: Without locality, Busse 3054 

(EA). TANGA: Tanga, Archbold 1491 (EA, MO); Totohovu 

[Mtotohovu] bf. Moa, Braun 1377 (EA-2 sheets); without 

locality, Busse 2281 (EA); 5 mi SE of Ngomeni, Drummond & 

Hemsley 3610 (B, BR, EA, K-2 sheets, S); 9 km TangaGombero road from Tanga-Mombasa road junction, -5"OO'S, 

38'58'E, Faden et al. 741326 (B, BR, C, COI, DSM, EA, FT, K, 

LISC, MO, P, PRE, S, US, WAG); Kange Estate, Faulkner 838 

(BR, K-2 sheets, S); Pongwe-Maweni, Faulkner 3670 (K, 

MO, SRGH) & Faulkner s.n. (EA, K); Amboni b. Tanga, 

Geilinger 133 (K, Z) & 155 (K, Z) & 642 (Z); Tanga, Volkens 

175 (BM, G, K). UZARAMO: 5 mi up Bagamoyo road [from Dar 

es Salaam], Batty 162 (K); Morogoro-Dar es Salaam road, 

Bamba, 6'44'S,38'513, Faden et al. 701686A (EA, K); near 

Dar es Salaam airport, Harris 1484 (EA) & BJH3069 (EA, 

MO); 3 mi NE [of] Dar es Salaam on Bagamoyo road, Harris 

2045 (EA); Dar es Salaam, Ubungo-Kigogo road, Harris 

BJH2848 (EA); Dar es Salaam, Harris BJH3413 (EA); 

Kunduchi, 16 km NNW of Dar es Salaam, 6'41'-43'S, 

39'13'-14'E, Harris & Tadros BJH5721 (EA); -9.5 km E of R. 

Ruhu [Ruvu], Milne-Redhead & Taylor 7437 (BR, EA, K-2 

sheets, LISC). ZANZIBAR: Without locality, 1847-1852, Boivin 

s.n. (P); Bord de la mer, Sacleux 1142 (P; photo K); Bet-el-ras, 

Vaughan 2106 (BM). 

MOZAMBIQUE. Without locality, Festland, an feuchten 

Orten, undated, Peters sen. (B). NIASSA: Mouth of Msalu River, 

Allen 94 (K); Mussovil [Mossuril] & Cabeceira, 1884-1885, 

Carvalho s.n. (COI). ZAMBEZIA: -9 km de Nicuadala [on] 

estrada para Namacurra, Torre & Correia 14296 (LISC). 

lb. Aneilema petersii (Hasskarl) C.B. Clarke subsp. 

pallidiflorum Faden, new subspecies 

Ab subsp. petersii habitu annuo, floribus parvioribus 

((6.5-)ll-15 mm latis), sepalis parvioribus (2-3.4(-3.8) mm 

longis); petalis lateralibus albis vel perpallide lilacinis et 

parvioribus (4.5-7(-7.8) mm longis) unguibus (1-)1.5-2 mm 

longis, stylo parviore (2.5-6.5 mm longo) differt. 

TYpE.-Kenya, Eita [Taita on label] District, Irima Rock 

above Irima Water Hole, 3'18'S,38'33'E, 24 Apr 1974, Faden 

& Faden 741237 (US, holotype; BR, EA, K, MO, PRE, WAG, 

isotypes) . HABITAT.-Deciduous bushland and bushland thickets commonly dominated by species of Commiphora and Acacia, 

frequently on rocky hills and outcrops, also recorded from 

grassland, woodland, riverine thickets, roadsides, and in 

swampy situations; sandy soil: partial shade to full sun; 

(125-)500- 1000 m. 

FLOWERING.-Flowering specimens have been seen from 

February to June and August (Kenya and Tanzania), September 

(Sudan and Uganda), and November (Somali Republic). 

Flowers of populations in the Tsavo area of Kenya open from 

0545 to 0630 hrs and fade 1030-1 100 hrs. 

CHROMOSOME NUm3m-n = 13,2n = 26. 

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 Smithsonian Institution (US). 

SOMALI REPUBLIC. BALAD: Hilweyne area along BaladMogadishu 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, Acheptopokwa 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). KIT~ANA 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). 

mu: Meru National Park, S of Point 40, new road crossing 

Kiolu River, Ament & Magogo 263 (EA). NORTHERN FRONTIER: 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, Huch 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 I1 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, subplanoconvex, lustrous, mother-of-pearl colored, strongly humpbacked, 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 inflorescence-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 populations 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, inflorescencebearing 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.pallidiflorum, 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. Observations 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, ascending (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 inflorescence or decreasing in the upper cincinni, exceeding the 

cincinnus bracts, 3-1 1(-12.5) mm long, puberulous. Cincinnus 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, mq): 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, 

frm Faden & Faden 741208; mq, 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 ovateelliptic, 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, obovateelliptic 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 

HABITAT.-Bushland and thickets of varied species compo- open 0600-0630(--0645) hrs and fade 1100-1230 hrs. 

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 

MAP 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 SEEN.-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); GarsenMalindi 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 NairobiMombasa 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 Joaao 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 irregulariter 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 (US, 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). 

SPECIMENSEEN.-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. Commonly 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 unknown 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 collections of subsp. indehiscens show some morphological tendencies 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 connectives, 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 triloculares, (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 glabrescent). 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 inflorescenceshoots from the inflorescence bract and upper leaves (occasionally 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 subverticillate (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 recwafwn Faden, new species: a, habit; b, inflorescence; 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, foliaceous 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 lanceolate (rarely the lowermost linear), 1.5-3(-9.5) mm long, 

glandular subapically, usually glabrous or subglabrous. Cincinnus 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 cupshaped, perfoliate, 1-2 mm long, 0.5-1.25 mm high, prominently 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), subapical 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, obovatecuneate 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 oblongelliptic, saddle-shaped, 1.5-2.2 mm long, 0.95- 1.4 mm wide, 

pollen concolorous with lateral anther pollen. Ovary substipitate, 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 42, 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, oblongelliptic 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 

SPECIMENSEEN.-ETHIOPIA. GEMU-GOFA: Rive del 

NW Tanzania (Map 12). 
NUMBER 76 109 

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MAP 12.- Distribution of Aneilema recwvatwn Faden, new species. 

Ghizo, Corradi 2159, p.p. F); base of escarpment W of Arba 

Minch, 6OOO'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); TambachTot 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 UrnRONDO: 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, lessobovate, 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 staminode, 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 lanceolato-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 numerosis 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 lode 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). pars = 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 (US, 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, pilosepuberulous, ciliate at the apex, laminae petiolate (except the 

uppermost leaves), sometimes reduced towards the inflorescence, 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 declinate), 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. Cincinnus 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 subapically 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 ovatelanceolate, 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 oblongelliptic, saddle-shaped, 1.7-2.5 mm long, 0.9-1.5 mm wide, 

pollen golden yellow. Ovary substipitate, obovate to obovateoblong, -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, sometimes (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 SEEN.-ETHIOPIA. GEMU-GOFA: Arba Minch, 

escarpment below the town, -6’OSN. 37’3% Gilbert & 

Thulin 266 (MO). SIDAMO: [“Somali-land” on label], Donuldson 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 cupshaped, 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. Aneilem 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. Internodes (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 (occasionally the lower becoming patent or declinate). Peduncles 

4-15(-22.5) cm long, puberulous. Inflorescence bract supramedial (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 lanceolateelliptic), (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 generally 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, concolorous 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 lanceolateelliptic 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 obovate-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 convexoplanar, 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 64 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; FilfilMarrao, 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, Schweinfurth & 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 prop 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 description 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 Hochstetter 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 ehrenbergii, but Ehrenberg 413 bears both Hasskarl’s and Schweinfurth’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 specimens 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 specimens 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 antherodeslobes 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 lanceolateelliptic, 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 

33-53 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), puberulous. 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 (symmetrically 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 staminode 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 * concolorous 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 Schweinfruit. 

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 collections. 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 peduncles, 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 

52~ 

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 ovatelanceolate, 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 foliaceous), 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 puberulous. 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 lanceolateelliptic, (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 ovateoblong, 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 smalleranthered 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, “greenishwhite 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 capsule 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 dimorphic. 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 subglabrous 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 ma 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 inflorescences 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. Inflorescence bract supramedial to subapical, herbaceous and foliaceous. 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 subglabrous. Cincinnus peduncles, except the lowermost, generally 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 cupshaped, 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 ovateelliptic 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 represented 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, saddleshaped, 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 whitefarinose in many of the depressions and around the embryotega, 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). 

SPECIMENSEEN.-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 FA, K, MO); GarsenMalindi 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 inflorescence 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 communication). 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 inflorescences (species 1-7) and those that never have them (species 

9-13). More precisely, it is intermediate between A. benadirense and A. calceolus in lamina length, inflorescence form, 

cincinnus number per inflorescence, distance between bracteoles, 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, consisting of 1-4(-6) cincinni that appear fasciculate in the ails 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 cupshaped, perfoliate, 1.1-1.9 mm long, 0.4-1 mm high, 

inconspicuously glandular near the apex, glabrous or subglabrous. 

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 obovateelliptic, 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) hemispherical, 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 hyphalike 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. 

DISTRIBUTION.~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. 34 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 uniseriate 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 membranous. 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 ovatesubreniform, -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 oblongelliptic, 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, prominently 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 depressions, 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 observations of three populations indicate that the flowers open 

0845-0930 hrs and fade 1200-1300 hrs. 

CHROMOSOME NUMBER.-2n = 78. 

DISTRIBUTION.<O~S~~~ Kenya and Tanzania (including 

Pemba Island) (Map 16). 

SPECIMENS SEEN.-KENYA. KILIFI: Sokoke-Arabuko Forest on Malindi-Mombasa road, just N of 50 mi sign, Evans & 

Maikweki 51 (EA); Pangani Forest at crossing of Lwandani 

Stream on Ribe-Chonyi road, 3'523, 39'4032, Fuden & 

Beentje 85149 (US); Sokoke Forest, road to Jilore Forest 

Station, 3.2 km from turnoff on Kilifi-Malindi road, Faden & 

Evans 711714 (DSM, EA, K, MO, US, WAG); Chasimba, 

Kaloleni-Kilifi road -14 km, 3'43'S, 39'42%, Faden & Evans 

711726 (EA, K); same locality, Fuden & Faden 711788 (C, EA, 

FT, K, MO, PRE, US); Sokoke Forest, 21.3 km S of Gede 

turnoff, -3"26'S, 39'5392, Faden et al. 711633 (BR, C, EA, F, 

G, K, MO, PRE, US, WAG); Mnarani, S side of Kilifi Creek, 

Faden & Faden 711809 (B, BR, EA, F1: K, LISC, MO, P, PRE, 

US, WAG); 1 km NE of Pangani on Chonyi-Ribe road, 3'529, 

39'4095 cultivated Missouri Botanical Garden, Faden & 

Faden 721226A (BR, EA, K, MO); Sokoke Forest, -3 km on 

track S of Gede to Jilore Forest Station, -3'18'S, 39'57% 

Faden & Faden 7411215 (BM, EA, K, MO); Pangani, crossing 

of Lwandani Stream on Chonyi-Ribe road, 3'51'S, 39'412, 

Faden et al. 77/523 (EA); Kilifi, Jej4er-y K315 (EA, G, K); 

Marafa, 25 mi NW of Malindi, Polhill & Paulo 796 (BR, EA, 

K, PRE). KWALE: Mwachi Forest, N end, along Mwachi River, 

3'59'30"S,39'32'30"E, Fuden & Fuden 771484 PA); N side of 

Gogoni Forest, -4024'S, 39'28'~~ Fuden & Faden 771629 (US); 

Mwena River S of Marenje Forest, N side of river, 4'325, 

39'12%. Faden & Faden 771749 (BR, EA, F, K, MO, US, 

WAG); Muhaka Forest, N end, 4'20'S, 39'312, Faden & 

Fuden 771775 (EA, US). LAMU: Mambasasa, Utwani Forest 

Reserve, Greenway & Rawlins 9344 (EA, K). TANA RIVER: 
132 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1 

4'0 I 4'2 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

0 Oo% 

0 

// 

3 

0 100 MILES 

I 

I 

0 100 KILOMETERS 

I I 1 I 

MAP 16.-Distribution of Aneilema clorkei Rendle (solid circles), A. lamuenre Faden, new species (solid 

squares), and A. succulentwn Faden, new species (open circles). 
NUMBER 76 133 

Lake Dumi, 13 Feb 1893, Gregory s.n. (BM). 

TANZANIA. UWO: Near Magogoni, Vaughan 2395 

@A). PEMBA: Ngezi Forest, Vaughan 621 @A, K) & 2064 

0. 

DISCUSSION 

Rendle’s figure 8, which accompanied the type description 

of A. clarkei, is not of this species but of A. tanaense. Gregory 

collected the latter the same day he collected the type of A. 

clarkei. The remaining figures published by Rendle do belong 

to A. clarkei and are reproduced from drawings on the type 

specimen. Only Rendle’s figure 8 is not on the type (see 

discussion under A. tanaense). 

Plants of A. clarkei tend to form mats where locally 

common. Vegetative shoots can attain at least one meter in 

length. 

Clarke (1901) observed that the leaves in this species were 

“somewhat polymorphic.” While they are in fact no more so 

than in many other Aneilemu species, several factors combine 

to make this polymorphism more obvious in A. clarkei. The 

relatively short flowering shoots have towards their bases the 

usual transition from prophyll to mature leaf. In addition, the 

leaves are generally reduced towards the terminal inflorescence. Thus in a moderately short flowering shoot there may be 

two transitions in lamina size and shape. The net effect is that 

leaf shape polymorphism is more concentrated, hence more 

striking, in A. clarkei than in most other species in the genus. 

Every cincinnus in A. clurkei is the homolog of an entire 

thyrse. This is evidenced by the presence of two bracts at the 

base of the cincinnus, the larger, lower one the inflorescence 

bract, the upper the cincinnus bract (Figure 9). The uppermost 

cincinnus is the terminal inflorescence; it lacks the basal 

prophyll that characterizes the other, lateral cincinni, and lateral 

shoots in general. 

Two cincinni are frequently closely associated at the end of 

the flowering shoot, suggesting a larger inflorescence unit than 

the single cincinnus proposed here. In these examples the true 

terminal cincinnus has just the two bracts associated with it, 

while the subterminal, sessile, or subsessile cincinnus appears 

to have four. The lowermost of these is a foliage leaf. The next, 

visible only upon dissection, is a prophyll. The upper two are 

homologous to the two bracts of the terminal cincinnus. The 

prophyll indicates that the subterminal cincinnus is actually 

terminal on a very reduced lateral shoot and is not a part of the 

same thyrse as the terminal cincinnus. Of course, the two 

cincinni may be functionally related-as in the attraction of 

pollinators by simultaneously producing flowers-but their 

morphological relationship is equivalent to the loose association of thyrses found at the ends of the flowering shoots in 

many Aneilemu species. 

The retention of the lateral stamen filaments by the middle 

petal is a well-developed and nearly constant character in A. 

clarkei. Field notes on Faden & Faden 711809 indicate that in 

a rare flower the stamens are not held. Also rarely, one stamen 

is not released. 

The presence of dorsal capsule valves in mature fruiting 

specimens of A. clarkei shows that this valve is not regularly 

deciduous, The same interpretation has been used for other 

species in section Lamprodithyros, such as A. recurvatum and 

A. zebrinum. These conclusions are supported by Observations 

of these species in cultivation. In contrast, the dorsal capsule 

valves in other species of this section, such as A. calceolus, A. 

lamuense, and A. petersii, are usually deciduous. The dehiscence of the dorsal valve, or lack thereof, is probably related to 

seed dispersal (see p. 38). 

Aneilemu clarkei occurs in a variety of habitats. All of them 

are similar in having at least 760 mm rainfall (National Atlas of 

Kenya, 1970) and probably calcareous soils. 

Aneilema clarkei is readily distinguished from A. calceolus, 

with which Brenan (1961) contrasted it, by its inflorescence 

structure, distinctly perennial habit, pedicels recurved and 

puberulous only near the apex, larger and bluer flowers that 

nearly always have a medial staminode, and larger capsules in 

which the dorsal valve is often persistent. 

Similar inflorescence structure and fruiting pedicel curvature 

to those of A. clarkei are found only in A. succulentum and A. 

lamuense, which must be considered its closest relatives. The 

three species may be worth treating as a series within section 

Lamprodithyros, but further studies are required of other 

species before the section can be satisfactorily subdivided. 

Aneilema clarkei differs from A. succulentum by the former’s 

petiolate leaves with symmetric bases and larger capsules and 

seeds, from A. lamuense by its perennial habit, lack of 

uniseriate hairs on the pedicels and sepals, and yellow medial 

anther with yellow pollen, and from both in its larger, bluer 

flowers, better-developed stamen-retention mechanism, more 

divergent lateral stamen filaments, and less regularly deciduous 

dorsal capsule valves. 

11. Aneilema lamuense Faden, new species 

Herbae annuae foliis sessilibus vel breviter petiolatis, 

spiraliter dispositis vel distichis laminis ovato-ellipticis ad 

ovatas, lanceolatas vel lanceolato-ellipticas, (1-)2-4.5(-5.8) 

cm longis, 1-2(-2.5) cm latis. Inflorescentiae terminales in 

caule principali caulibusque lateralibus principalibus etiam in 

caulibus brevibus e nodis 1-3 summis caulium principalium 

prodientibus, cincinnis brevibus solitariis jug0 bractearum 

subtentis et partim inclusis compositae. Pedicelli 7-10 mm 

longi, in fructu valde recurvati (-180’) tantum prope apicem, 

piloso-puberuli prope apicem. Petalum medium cymbiforme 

vel cupulatum. Antherae omnes 3 pollen album continentes. 

Capsulae 3.6-4.3(-5) mm longae, 1.95-2.1(-2.55) mm latae, 

valva dorsali decidua, loculo dorsali 1-seminali, loculis 

ventralibus uterque 2-seminalibus. Semina loculorum ventralium ovata vel subtriangularia, (1.35-) I .45-1.7(-1.8) mm 
134 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

longa, 1.4-1.75 mm lata, testa scrobiculata non profunda vel 

sulcato-scrobiculato (Figure 54). 

TYPE.-Kenya, Lamu District Pana River District on label], 

Kitwa Pembe Hill and vicinity, 2027'S, 40'423, 15-16 Jul 

1974, Faden & Faden 7411083 (US, holotype; BR, C, EA, lT, 

K, MO, PRE, UPS, isotypes). 

Annual herbs (habit type IC, or occasionally IIIA, Figures 1, 

54u; Plate 4a). Roots fibrous. Shoots erect to ascending or 

sometimes decumbent, usually much branched, to -2q-40) 

cm tall, all shoots eventually terminating in inflorescences. 

Internodes (0.5-)1.5-6(-9) cm long, puberulous (sometimes 

only just below the nodes). Leaves spirally arranged on the 

main shoot, distichous (the 2 ranks at an obtuse angle to each 

other) on the lateral shoots (only -4-5 leaves produced on the 

larger lateral shoots), not reduced towards the terminal 

inflorescence, sheaths 3-9 mm long, puberulous and usually 

with a few uniseriate hairs (lower leaves) or densely pilosepuberulous (uppermost leaves), apex ciliate, laminae shortly 

petiolate or sessile, ovate-elliptic to ovate, lanceolate or 

lanceolate-elliptic, (1-)2-4.5(-5.8) cm long, 1-2(-2.5) cm 

wide, apex acute to acuminate, margin usually planar, scabrid, 

sparsely ciliate basally or on the petiole (when present), adaxial 

surface dull, abaxial lustrous, both surfaces pilose-puberulous, 

the uniseriate hairs longer on the adaxial surface than the 

abaxial. 

Inflorescences (Figure 54b) terminal on the main shoots and 

also on short, non-perforating shoots produced from the 

uppermost 1-3 nodes on the main and major lateral shoots 

(sometimes also terminal on more elongate, * leafy shoots 

produced from lower nodes), each inflorescence consisting of a 

short, solitary cincinnus subtended by and partially enclosed in 

a subopposite pair of herbaceous bracts, the larger, lower one 

the inflorescence bract, the upper the cincinnus bract. Peduncles to -1 cm long, puberulous. Inflorescence bract terminal, 

herbaceous. Cincinni to -5 mm long and 10-flowered. 

Cincinnus bracts herbaceous, lanceolate, falcate, erect to 

ascending, -8-20 mm long (including the sheath) x 2-5 mm 

wide, eglandular, both surfaces pilose-puberulous, margins 

longly ciliate. Cincinnus peduncles not developed. Cincinnus 

axes very short, completely covered by overlapping bracteoles, 

glabrous. Bracteoles attached up to 0.2 mm apart, membranous, 

lanceolate-ovate to ovate, often eccentrically cup-shaped, 

perfoliate or not, -4-6 mm long, to -2 mm high, eglandular but 

the apex thickened, lower (outer) ones pilose-puberulous, 

upper (inner) ones pilose only at the apex or completely 

glabrous. 

Flowers perfect and staminate, odorless, 10.5-14(-15.5) 

mm wide (Figure 54c,d; Plate 46,~). Pedicels 7-10 mm long, 

arcuate-erect to gently S-shaped in flower, in fruit, strongly 

recurved (-180') only near the apex and becoming much 

thickened in that region, persistent, pilose-puberulous near the 

apex, otherwise glabrous (Figure 54j). Sepals not clearly 

glandular; medial sepal lanceolate to lanceolate-ovate or 

lanceolate-elliptic, 2.8-3.6 mm long, 1.6-2.1 mm wide, 

generally glabrous, occasionally sparsely puberulous at the 

base: lateral sepals elliptic to obovate-elliptic or ovate-elliptic, 

3.5-4.3 mm long, 2-2.6 (-2.9) mm wide, pilose-puberulous 

along the basal '/2 of the medial (lower) edge, the long, 

uniseriate hairs very few. Paired petals (Figure 54e) 6-9 mm 

long, 5-7(-8) mm wide, limb ovate 4-5.5(-6) mm long, lilac 

(RHS color: 81C, C-D, D, Faden & Faden 74/1083), apex 

acute to obtuse or rounded, margin crenulate, claw 2-3 mm 

long, concolorous with limb apically, whitish basally, glabrous. 

Medial petal boat or cup-shaped, ovate, broadest at the margin 

(viewed from the apex), retaining the lateral stamens when the 

flower opens, 5-6(-7.5) mm long, 2-4.5(-6) mm wide, 

2.8-3.7(-3.9) m deep, whitish except basally where rt 

Lateral stamen filaments fused basally to medial stamen 

filament and lateral staminode filaments (Figures 28c,d, 54h), 

stamen filaments shortly adnate to medial petal base. Medial 

staminode filament 0.3-1(-1.5) mm long, pinkish purple, 

antherode bilobed, lobes sessile or subsessile, broadly ovate to 

reniform, 0.5-0.7 mm long, yellow with pinkish purple bases, 

connective slightly elongate (Figure 54f). Lateral staminodes 

with filaments 3.5-4.7 mm long, ventrally displaced, lilac, 

shading to white near the apex, antherodes bilobed, lobes 

stipitate, obovate to broadly ovate, reniform or transversely 

elliptic, 0.45-0.65 mm long, yellow with a pinkish purple or 

violet, cuneate base, connective very slightly elongate (Figure 

54g). Lateral stamens with filaments parallel in the perfect 

flowers, divergent above the middle (and somewhat longer) in 

the staminate flowers (Figures 28c,d, 54h), (5-)6-8.5 mm 

long, k S-shaped, decurved about the middle, glabrous, anthers 

lanceolate-elliptic to elliptic, 1.4- 1.7 mm long, 0.8-1 mm 

wide, pollen white. Medial stamen with filament 4.5-6(-6.3) 

mm long, anther lanceolate to oblong, saddle-shaped, 1.6-2 

mm long, 1-1.35 mm wide, entirely creamy white, pollen 

white. Ovary substipitate (Figure 549, obovate-elliptic to 

obovate, 1.5-1.75 mm long, 1-1.2 mm wide, densely and 

uniformly covered with patent, glandular hairs (mixed with 

some hook-hairs, best seen in fruit), apex rounded to truncate, 

dorsal locule very prominent, 1-ovulate, ventral locules each 

2-ovulate; style 7-8 mm long, nearly straight to gently 

arcuate-decurved for most of its length, also gently curved out 

of the floral midplane, sometimes with a few hook-hairs and/or 

glandular hairs near the base, stigma capitate, held slightly in 

front of the anthers and generally at the same level as them. 

concolorous with 3l e limbs of the paired petals (Figure 24H). 

ROWRE 54.-Aneilemu Iamuense Faden. new species: (I, habit; b, end of 

flowering shoot showing terminal inflorescence and uppermost lateral 

inflorescence; c, perfect flower, front view; d, perfect flower, side view; e, 

paired petal,f, medial staminode; g, lateral staminode, h, androecium of perfect 

flower, top view, medial staminode omitted, gynoecium removed; i, ovary and 

base of style, side view; j, capsule attached to dncinnus, side view; k, dorsal 

capsule valve, dorsal view; I, dorsal locule seed, ventral view; m, dorsal locule 

seed, dorsal view; n, ventral locule seed, ventral view; 0, ventral locule seed, 

dorsal view (all from Foden & Faden 7411083). (Bars = 20 mm foro; 4 mm for 

b; 2 mm for c-e; 1 mm forJh,j,k; 0.5 mm for ik) 
NUMBER 76 135 
136 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

Capsules substipitate (Figure 54j), obovate to obovateelliptic, dehiscent, bivalved, trilocular, 3.6-4.3(-5) mm long, 

1.95-2.1(-2.55) mm wide, tan with dark brown spots, 

lustrous, puberulous, apex emarginate, dorsal valve (Figure 

54k) deciduous, dorsal locule very prominent, 1- 

seeded (or occasionally empty), ventral locules each 2-(or, by 

abortion, l-)seeded, cells of the capsule wall transversely 

elongate. Dorsal locule seed (Figure 541,m; Plate 62) broadly 

elliptic, 1.65-2.15 mm long, 1.35-1.55 mm wide, -1 mm 

thick, hilum dark brown, arising from a very shallow groove, 

much shorter than the seed, testa orange-buff, heavily spotted 

and striped with dark brown (or maroon?), nearly smooth on 

the dorsal surface, not at all white-farinose to sparsely so 

around the embryotega and hilum, hypha-like filaments absent. 

Ventral locule seeds (Figure 54n,o; Plate 6i) ovate to 

subtriangular (sometimes somewhat 3-lobed), apical seeds 

rounded apically, basal seeds angular basally, (1.35-)1.45- 

1.7(-1.8) mm long, 1.4-1.75 mm wide, 0.9-1.2 mm thick, 

hilum dark brown, prominently raised within a groove, f not 

extended onto apical and basal surfaces, testa tan, orange-buff 

or orange-brown, usually heavily spotted and striped with dark 

brown (or maroon?), shallowly scrobiculate or sulcatescrobiculate on the dorsal surface, sparsely white-farinose in 

many of the depressions, densely so around the embryotega and 

hilum, hypha-like filaments sometimes present among the 

farinose granules. 

HABITAT.rOaStal sand dunes; -0-50 m. 

lXowEruNc.-Flowering specimens have been seen from 

July and August. In the field the flowers open 0900-0930 hrs 

and fade - 1300 hrs. 

CHROMOSOME NUMBER.-2n = 26. 

DISTRIBUTION.-N Kenya coast (Map 16). 

SPECIMENS SEEN.-KENYA. LAMU: Kitwa Pembe Hill and 

vicinity, 2'273, 40'42%, Fuden & Fuden 7411 083 (BR, C, EA, 

FT, K, MO, PRE, US, UPS); Lamu Island, Schlieben 12121 

6). 

DISCUSSION 

Schlieben's plants differ from the Fadens' in having shorter 

internodes and apparently somewhat fleshy leaves. They were 

probably growing more exposed to the effects of salt-laden sea 

breezes than were the Fadens' plants which came from the 

sheltered, leeward side of 45-60 m high sand dunes. 

Plants of A. lumuense show great variation in habit, ranging 

from unbranched, erect individuals a few centimeters tall to 

much-branched ones up to about 40 cm tall and 50 cm in 

diameter. In general, the shoots are erect to ascending; only in 

the largest plants do they become somewhat decumbent. 

The inflorescences have the same basic structure as those of 

A. clarkei and A. succulentum (Figure 9). Unlike either of these 

species, those of A. lamuense are nearly all sessile or subsessile. 

They show the same tendency as A. clurkei for the terminal 

inflorescence of the larger flowering shoots to become 

associated with the subterminal inflorescence-shoot (see 

discussion under A. clurkei). In A. lamuense, secondary 

inflorescences are often also produced in the mils of the 

inflorescence bracts of these two inflorescences. This can make 

interpretation of the inflorescence structure very difficult, as 

under these circumstances, 11 bracts (four inflorescence bracts, 

four cincinnus bracts, and three prophylls) plus a foliage leaf 

are present in addition to the four cincinni. The production of 

numerous inflorescences in A. lamuense (as compared with A. 

clarkei and A. succulentum) is consistent with its annual habit 

that places a high premium on seed production. 

The stamen retention mechanism is poorly developed in A. 

lamuense. In the field the period for which the lateral stamens 

were held by the medial petal could not be determined because 

visiting bees appeared to be instrumental in their early release. 

However, in cultivation the stamens were found to be released 

in less than an hour and often after only a few minutes, 

suggesting that the pollinators were not the cause of the early 

release. The anthers escape from a very narrow gap between the 

still involute, lateral margins of the medial petal. 

Aneilema lamuense is highly distinctive because of its 

inflorescence structure and fruiting pedicel curvature, charac - 

ters it shares only with A. clarkei and A. succulentum, its closest 

relatives, Aneilema lamuense differs from the former by its 

smaller flowers and narrower capsules, from the latter by its 

petiolate leaves and larger capsules and seeds, and from both by 

its annual habit, long, uniseriate hairs on the pedicels and 

lateral sepals, petal color, staminode lobes with pinkish purple 

or violet spots at their bases, and white medial anther with 

white pollen. It is also ecologically distinct from both. 

12. Aneilema succulentum Faden, new species 

Herbae perennes internodiis succulentis foliis succulentis 

distichis laminis sessilibus anguste lanceolato-ellipticis vel 

ellipticis, (1.3-)1.5-4 (-6) cm longis, (0.4-)0.7-1.3(-1.9) cm 

latis. Inflorescentiae terminales in caulibus floriferis etiam in 

caulibus brevibus e nodis 1-4 summis caulium floriferorum 

prodientibus, cincinnis brevibus solitariis jug0 bractearum 

subtentis et partim inclusis compositae. Pedicelli 7- lo(- 13) 

mm longi, in fructu valde recurvati (-180') tantum prope 

apicem. Petalum medium cymbiforme vel cupulatum. Filamenta stamineum et staminodiorum basi connata. Capsulae 

(2.8-)3.2-3.7(-4.3) mm longae, (1.5-)2.1-2.3(-2.5) mm 

latae valva dorsali saepe decidua, loculo dorsali 1-seminali, 

loculis ventralibus uterque 2-seminalibus. Semina loculorum 

ventralium subtriangularis vel ovata, 1.3-1,5(-1,6) mm longa, 

1.3-1.5 mm lata, testa scrobiculata non profunda (Figure 55). 

TYPE.-Kenya, Tana River District Lamu District on label], 

Garsen-Witu road, near Nyangoro Bridge, 2'18'S, 40'20'E, 

20-21 Jul 1974, Fuden & Fuden 7411152 (US, holotype; BR, 

EAm, K, MO, PRE, isotypes). 

Perennial herbs (habit type IIA2, Figures 2, 551). Roots 
NUMBER 76 137 

fibrous. Vegetative shoots repent, sometimes with looping 

internodes, much branched, often forming mats; flowering 

shoots ascending, unbranched or little branched, to -15-30 cm 

tall. Internodes (0.5-)1-5(-8) cm long, often suffused with 

maroon, puberulous, very succulent, circular to semicircular or 

elliptic in transverse section, becoming swollen in the rainy 

season, the nodes then appearing as constrictions. Leaves 

distichous (the 2 ranks at acute or right angles to each other), 

usually not reduced apically on the flowering shoots, sheaths 

2-6(-10) mm long, pilose-puberulous, ciliate at the apex, 

laminae succulent, often slightly falcate, often somewhat 

conduplicate, sessile, narrowly lanceolate-elliptic to elliptic 

(rarely ovate-elliptic), (1.3-) 1.5-4(-6) cm long, (0.4-)0.7- 

1.3(-1.9) cm wide, apex acute, base slightly asymmetric, 

cuneate, margin planar or undulate, scabrid, sparsely ciliate 

towards the base, often maroon, both surfaces slightly lustrous, 

adaxial frequently mottled with maroon, puberulous or pilosepuberulous, abaxial pilose-puberulous, the uniseriate hairs 

generally shorter (occasionally equal to) and more numerous 

than those of the adaxial surface. 

Inflorescences (Figure 55b; Plate 4e) terminal on the 

flowering shoots and also on short, non-perforating shoots 

produced from the uppermost 1-4 nodes on the flowering 

shoots, sometimes longer lateral shoots, produced from the 

lower nodes of the main flowering shoot, also bearing 1 or 2 

inflorescences in the same manner as the main flowering shoot; 

each inflorescence consisting of a short, solitary cincinnus 

subtended by, and partially enclosed in, a subopposite pair of 

herbaceous (at least at their apices) bracts, the larger, lower one 

the inflorescence bract, the upper the cincinnus bract. Peduncles -0-4 cm long (to 7 cm long in cultivation), green, 

puberulous. Inflorescence bract terminal, herbaceous, slightly 

falcate. Cincinni to -5 mm long and 10-flowered (to 7 mm long 

and 20-flowered in cultivation). Cincinnus bracts completely 

membranous or herbaceous apically and membranous basally, 

linear-lanceolate to lanceolate, falcate, erect to ascending, 

10-22 mm long (including the sheath), 1.5-5 mm wide, 

eglandular, densely pilose-puberulous on the adaxial surface, 

puberulous (occasionally sparsely pilose as well) on the abaxial 

surface, margin ciliate. Cincinnus peduncles not developed. 

Cincinnus axes very short, completely covered by overlapping 

bracteoles, glabrous. Bracteoles attached up to 0.2 mm apart, 

membranous, strongly eccentrically cup-shaped, perfoliate, 

3.25-7 mm long, 1-3 mm high, slightly thickened apically but 

apparently eglandular, sparsely puberulous apically with 

uniseriate hairs and/or hook-hairs, rarely glabrous. 

Flowers perfect and staminate, odorless, 9-14(-17) mm 

wide (Figure 55c,d; Plate 4f). Pedicels 7-lo(-13) mm long, 

arcuateerect to gently S-shaped in flower, in fruit, strongly 

recurved (-180') only near the apex and becoming much 

thickened in that region (Figure 55j), persistent, puberulous 

near the apex, otherwise glabrous. Sepals inconspicuously 

glandular or apparently eglandular near the apex (glands, when 

present, unlobed) glabrous, or puberulous basally; medial sepal 

lanceolate to lanceolate-elliptic, 3.5-4(-4.5) mm long, 1.8- 

2.3 mm wide; lateral sepals elliptic to ovate, 4-4.5(-5.5) mm 

long, 2.5-3.2 mm wide. Paired petals 7-8(-10.5) mm long, 

5-7(-9) mm wide (Figure 554, limb ovate-elliptic to ovate or 

reniform, 4.5-53-7) mm long, white to pale lilac (RHS 

colors (those too pale to be measured omitted): 76C, Faden et 

al. 701937 and 72139, both cultivated; 76C-D, Evans & 

Maikweki 54, cultivated, Faden & Faden 7411 152 and 7411 186, 

both field & cultivated; 76D, Faden & Faden 7411152, 

cultivated), apex subacute to obtuse or rounded, claw 1.5-2.5 

(-3.5) mm long, white or whitish, glabrous. Medial petal 

boat-or cup-shaped, sometimes k carinate medially, ovate to 

obovate, broadest at the margin (viewed from the apex), which 

is generally slightly revolute towards the apex, frequently 

retaining the lateral stamens when the flower opens, 5.5-7 

(-9.5) mm long, 3-5(-7) mm wide, (2-)2.5-3(-5) mm deep, 

concolorous with the limbs of the paired petals when they are 

whitish, distinctly paler than the latter when they are pale lilac 

(Figure 24~'). Lateral stamen filaments fused basally to 

medial stamen filament and lateral staminode filaments 

(Figures 28b, 55h), stamen filaments shortly adnate to the 

medial petal base. Medial staminode filament 0.5-2 mm long, 

white or greenish, often shading to yellow at the apex, 

antherode bilobed, lobes sessile, ovate, obovate, reniform, 

lanceolate or transversely elliptic, 0.15-0.4 mm long, connective not at all to slightly elongate (Figure 55f). Lateral 

staminodes with filaments 3-4(-5) mm long, ventrally 

displaced, white or whitish, shading to yellow apically, 

anthercdes bilobed, yellow, lobes sessile to shortly stipitate, 

obovate-cuneate to transversely elliptic or subreniform, 0.4- 

0.9 mm long, connective slightly to not at all elongate (Figure 

55s). Lateral stamens with filaments parallel to slightly 

divergent, sometimes (always?) slightly longer and more 

divergent in staminate than in perfect flowers (Figures 28b, 

55h), 5-6(-7) mm long, S-shaped, arcuatedecurved or 

sharply geniculate near the middle, anthers ovate to elliptic or 

lanceolate-elliptic, (0.7-)0.9-1.4 mm long, (0.45-)0.6-1 mm 

wide, pollen white or, less commonly, orange-yellow. Medial 

stamen with filament 4-5(-5.5) mm long, anther lanceolate to 

ovate or ovate-elliptic, saddle-shaped, (1.2-)1.5-2.3 mm long, 

1-1.2 mm wide, pollen orange-yellow. Ovary substipitate to 

shortly stipitate (Figure 55i), obovate to obovate-oblong or 

obovate-elliptic, 1.3-1.6 mm long (including the stalk or 

cuneate base), 0.9-1.1 mm wide, densely and uniformly 

covered with patent, glandular hairs, rarely mixed with 1 (or 

more?) hook-hair(s), apex rounded to truncate, dorsal locule 

prominent, 1-ovulate, ventral locules each 2-ovulate; style 

5.5-7 mm long, very gently arcuate-decurved, usually held 

slightly out of the floral midplane, sometimes with a few 

glandular hairs towards the base, stigma capitate, held in front 

of the anthers and slightly above to slightly below them. 

Capsules (Figure 55j) substipitate to shortly stipitate, elliptic 

to obovate-elliptic, carinate midventrally, dehiscent, bivalved, 

trilocular, (2.8-)3.2-3.7(-4.3) mm long, (1.5-)2.1-2.3(-2.5) 
138 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

mm wide, grayish tan, frequently with small, dark brown spots 

or mottling, lustrous, puberulous, apex truncate to emarginate, 

dorsal valve (Figure 55k) frequently deciduous, dorsal locule 

prominent, 1-seeded (rarely empty), ventral lccules each 2-(or, 

by abortion, 1-) seeded, cells of the capsule wall transversely 

elongate. Seeds (Figure 55l-o; Plate 6j) broadly elliptic to 

broadly ovate, 1.35-1.55 mm long, 1.2-1.25 mm wide (dorsal 

locule seed) or subtriangular to ovate (sometimes somewhat 

3-lobed), with apical seeds rounded apically and basal seeds 

angular basally, 1.3-1.5(-1.6) mm long, 1.3-1.5 mm wide 

(ventral locule seeds), 0.8-1 mm thick, hilum dark brown, 

prominently raised within a groove, not extended onto apical 

and basal surfaces, testa tan or grayish tan, sometimes marked 

with brown spots and streaks, shallowly scrobiculate (the pits 

sometimes distinctly elongate), white-farinose around the 

embryotega and hilum and frequently sparsely so in some of 

the depressions, hypha-like filaments sometimes present 

among the farinose granules. 

I-IABITAT.--D~~ deciduous bushland and bushland thicket 

dominated by species of Comphora, Euphorbia, Lmnea, 

Boswellia, Combretum, Grewia, Sansevieria, or, less commonly, Diospyros or Acacia; also in Brachystegia forest or 

woodland and along roadsides: sandy or clayey soils, occasionally on rocks; usually in partial shade or full sun; -10-600 m. 

FLowERING.-Flowering specimens have been seen from 

January, April, May, July, October, and December. In the field 

the flowers open 1130-1230 hrs and fade 1445-1530 hrs 

(Figure 25). 

CHROMOSOME NUMBER.-" = 13,2n = 26. 

DISTRIBUTION.-E Kenya (might be expected in NE 

Tanzania) (Map 16). 

SPECIMENS SEEN.-KENYA. KILIFI: Kaloleni-Mariakani 

road, Evans & Maikweki 54, cultivated in Nairobi (EA); Sala 

Gate-Malindi road, 7 km towards Sala Gate from junction with 

Bamba-Kakoneni road, -3'113, 39'452, Faden & Faden 

72/30 (EA, K, MO); Kakoneni-Jilore Forest Station road km 

2.5, 3'113, 39'52.5'E, Faden et al. 72139 (EA, K, MO, P, US, 

WAG); Malindi, Rawlins 863 (EA, K). KWALE: Near Tam, 

Samburu-Mackinnon Road, km 8, Faden & Evans 701782 

(EA, K, MO); 1 km S of Taru on Mombasa-Nairobi road, 

3'463, 39'072, Faden & Faden 72191 (EA, K); NairobiMombasa road, 5.3 km towards Mombasa from Taru, -3'463, 

39"10'E, Faden & Faden 7411281 (DSM, EA, K, MO); 

Mombasa-Nairobi road, 2.5 krn towards Mombasa from 

turnoff to Maji ya Chumvi Railway Station, 3'49'S,39"2O'E, 

Faden & Faden 771574 (EA, F, US); Mackinnon RoadSamburu, 19.4 km, Faden et al. 701937 (B, EA, FT, K, MO, 

US, UPS); Taru Desert, Taru, Tweedie 1202 6). TANA RIVER: 

Garsen-Witu road, near Nyangoro Bridge, 2'189, 40020'E, 

Faden & Faden 74/1152 (BR, EA, FT, K, MO, PRE, US); 

Garsen-Malindi road, 1.5 km towards Malindi from turnoff to 

Oda, 2'323, 4Oo07'3o"E, Faden &'Faden 7411186 (C, EA, K, 

MO, US, WAG); Garsen-Malindi road, -8 km from Garsen, 

-2'229, 40'07'E. Faden et al. 721227 (EA, K); Kurawa, 30 mi 

S of Garsen, Polhill & Paulo 629 (BR, EA, K, PRE); Lamu 

area, Malango ya Simba Forest, Rawlins 454 (EA, K). TEITA: 

Mackinnon Road-Bachuma, 9.6 km, 3'405, 38'58'E, Faden 

& Evans 71/620 (BR, EA, K); Nairobi-Mombasa road, 11.7 

km towards Mombasa past Maungu Station, -3'37'S,38'5O'E, 

Faden & Faden 741524 (EA, K, MO, US); Mt. Kasigau, lower 

slopes of S side above Bungule, Faden et al. 701969 (EA); Tam 

desert near Maungu, Tweedie 3364 (K). 

DISCUSSION 

Aneilema succulentum is one of the most succulent species in 

the genus, although much less so than some other Commelinaceae. In the dry season the thick internodes become laterally 

compressed due to the loss of water. The leaves also store 

water, particularly in the vertically elongate cells of the upper 

epidermis. In this character and in their general fleshiness they 

resemble the leaves of A. zebrinum. 

The inflorescences are identical in structure to those of A. 

clarkei and A. lamuense (see discussion under the former and 

Figure 9). The flowering shoots of A. succulentum tend to be 

more branched and therefore to produce more inflorescences 

than those of A. clarkei. In addition, the last two inflorescences 

on a flowering shoot show very little tendency to become 

paired as frequently happens in both A. clarkei and A. 

lamuense. 

The flowers of A. succulentum are considerably larger in 

cultivation (under a daily watering regimen) than in the field. 

The greatest dimensions for various floral parts given above 

(usually in parentheses) are mostly from flowers in cultivation. 

The normal range is taken from field collected flowers. 

The presence or absence of subapical glands on the sepals is 

not always readily determinable. In some populations they are 

clearly present on all three sepals. In others they appear to be 

lacking on the lateral sepals and perhaps also on the medial. 

The distinct absence of glands from the lateral sepals in the 

latter case can be observed in flowers preserved in FA A, which 

acts as a partial clearing agent for sepals. Anatomical studies 

are needed to determine more precisely what constitutes 

"presence" or "absence" of glands. 

The stamen retention mechanism of the medial petal is 

poorly developed in A. succulentum. It has been noted both in 

field populations and cultivated plants that frequently some 

flowers have one or both stamens free when the flowers open. 

FIOURe 55.-Aneilema succdentum Faden. new species: a, habit; b, dry 

inflorescence; c, perfect flower, front view; d. perfect flower, side view; e, 

paired petal,f, medial staminode, g, lateral staminode; h, androecium of perfect 

flower, top view, medial staminode omitted, gynoecium removed; i, ovary and 

base of style, side view; j, mature bud and capsule attached to cincinnus, side 

view; k, dorsal capsule valve, dorsal view; I, dorsal locule seed, ventral view; rn, 

dorsal locule seed, dorsal view; n, ventral locule seed, ventral view; 0, ventral 

locule seed, dorsal view (all from Faden & Faden 74/1152). (Bars = lorn for 

a; 4 mm for b; 2 mm for c-e; 1 mm for fAj,k; 0.5 mm for i,cO.) 
rW 
140 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Furthermore, in no case are the stamens held for more than half 

an hour (Figure 25). 

Pollen grains of the lateral anthers may be white or 

orange-yellow. In the latter case they are concolorous with the 

pollen of the medial anther. White appears to be the common 

(or exclusive?) color of lateral anther pollen observed in the 

field. Orange-yellow pollen is common in cultivated plants 

(including those observed in the field to have white pollen). 

The possibility of the orange-yellow color being an artifact of 

cultivation needs further investigation. 

The occurrence of A. succulenturn in Brachystegia woodland 

in the Kilifi District is interesting, because the closely related A. 

clarkei also frequently grows under Brachystegia in the same 

district. However, the soils and understory shrubs and herbs are 

quite different where each of the two occurs. Most notable is 

the presence of numerous succulents--e.g., Euphorbia grandicornis (= breviarticulata) is the dominant understory shrub-in 

those Brachystegia habitats where A. succulentum is present. 

Succulents are completely lacking in the Brachystegia woodland containing A. clarkei. The two species have not been 

found together. 

In the inland localities (Teita and Kwale districts) and in 

Kilifi District, A. succulentum occurs only in sandy soil or 

occasionally in well-drained rocky habitats. In Tana River 

District it grows mostly on a clayey alluvium mixed with sand. 

Aneilema succulentum appears to have two flowering 

periods inland, April to June and October to December or 

January, corresponding to the two rainy seasons in this region. 

In the Tana River District it may have one long flowering 

season (April to November?) or perhaps two. 

Sterile plants of A. succulentum can readily be confused with 

A. zebrinum, and the two have been found growing together 

near Taru and not far from one one another on Mt. Kasigau. 

Aneilema succulentum can be distinguished by its thicker 

internodes, proportionally narrower leaves with asymmetric 

bases, and the two ranks of leaves at acute or right angles to one 

other. 

Aneilema succulenturn is distinctive because of its vegetative 

and inflorescence morphology and fruiting pedicel curvature. 

The latter two characters are otherwise known only in A. clarkei 

and A. lamuense. It differs from the former by its smaller 

flowers, from the latter by its perennial habit, and from both by 

its more succulent internodes, the two ranks of leaves arranged 

at acute or right angles to each other, more succulent and sessile 

leaves with asymmetric bases, color of the paired petals, 

general lack of hook-hairs on the ovaries and capsules, and 

smaller capsules and seeds. 

13. Aneilema zebrinum Chiovenda 

Aneilema zebrinwn Chiovenda [“zebrina”], Webbia, 8:38, fig. 12, p. 39. 

1951.4bemeyer and Faden in Leistner, F1. Southem Africa, 4(2):40, figs. 

8,9-3a,b. 1985. [Syntypes: Ethiopia, Gemu-Gofa Prov., rive del Caschei, 5 

Jul 1939, Corradi 2154 (FT; photo K); same locality, 4 Jul 1939. Corradi 

2163 (FT; photo K); Cashei, 6 Jul 1939, Corradi 2155 (W, photo K: 

lectotype of Brenan, Kew Bull., 19:67,1%4).] 

Ballya zebrim (Chiovenda) Brenan, Kew Bull., 1964, map 1, p. 65. fig. 1, p. 

66, 1964.--Cufodontis, Bull. Jard. Bot. Natl. Belgique, 41, suppl., 1519, 

1971.-Ross, F. Natal, 117, 1972.-Faden in Agnew, Upland Kenya Wild 

Fl., 653. 1974. 

Perennial herbs (habit type 1%. Figure 2). Roots fibrous. 

Shoots repent, sparsely branched (usually unbranched for some 

distance behind the shoot apex) (Plate 4s). Internodes slender, 

(0.6-)1-3(-4) cm long, puberulous. Leaves distichous (the 2 

ranks usually at an obtuse, or rarely acute, angle to each other), 

sheaths 3-9 mm long, puberulous and usually sparsely pilose, 

ciliate at the apex, laminae somewhat succulent, sessile, ovate 

to ovate-elliptic or occasionally elliptic, 1-3.5(-4) cm long, 

0.5-2 cm wide, apex acute or occasionally obtuse, thickened 

and usually mucronate, base symmetric, usually amplexicaul, 

margin usually planar, ciliolate, scabrid in dried specimens but 

not in life, sparsely ciliate towards the base, both surfaces 

lustrous, puberulous with a mixture of equal-sized hook-hairs 

and uniseriate hairs, adaxial surface with very pale veins, often 

mottled with maroon between the veins when not so mottled, 

often gray-green (Plate 4g). 

Inflorescences axillary , perforating the leaf sheaths (Plate 

4h), generally consisting of one (rarely 2), short cincinnus 

subtended by 3 bracts, the basal a prophyll, the middle the 

inflorescence bract, the apical the cincinnus bract, frequently a 

secondary inflorescence, consisting of a single cincinnus with 

2 or 3 bracts at its base, developing from the axil of the 

inflorescence bract, rarely another secondary inflorescence, 

consisting of one cincinnus, developing from a prophyllar bud. 

Peduncles up to 8 mm long (often not developed), puberulous. 

Inflorescence bract basal to apical, bract-like, membranous. 

Inflorescence axis puberulous. Cincinni to -2 cm long and 

4(-6)-flowered. Cincinnus bracts erect to ascending, membranous, lanceolate to ovate, -2-3.5 mm long, eglandular, 

puberulous with ciliate or ciliolate margins. Cincinnus peduncles 1-5(-8) mm long, puberulous. Cincinnus axes puberulous, Bracteoles attached 0.5-2(-4.5) mm apart, membranous, 

eccentrically cup-shaped, perfoliate, 1.2-2(-2.2) mm long, 

0.3-1 mm high, eglandular, densely puberulous with hookhairs and uniseriate hairs. 

mm wide (Plate 4h). Pedicels (0.5-)1.5-3.5 mm long (to 5 mm 

long in fruit), ascending to erect in flower, erect to slightly 

further recurved in fruit, puberulous. Sepals apparently 

eglandular, puberulous with a mixture of hook-hairs and 

uniseriate hairs, the latter sometimes absent from the lateral 

sepals; medial sepal ovate-elliptic to lanceolate-elliptic or 

ovate, 2-2.7 mm long, (1-)1.5-1.6 mm wide; lateral sepals 

ovate-elliptic to elliptic or oblong-elliptic, 2.3-3 mm long, 

1.5-1.9 mm wide. Paired petals held in a lateral position, 4-5.7 

mm long, 3.1-4.5 mm wide, limb ovate to broadly ovate, 

2.5-3.7 mm long, pale lilac (RHS colors: 75B, Faden & Faden 

74482; 76B-C, Same collection, cultivated, Faden & Faden 

Flowers perfect and (very rarely) staminate, odorless, 7-10 ’ 
NUMBER 76 141 

7411052; 76C, Fa&n et al. 741205, Faden et al. 741330, 

cultivated, Faden & Faden 741533), apex rounded, margin k 

entire, claw 1.25-2 mm long, whitish, glabrous. Medial petal 

cup-shaped, ovate, broadest at or near the margin (viewed from 

the apex), not retaining the stamens when the flower opens, 

3.5-4.7 mm long, 2.7-4 mm wide, 1.7-2 mm deep, 

concolorous with the limbs of the paired petals (Figure 24j-1). 

Lateral stamen filaments fused basally to medial stamen 

filament and to the lateral staminode filaments, medial 

staminode nearly free (Figure 27c). Medial staminode usually 

completely hidden within the medial sepal, filament (1-)1.5- 

1.8 mm long, white, shading to yellow at the apex, antherode 

bilobed, yellow, lobes sessile, obovate-cuneate to suborbicular 

or transversely elliptic, 0.15-0.3 mm long, connective not 

elongate. Lateral staminodes with filaments (2-)2.2-2.6 (-3.5) 

mm long, displaced ventrally, white to very pale lilac, shading 

to yellow at the apex, antherode bilobed (or occasionally 

unlobed and clavate), yellow, lobes sessile, ovobate-cuneate, 

0.1-0.4 mm long, connective not elongate. Lateral stamens 

nearly parallel for their entire length or, more commonly, 

parallel or divergent from the fused portion to above the 

middle, then convergent apically, 3-3.5(-4.5) mm long, * 

S-shaped, decurved above the middle, glabrous, anthers often 

in contact with each other and with the medial anther, ovate to 

broadly ovate, 0.5-0.55 mm long, 0.45-0.6 mm wide, pollen 

yellow. Medial stamen with filament 2.5-2.8(-3.6) mm long, 

anther ovate to broadly ovate or ovate-quadrate, slightly 

saddle-shaped, 0.6-0.85 mm long, 0.6-0.8 mm wide, pollen 

yellow. Ovary substipitate, obovate to obovate-elliptic, 

(1-)1.2-1.5 mm long, (0.75-)0.9-1 mm wide, densely and 

uniformly covered with a mixture of glandular hairs and 

hook-hairs (the latter generally more numerous) (Figure 421,m), 

mely mixed with a few uniseriate hairs, apex rounded, dorsal 

locule l-ovulate, ventral locules each 2-ovulate, style 2.7-3.2 

(-3.6) mm long, arcuate-decurved for most of its length, held 

in the floral midplane or slightly curved laterally out of it, 

stigma capitate, often touching the anthers, but sometimes well 

below them. 

Capsules substipitate, obovate-elliptic, indehiscent or partially dehiscent along the lateral sutures, beginning at the base, 

bivalved (when partially dehiscent), trilocular, 3-4 mm long, 

2.1-2.5(-3) mm wide, stramineus or rarely chocolate brown, 

lustrous, puberulous, apex obtuse to rounded or truncate, valves 

persistent, dorsal locule a low hump, 1-seeded (or occasionally 

empty), ventral locules each 2-(or, by abortion, 1-)seeded, cells 

of the capsule wall transversely elongate. Seeds (Plate 7a) 

broadly elliptic to reniform, 1.4-1.8 mm long, 1.05-1.25 mm 

wide (dorsal locule seed), or ovate to ovate-cordate, with apical 

seed rounded apically and basal seed rounded or angular 

basally, 1.25-1.6 mm long, (1.1-)1.2-1.5 mm wide (ventral 

locule seeds), 0.5-0.6 mm thick, hilum slate gray or black, in 

a very shallow groove or not in a grwve, not extended onto 

apical or basal surfaces, testa gray or grayish brown, shallowly 

reticulate on the dorsal surface, sparsely tan-farinose around the 

embryotega and occasionally around the hilum, or occasionally 

farinose granules completely lacking. 

HABITAT.-Deciduous or semi-evergreen bushland and 

bushland thickets often dominated by species of Acacia and/or 

Commiphora, also in woodland, woodland thickets and light 

forest, sometimes in rocky arms, occasionally on steep slopes; 

sandy or clayey soils; partial or dense shade; 10-1150 m. 

RomRINa-Flowering specimens have been seen from 

January, February, April, May, July, and December. In the field 

the flowers open 1030-1115 hrs and fade 1315-1430 hrs. 

CHROMOSOME NUMBER.-n = 13,2n = 26. 

DISTRIBUTION.-SW Ethiopia to Tanzania, also in South 

Africa (N Natal) (Map 17). 

SPECIMENS SEEN.-ETHIOPIA. GEMU-GOFA: Rive del 

Caschei, Corradi 2154 (lT; photo K) & 2163 (lT; photo K); 

Caschei, Corradi 2155 (lT; photo K). 

KENYA. ELGEYO: Kerio Valley, Tot, Bully B12348 (K); 2.1 

km S of Tot, Faden et al. 701888 (EA, K, US, WAG). KWALE: 

Tanga-Mombasa road, mi 36, Bully B12193 (K); MombasaNairobi road, mi 30, Bully 813931 (EA, G); near Maji ya 

Chumvi, between Samburu and Mariakani, Bully & Smith 

B14320 (K); -1 km S of Taru on Mombasa-Nairobi road, 

3'46'S,39'07%, Faden & Faden 72192 (BR, EA, K, MO, US); 

5.3 km towards Mombasa from Taru on Nairobi-Mombasa 

road, -3'46'S,39"10'E, Faden & Faden 7411283 (EA, FT, K, 

MO, US); Lungalunga-Vanga road, -0.5 km, 4'333, 

35'07'30"E, Faden & Faden 771364 (EA, F, K, US, WAG); 

Mombasa-Nairobi road, 2.5 km towards Mombasa from 

turnoff to Maji ya Chumvi Railway Station, 3'49'S,39'203, 

Faden & Faden 771585 (EA, F, US). TANA RIVER: 13 km from 

Galole on Galole-Garsen road, -1'37'S, 39'59E', Faden & 

Faden 7411052 (BR, C, EA, FT, K, MO, PRE, UPS, US); 0.8 

km towards Garsen from turnoff to Kibusu on Malindi-Garsen 

road, 2'219, 40'07'E. Faden & Faden 7411176 (EA, K, MO, 

US). TEITA: 28 km on Voi-Taveta road from NairobiMombasa road turnoff, 3"30'S, 38'19'E, Faden & Faden 

741482 (MO); 18 km on Voi-Taveta road from NairobiMombasa road turnoff, -3'309, 38'24'E, Faden & Faden 

741533 (B, EA, K, MO, US); same locality, Faden & Faden 

771311 (EA, F, US); Maungu Hills, 3'38's. 38'45'E, Faden et 

al. 701183 (EA, K); Mt. Kasigau, lower slopes of S side above 

Bungule, Faden et al. 701966 (EA, K, MO); Mzinga Hill, Voi, 

Verdcourt 3892 (EA, K). 

TANZANIA. BAGAMOYO: Dar es Salaam-Morogoro road, 

-10 km E of Chalinze (-85 km W of Dar). Mhoro & Wingfield 

BM2411 (EA). PARE: Between N Pare Hill & N end of Lake 

Jipe, Bully B12145 (K; also cultivated at K). TANGA: 3krn 

towards Maramba from turnoff to Lelwa on Maramba 

(Ma1amba)-Korogwe road, 4'46's. 38'473, Faden et al. 

741330 (DSM, EA, K, MO, US). 

SOUTH AFRICA. NATAL: 2632 (Bela Vista): Ndumu Game 

Reserve, Mahemane Thicket, -26'53'S, 32'15'E (-CD), Faden 

et al. 741205 (BOL, EA, K, MO, NH, NU, PRE, US); Ndumu 

Game Reserve, E of Shokwe (-CD), Poole 1255 (K, MO). 2732 
142 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Lh 

I 

10 

1 

8 

1 

I 

j 3/ 40 

MAP 17.-Distributim of Aneiiema zebrinwn Chiovenda. 

(Ubombo): Mkuzi flats E of Jozini (-AC), Brenan 14243 (K); 

Sandforest (-AD), Strey 4752 (Pm). 

DISCUSSION 

Corradi 2155 was selected as the lectotype of this species by 

Brenan (1964), whose choice must be followed (International 

Code of Botanical Nomenclature, Article 8, 1983). This was 

clearly the best choice, as it is the sheet illustrated in the figure 

accompanying the type description, The selection of Corradi 

2154 by Cufodontis (1971) as the lectotype should be ignored. 

The monospecific genus Ballya, based on A. zebrinum, was 

distinguished from Aneilema by Brenan (1964) primarily on 

the basis of three characters: (1) Ballya flowers having an 

inverted symmetry of Aneilema flowers, i.e, medial petal 

uppermost, staminodes lowermost, etc., (2) Ballya inflorescences perforating the leaf sheaths vs. not perforating in 

Aneilema, and (3) Ballya stamen and lateral staminode 

filaments being fused basally vs. free in Aneilema. Of these 

characters the first has been found to be atypical of A. rebrinum 

itself. Its flowers usually have the same orientation as those of 

Aneilema, or at least not an inverted one. This has been 

observed in the field in a number of populations as well as in 

cultivation. The flowers in Brenan’s plant were inverted 

because the plant was cultivated as a hanging basket, and its 

unusually short pedicels prevented reorientation of the flowers 

to a more normal position. Such reorientation has been 

observed to occur in other species of Aneilema and even in 

other populations of A. rebrinum with longer pedicels. 

The remaining two characters, supposedly unique to Ballya, 

also occur in species of Aneifema. The stamen filament bases 

are fused in all species of section Lamprodithyros as well as in 

A. hockii of section Amelina. Most of these species also have 

the lateral staminode filaments at least shortly fused basally to 

the stamens, as in A. zebrinum. Perforating inflorescences occur 

in A. petersii subsp. pallidijlorum and A. tanaense (both section 

Lamprodithyros), although no Aneilema species produces them 

almost exclusively except A. rebrinum. A single terminal 

inflorescence noted on a long shoot of this species is further 

evidence of its basic similarity to other Aneifema species. 

Tomlinson (1966:387) used anatomical data to try to 

separate Ballya from Aneilema. He stated that Ballya differed 

from Aneilema by the former’s glandular microhairs having an 

indistinct basal septum and the distal cell usually being longer 

than the middle cell, in contrast to the basal septum “obvious” 

and the “distal [cell] often shorter than [the] middle cell” in 

Aneilema. He further distinguished A. zebrinum by its 

epidermal cells vertically elongate in transverse section, 

whereas those of other Aneilema species were not elongate. 

Tomlinson’s conclusions were based on a sample of 

Aneilema species that was small and unrepresentative of the 

genus as a whole. The seven or eight species he examined 

(listed in Tomlinson, 1969) did not include any of section 

Lamprodithyros. The present study does not support Tomlin- 
NUMBER 76 143 

son's conclusion about the distinctions between Ballya and 

Aneilema. The glandular microhairs of A. zebrinum have been 

found to usually have the distal cell shorter than the middle cell 

(see Tomlinson, 1966, figs. 6, 114), a character that is found in 

a number of Aneilema species, including A. succulentum 

(section Lamprodithyros). The distinctness of the basal septum 

is open to subjective interpretation. I could find no difference in 

this character between A. zebrinum and A. succulentum. The 

latter species also has vertically elongate upper epidermal cells 

very much like those of A. zebrinum. Thus Ballya is not distinct 

from Aneilema. 

The flowers of A. zebrinum are often borne under the leaves. 

The small space available for them probably accounts for their 

being tilted or more or less vertical rather than horizontal as in 

most other Aneilema species. The lateral position of the paired 

petals may also be the consequence of space constraints. 

Individual flowers remain open for about three hours. Aneilema 

zebrinum has probably been overlooked by collectors because 

of its prostrate habit and tendency to grow under shrubs. Even 

when spotted the plants may be passed over as sterile because 

of the inconspicuousness of the flowers and capsules. In East 

Africa there is the further problem that A. zebrinum bears a 

striking resemblance vegetatively to and may occur with A. 

succulentum and an undescribed species of Cyanotis. The 

undercollection of A. zebrinum undoubtedly explains the 

apparent range disjunction of 2400 km between northern 

Tanzania and South Africa recorded by Faden (1975). The 

recent report of this species from the Selous Game Reserve in 

southern Tanzania (Vollesen, 1980) has considerably reduced 

the gap and and has added evidence that the remaining gap is 

only an artifact due to the lack of collections. 

Aneilema zebrinum exhibits a number of characters that are 

unusual for the genus as a whole and clearly demonstrate its 

relationship to species of section Lamprodithyros. In addition 

to the fused filament bases, large, cup-shaped medial petal, 

perforating inflorescences, and vertically elongate epidermal 

cells mentioned above, these characters include the repent 

habit, eglandular bracteoles and sepals, ovary and capsule 

pubescence frequently composed predominantly of hook-hairs, 

indehiscent or partially dehiscent capsules, and strongly 

dorsiventrally compressed seeds. 

Although the strictly repent flowering shoots of A. zebrinum 

are unique in Aneilema, its mat-forming habit occurs in several 

species. The particularly strong vegetative similarity to A. 

succulenturn has been noted above. As in that species, the 

primary shoot has spirally arranged leaves. It branches 

profusely from the lower nodes, producing thereafter only 

repent shoots with distichous leaves. 

The bracteoles in A. zebrinum are eglandular, as in A. clarkei, 

A. lamuense, and A. succulentum, and unlike nearly all other 

Aneilema species. The absence of sepaline glands from A. 

zebrinum suggests an endpoint in the reduction series shown by 

the same three species: glands are normally present on all 

sepals in A. clarkei; they may be lacking on some or all of the 

sepals in A. succulentum; they are regularly absent from all 

three sepals in A. lamuenre. It should be noted that presence or 

absence of sepaline glands must be used with some caution: 

sepals with inconspicuous glands may be difficult to distinguish from truly eglandular ones. 

The abundance of hook-hairs on the ovaries and capsules of 

A. zebrinum, although very variable, is not matched by any 

other species in the genus. However, smaller numbers of 

hook-hairs among more numerous glandular hairs also occur in 

a number of species of section Lamprodithyros (A. tanaense, A. 

calceolus, A. lamuense (regularly), A. clarkei (frequently), and 

A. succulentum and A. indehiscens (rarely)). Ovarian hookhairs have otherwise been noted in Aneilema only in A. gillettii 

(section Amelina) and very rarely in A. somaliense (section 

Somaliensia). 

Indehiscent or partially dehiscent capsules and strongly 

dorsiventrally compressed seeds, present in A. zebrinum, also 

occur together in A. indehiscens (section Lumprodithyros). 

In addition to all of the above characters, several others 

suggest affinities of A. zebrinum with section Lamprodithyros 

and support its placement in that section. Its very reduced 

inflorescences are homologous to the lateral inflorescenceshoots of A. clarkei, A. lamuense, and A. succulentum. Its 

five-seeded capsules are common to nearly all species of the 

section, and its basic chromosome number, x = 13, is the same 

as that of the other species of section Lamprodithyros. One may 

also note that A. zebrinum, although very widespread, occurs 

entirely within the geographic ranges of other species of this 

section. 

Within section Lamprodithyros, A. zebrinum is most closely 

related to A. clarkei, A. lamuense, and A. succulentum and 

especially to the last species. The repent flowering shoots and 

strictly lateral inflorescences of A. zebrinum readily distinguish 

it from these species. 

Section 6: Brevibarbata Faden, new section 

Herbae perennes vel annuae foliis spiraliter dispositis. 

Inflorescentiae thyrsi plerumque densi vel modice densi. 

Bracteolae cupulatae, perfoliatae vel non perfoliatae. Petalum 

medium deminutum. Filamenta libra. Stamina lateralia filamentis sigmoideis, in dimidio distali subtiliter barbatis. 

Stamen medium anthera plerumque in magnitudine antheras 

staminum lateralium subaequanti sed in forma differens. 

Ovarium glabrum vel puberulum. Capsulae dehiscentes, lucentes, glabrae vel puberulae, valvis persistentibes, loculo dorsali 

haud evoluto vel 1-seminali, loculis ventralibus uterque 

1-3-seminalibus. 

TYPE SPECIES.-Aneilema beniniense (F'. de Beauvois) 

Kunth. 

Perennial or annual herbs with fibrous or tuberous roots. 

Leaves spirally arranged. 

Inflorescences thyrses, terminal and occasionally axillary 

from the inflorescence bract and upper leaves, usually dense or 
144 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

moderately dense. Inflorescence bract herbaceous or membranous, when membranous, commonly enclosing the young 

inflorescence. Inflorescence axis puberulous (rarely subglabrous) with hook-hairs of uniform length or of two sizes 

(rarely some long, uniseriate hairs also present). Bracteoles 

cup-shaped, perfoliate or not. 

Flowers usually perfect and staminate (rarely pistillate, 

perfect and staminate or all perfect). Fruiting pedicels generally 

erect or recurved to 150'(-180'), rarely decurved. Sepals 

glabrous or puberulous, distinctly glandular subapically. Paired 

petals white to lilac or lavender, or yellow to orange, glabrous. 

Medial petal reduced, generally inconspicuous. Filaments free. 

Staminodes with bilobed, yellow antherodes. Medial staminode reduced or absent, when present, its antherode commonly 

different in form from those of the lateral staminodes. Lateral 

staminodes with straight or arcuate filaments. Lateral stamens 

with filaments S-shaped, closely parallel or, more commonly, 

divergent, finely bearded in the distal half on the lower surface, 

generally with minute, inconspicuous hairs. Medial stamen 

with anther subequal to the lateral anthers in size but differing 

in shape (usually with a broader connective) and orientation. 

Ovary sessile to stipitate, glabrous or puberulous with 

glandular hairs, dorsal locule usually not developed, occasionally 1-ovulate, ventral locules each 1-3-ovulate, style arcuatedecurved to straight, generally recurved apically, stigma 

capitate. 

Capsules sessile to stipitate, dehiscent, bivalved, bilocular or 

occasionally trilocular, lustrous, glabrous or puberulous, valves 

persistent, dorsal locule (when developed) l-seeded or, by 

abortion, empty, ventral locules each 1-3-seeded, cells of the 

capsule wall commonly transversely elongate, occasionally 

isodiametric or rarely longitudinally elongate. Seeds very 

variable, in outline circular, elliptic, oblong, ovate or trapezoidal, hilum in a groove or not, testa smooth, rugose, ribbed, 

sulcate, reticulate, tuberculate or scrobiculate, farinose granules 

sometimes present, entire testa or parts of it sometimes covered 

with a thin whitish coating (fused farinose granules?). 

BASIC CHROMOSOME NUMBERS.-% = 10,13, and 15. 

DISTRIBUTION.-West Africa from N limits of the genus E to 

S Sudan, SW Ethiopia, W Kenya, and E Tanzania S to Angola, 

Zambia, Zimbabwe, Swaziland, and South Africa (E Cape 

Province); Aneilema umbrosum also occurs in the neotropics. 

Key to the Species of Section Brevibarbata Faden 

1. Paired petals yellow to orange. 

2. Fruiting pedicels recurved 270"; inflorescences lax or moderately lax; roots 

2. Fruiting pedicels erect or recurved to -150"; inflorescences dense; roots tuberous; 

laminae sessile. 

3. Testa smooth, lacking farinose granules .............. A. homblei 

3. Testa furrowed, with farinose granules. 

4. Leaves linear, less than 1 cm wide; flowers 5.5-8.5 mm wide ....... .............................. A. macrorrhizum 

4. Leaves lanceolate to lanceolate-elliptic, more than 1 cm wide; flowers 9-14 

mm wide .......................... A. welwitschii 

fibrous; laminae petiolate ..................... A. angolense 

1. Paired petals white to lilac or lavender. 

7. At least the lower laminae petiolate; cincinni always several-flowered; testa 

6. Seeds one per ventral locule; capsules as wide as long or wider. 

foveate-tuberculate or foveate-reticulate, brown. 

8. 

8. Sepals puberulous; annuals. 

Sepals glabrous; decumbent perennials ........... A. dispermum 

of the capsule wall transversely elongate ......... A. arenicola 

9. Pedicels 1.5-3 mm long, erect to slightly further recurved in fruit; cells 

9. Pedicels 2-8 mm long, strongly recurved (120-270') in fruit; cells of the 

10. Cells of the capsule wall longitudinally elongate; inflorescences 

capsule wall longitudinally elongate or isodiametric. 

composed of (6-)lo-20(-29) cincinni; seeds dark brown ...... ......................... A. brunneospermum 

10. Cells of the capsule wall f isodiametric to slightly longitudinally 

elongate; inflorescences composed of 3-13 cincinni; seeds pale 

pinkish gray ...................... A. schlechten' 

7. All laminae sessile; cincinni 1- or several-flowered; testa ribbed, whitish or 

pinkish. 

11. Slender annuals; cincinni all l-flowered; seeds globose. 
NUMBER 76 145 

12. 

12. 

Cincinnus bracts linear, 5-7 mm long ............ A. mortonu 

Cincinnus bracts ovate, 1-3 mm long .......... A. paludosum 

11. Tbberous rooted perennials; cincinni several-flowered, seeds longer than 

13. Stems slender, internodes mostly exposed near the inflorescence; bristly 

red hairs absent ...................... A. paludosum 

13. Stems more robust, internodes usually not exposed, if more slender and 

the internodes exposed, then bristly red hairs present on the internodes 

and sheathes ........................ A. setiferum 

14. Shoots decumbent; roots fibrous; laminae petiolate, lanceolate to ovate; 

wide. 

6. Seeds 2-3 per ventral locule; capsules longer than wide. 

capsules glabrous or puberulous; forest species. 

15. Seeds rugose-tuberculate; capsules puberulous; sepals puberulous ..... ................................ A. dregeanum 

15. Seeds reticulate-foveate; capsules glabrous or subglabrous; sepals glabrous 

or sparsely puberulous. 

16. Inflorescences moderately lax to moderately dense; fruiting pedicels 

often decurved; capsules with dorsal locule not developed, with ventral 

locules each 2-seeded; sepals sometimes sparsely puberulous ..... ............................... A. umbrosum 

16. Inflorescences dense; fruiting pedicels erect or further recurved; capsules 

with dorsal locule frequently developed, 1-seeded or empty, with ventral 

locules each 2-3-seeded; sepals glabrous. 

17. Leaf margins ciliate; capsules acute at apex, very sparsely puberulous ............................. A. silvaticum 

17. Leaf margins scabrid, not ciliate; capsules rounded to truncate at apex, 

glabrous ........................ A. beniniense 

14. Shoots erect to ascending, rarely decumbent; roots tuberous; laminae sessile or 

18. Plants with basal rosettes of leaves; leaf margins ciliate; flowers opening in 

the afternoon ...................... A. pomeridianum 

18. Plants lacking basal rosettes; leaf margins scabrid but not ciliate; flowers 

opening in the morning .................. A. lanceolatum 

petiolate, linear to lanceolate; capsules puberulous; savannah species. 

SPECIES CITATIONS AND DISTRIBUTIONS 

Aneilema angolense C.B. Clarke in Thiselton-Dyer, Fl. Tmp. Africa, 8:67,381, 

1901. [Angola.] 

Aneilema arenicola Faden, Bothalia, 15:94, 1984. [Mozambique to South 

Africa (N Natal).] 

Aneilema beniniense (F'. de Beauvois) Kunth, Enum. PI.. 4:73, 1843. 

[Basionym: Commelina beniniensis P. de Beauvois, Fl. Owar.,2:49, tab. 87, 

1816. Senegal to SE Ethiopia S to Angola and Zambia (Plate 443.1 

Aneilema brunneospermum Faden, Bothalia, 15:91, 1984. [Mozambique to 

South Africa (Transvaal and N Natal) and Swaziland (Plate 41).] 

Aneilema dispermum Brenan, Kew Bull., 7:198, 1952. [Tanzania to Malawi 

(plants from Cameroon and Fernando Po may not be conspecific).] 

Aneilema dregeanwn Kunth, Enum. Pl., 4:73. 1843. [Mozambique to South 

Africa (Cape and Natal).] 

Aneilema homblei De Wildeman, Feddes Repert. Spec. Nov. Regni Veg.. 

12:290, 1913. [Zaire.] 

Aneilema lanceolatum Bentham in Hooker & Bentham. Fl. Nigrit., 546, 1849. 

[Mali to Sudan S to Congo, Zaire, Uganda and NW Kenya.] 

Aneilema macrorrhizum Th. Fries, jr,, in R.E. Fries, Schwed. Rhodesia-Kongo 

Exped., 1:220, fig. 19a-b, p. 221, 1914. [SE Zaire and Zambia to S 

Tanzania.] 

Aneilema morfonii Brenan, Kew Bull., 22:387, 1968. [Ghana.] 

Aneilema paludosum A. Chevalier, Mem. Soc. Bot. France, 8:215. 1912. 

Aneilema pomeridianwn Stanfield & Brenan in Brenan, Kew Bull., 15:217, 

Aneilema schlechteri K. Schumann, Bot. Jahrb. Syst., 33:376, 1903. [SE 

Aneilema sefiferwn A. Chevalier, Mem. SOC. Bot. France, 8:215, 1912. [Mali 

Aneilema silvaficwn Brenan, Kew Bull., 7:203. 1952. [Nigeria to N Zaire.] 

Aneilema wnbroswn (Vahl) Kunth, Enum. PI., 4:71, 1843. [Basionym: 

Commelina umbrosm Vahl, Enum. Pl., 2:179, 1805-1806. Senegal to S 

Sudan S to Congo, Zaire and Uganda; also Panama to Venezuela and 

Grenada S to Bolivia and Brazil.] 

Aneilema welwifschii C.B. Clarke in De Candolle, Monogr. Phan., 3:229,1881. 

[Angola to S Tanzania and Mozambique (Plate 413.1 

[Senegal to Chad.] 

1961. [Ghana and Nigeria (Plate 4k).] 

Zimbabwe to South Africa (Transvaal).] 

and Guinea Bissau to Nigeria.] 

DISCUSSION 

Section Breviburbata is the largest and most complex 

section in the genus. It is characterized by small- to 

medium-sized flowers in which the distal half of the lateral 
146 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

stamen filaments is minutely and inconspicuously bearded and 

by the medial anther more or less equal to the lateral anthers in 

size (but differing in form). The section is mainly centered in 

western Africa, and it includes several widespread species that 

are morphologically and cytologically diverse, e.g., A. beniniense, A. umbrosum, and A. welwitschii. Section Brevibarbata 

comprises four species groups: a western African forest group 

(A. beniniense, A. silvaticum, A. umbrousm, and west African 

A. dispermum, which is probably distinct from the eastern 

African plant of the same name); a western African savannah 

group (A. lanceolatum, A. pomeridianum, A. setiferum, A. 

paludosum, and A. mortonii); a central African savannah and 

woodland group with yellow to orange flowers (A. welwitschii, 

A. homblei, A. macrorrhizum, and A. angolense); and an eastern 

and southern African group of diverse habitats (A. dispermum, 

A. dregeanum, A. schlechteri, A. arenicola, and A. brunneospermum). Although many of these taxa have been studied, 

especially by Morton (1966) and Faden (1984), and a number 

of infraspecific ma have been described, we are far from 

understanding the total variation in this section. 

Section 7: Pedunculosa Faden, new section 

Herbae plerumque annuae radicibus fibrosis, foliis spiraliter 

dispositis. Inflorescentiae thyrsi densi. Axis inflorescentiae et 

axes atque pedunculi cincinnorum puberuli trichomatibus 

unciformibus plerumque biformibus praediti, trichomatibus 

uniseriatis longisque saepe adjectis. Bracteolae haud perfoliatae apicibus longis, linexibus in glandibus clavatis terminantibus saepe praeditae. Petalum medium deminutum. Filamenta 

libra. Stamina lateralia filamentis sigmoideis, in dimidio distali 

dense barbatis. Stamen medium deminutum. Ovarium puberulum trichomatibus glandulosis praeditum sed medidorsaliter 

glabrum. Capsulae dehiscentes, lucentes, puberulae, loculo 

dorsali haud evoluto vel 1 -seminali, loculis ventralibus uterque 

1-3-serninalibus, cellulis parietis capsulae transverse elongatiS. 

TYPE SPECIES.-Aneilema pedunculosum C.B. Clarke. 

Annual or less commonly perennial herbs with fibrous roots 

and spirally arranged leaves. 

Inflorescences dense thyrses, terminal and axillary from the 

inflorescence bract and upper leaves. Inflorescence bract basal 

to medial, membranous, enclosing the young inflorescence. 

Inflorescence axis and cincinnus peduncles and axes puberulous with hook-hairs generally of two sizes, frequently a small 

number of long, uniseriate hairs also present. Cincinnus axes 

often abbreviated. Cincinnus bracts and bracteoles frequently 

with long, linear apices terminating in k clavate glands. 

Bracteoles never perfoliate. 

Flowers perfect and staminate (rarely some pistillate). 

Fruiting pedicels erect or recurved to -150°, persistent. Sepals 

prominently glandular subapically . Paired petals whitish to 

lilac or bluish purple, glabrous. Medial petal reduced, but 

sometimes conspicuous. Filaments free. Staminodes with 

bilobed, yellow antherodes; medial staminode (occasionally 

absent) similar to the lateral staminodes but filament shorter; 

lateral staminodes with filaments gently arcuate-decurved. 

Lateral stamens with filaments S-shaped, densely bearded in 

the distal half with long, usually brightly colored hairs attached 

mostly on the lower surface, the part of the filament to which 

the hairs are attached broadened laterally, anthers small, ovate. 

Medial stamen shorter than the lateral stamens, with a small, 

usually dumbbell-shaped anther with small, slightly polleniferous, distal anther sacs. Ovary sessile or subsessile, puberulous 

with forward-pointing to patent, capitate, glandular hairs 

predominantly on the ventral and lateral surfaces, glabrous 

middorsally , dorsal locule usually not developed, occasionally 

present and containing an ovule, ventral locules each 1-3- 

ovulate, style arcuate-decurved to straight, sometimes recurved 

apically, stigma capitate or not enlarged. 

Capsules sessile or subsessile, dehiscent, bivalved, bilocular 

or occasionally trilocular, lustrous, puberulous, apex rounded 

to truncate or emarginate, valves persistent, rarely the dorsal 

valve tardily deciduous, dorsal locule (when developed) 

l-seeded or, by abortion, empty, ventral locules each 1-3- 

seeded, cells of the capsule wall transversely elongate. Seeds 

mostly ovate to trapezoidal or elliptic, apical ventral locule 

seeds usually rounded apically, basal ventral locule seeds 

commonly angular basally, hilum in a deep groove or pit, testa 

rugose or shallowly scrobiculate, sulcate or reticulate-foveate, 

farinose granules commonly present. 

BASIC CHROMOSOME NUMBER.-x = 9. 

DISTRIBUTION.-SE Sudan and W Ethiopia, S to Namibia 

(Caprivi Strip), Botswana, Zimbabwe, and Mozambique. 

Key to the Species of Section Pedunculosa Faden 

1. Cincinnus bracts and bracteoles lacking linear, gland-tipped apices; inflorescences 

lacking long, uniseriate hairs. 

2. Leaves linear to linear-lanceolate, 0.3-1 cm wide; leaf sheaths lacking long hairs 

at the apex; hilum in a pit . . . . . . . . . . . . . . . . . . . . A. richardsiae 

2. Leaves ovate to lanceolate-elliptic, mostly 1.5-3.2 cm wide; sheaths with long 

hairs at the apex; hilum in a groove. . . , . . . , . . , . . , . . . . A. spekei 

1, Cincinnus bracts and bracteoles with linear, gland-tipped apices; inflorescences 

commonly with long, uniseriate hairs. 
NUMBER 76 147 

3. Leaves with sessile laminae. 

4. Flowers 8-10 mm wide; stamen filaments bearded with maroon hairs; capsules 

with ventral locules each 1-2-seeded ............... A. hirtum 

4. Flowers 13-15 mm wide; stamen filaments bearded with yellow hairs; capsules 

with ventral locules each 2-seeded ............. A. chrysopogon 

5. Ventral locules usually 1-seeded; solitary cincinnus present at base of peduncle 

of terminal thyrse, widely separated from other cincinni; petals subequal in 

width and k concolorous .................. A. minutiflorum 

5. Ventral locules 2-3-seeded (1-seeded only through abortion); no cincinnus 

present at base of peduncle of terminal thyrse; medial petal narrower than 

paired petals, differing in color from them. 

6. Seeds 3 per ventral locule, 3-lobed .............. A. nicholsonii 

6. Seeds 2 per ventral locule, unlobed or 3-lobed. 

7. Flowers 4-7.5 mm wide; medial staminode lacking; filaments bearded 

with reddish purple or maroon hairs; seeds 3-lobed . . A. termitarium 

7. Flowers 9-13 mm wide; medial staminode present; filaments bearded with 

bluish purple hairs; seeds unlobed. 

8. Annuals; dorsal locule well-developed, usually containing a seed , . , 

........................... A.pedunculosum 

8. Decumbent perennials; dorsal locule not developed .... A. leiocaule 

3. Leaves with petiolate laminae. 

SPECIES CITATIONS AND DISTRIBUTIONS 

Aneilema chrysopogon Brenan, Kew Bull., 15:222, 1961. [S Tanzania to N 

Zambia (Plate 4n).] 

Aneilema hirtum A. Richard, Tent. Fl. Abyssinicae, 3(5):343, 1850. [Ethiopia 

to Zaire, Zambia, Malawi, and Tanzania (Plate 40).] 

Aneilema leiocaule K. Schumann in Engler, Pflanzenwelt Ost-Afrikas C, 136. 

1895. [S Ethiopia S to E Zaire and Tanzania (and Zambia?) (Plate 4m).] 

Aneilema minutiflorum Faden, ined. [(= Aneilema sp. B of Faden in Agnew, 

Upland Kenya Wild Fl., 664,667,1974].-Faden &Evans 70J899; Tweedie 

759). W Kenya to Uganda and Tanzania.] 

Aneilema nicholsonii C.B. Clarke in Thiselton-Dyer, Fl. Trop. Africa, 8:70. 

1901. [S Kenya S to Namibia (Caprivi Strip), Botswana, Zimbabwe, and 

Mozambique (Plate 4p).] 

Aneilema pedunculoswn C.B. Clarke in De Candolle, Monogr. Phan., 3:228, 

1881. [S Tanzania to Zimbabwe and Mozambique.] 

Aneilema richardsiae Brenan, Kew Bull.. 15:219,1961. [S Tanzaniato E Zaire 

and N Zambia.] 

Aneilema spekei C.B. Clarke in Thiselton-Dyer, Fl. Trop. Africa, 8:71, 1901. 

[Sudan and Ethiopia S to Zaire, Zambia, and Tanzania.] 

Aneilema iermitarium Faden, ined. [(Faden & Faden 741139). S Tanzania and 

E Zaire to Zambia.] 

DISCUSSION 

Section Pedunculosu is very natural. Several unusual 

characters clearly define it: annual habit (except A. leiocaule), 

dense inflorescences, non-perloliate bracteoles that often 

terminate in a clavate, glandular apex, densely bearded (with 

long, brightly colored hairs) lateral stamen filaments, and 

reduced medial anther. The basic chromosome number, x = 9, 

is unique to this section and is recorded from all species that 

have been counted. Although the section is easy to recognize, 

separating the species is difficult, especially with dried 

specimens. The main taxonomic problems are the determination of the status of plants that resemble A. spekei but have 

bracteoles with linear apices, e.g., Renvoize & Abdallah 2251; 

the identification of a plant from northern Zambia that 

resembles A. leiocaule (Richards 1531); and a study of the 

variation in A. hirtum for which two chromosome numbers are 

known. 

Species of Uncertain Generic Position 

In addition to the species included in the seven sections of 

Aneilema recognized above and the species once placed in the 

genus but here with certainty or high probability excluded from 

Aneilema (see Appendix 11), there remain two species that 

cannot easily be included in the genus but neither can they be 

readily assigned to other genera. These are described and 

discussed below. 

Aneilema calandrinoides F. Mueller 

Aneilema calandrinoides F. Mueller, Frag.. 9:191. 1875. 

TYPE.-Australia, Queensland, between the Norman and 

Gilbert rivers, T A. Gulliver s.n. (MEL, holotype, not seen; 

BRI, K-2 sheets, isotypes). 

Decumbent herbs rooting at the nodes. Roots fibrous, 

moderately thick (-1-1.5 mm wide). Shoots unbranched (in 

specimens), up to 14 cm long. Internodes 2-7 cm long, 1-2 

mm in diameter, glabrous. Leaves wilh sheaths 5-14 mm long, 

glabrous or with some very small, scattered, uniseriate hairs, 

laminae sessile, apparently slightly fleshy, linear, 1.5-4 cm 

long (or perhaps longer because none has an apex), 1-3 mm 
148 SMITHSONAN CONTRIBUTIONS TO BOTANY 

m 

~ I 

a b C 

FIGURE 56.-Antherode and anthers of Aneilema calandrimides F. Mueller: a, 

antherode; b, lateral anther. E, medial anther (from Gul1ivers.n.-K), (Bar = 0.5 

mm.1 

wide, base cuneate, margin thickened, glabrous or with a few, 

minute, uniseriate hairs (and rarely 1 or more hook-hairs?) 

towards the base, both surfaces glabrous. 

Inflorescences consisting of solitary, terminal flowers, which 

become leaf-opposed. Bracts and bracteoles absent. Flowers 

apparently -2.5-3 cm wide. Pedicels erect, 11-22 mm long, 

glabrous. Sepals sepaloid, subequal, free, one (probably the 

outer) more strongly hooded than the other two, 8.5-9 mm 

long, glabrous. Petals free, equal or subequal, 2 cm long, not 

distinctly clawed, bluish purple. Staminodes 3, all on the side 

of the flower towards the strongly hooded sepal, subequal, 

filaments slender, -5 mm long, glabrous, antherodes bilobed, 

yellow, lobes sessile, reniform to transversely elliptic, -0.8 mm 

long, 1.4-1.6 mm wide (Figure 56u). Stamens 3, all on the side 

of the flower away from the strongly hooded sepal: lateral 

stamens with filaments -10 mm long, glabrous, anthers 

oblong-elliptic, 2-2.8 mm long, connective narrow, anther sacs 

with basal, free ends divaricate or not (Figure 56b); medial 

stamen with filament -6 mm long, glabrous, anther oblong, 

3-3.8 mm long, connective narrow, anther sacs with basal free 

ends slightly divaricate (Figure 56c). Ovary -1.5 mm long, 

3-angled and probably trilocular, densely covered with papillae 

or wart-like glands, particularly along the angles, tapering into 

the style, style S-shaped, -11 mm long, glabrous, stigma 

capitate. 

Capsules and seeds unknown. 

DISTRIBUTION.-Australia (N Queensland). 

SPECIMENS SEEN.-AUSTRAL,IA. QUEENSLAND: Between 

the Norman and Gilbert rivers, Gulliver s.n. (BRI, K-2 

sheets); Carpentaria, Canon’s Creek, Gulliver s.n. (K) [part of 

the previous collection?]. 

DISCUSSION 

The flowers of this species agree with those of Aneilemu in 

stamen and staminode arrangement, form of the staminodes, 

and differentiation of the medial from the lateral stamens. In 

some details, e.g., bilobed antherodes and differentiated medial 

stamen, they are closer to typical Aneilema species than are the 

flowers of A. neoculedonicum, a true Aneilemu. Indeed, the 

only observable floral difference from Aneilema is the papillae 

or wart-like glands on the ovary. It has not been possible to 

determine the numbers of locules and ovules because of the 

limited material. 

Although A. culundrinoides could readily be accepted as an 

Aneilema on the basis of its floral structure, its inflorescence 

structure-ebracteate and ebracteolate, one-flowered cincinni-is not only totally unlike that of any species of the 

genus, but it is also unique in the Commelinaceae. In Aneilemu 

the cincinni always have bracts at their bases, and the pedicels 

are bracteole-opposed. The cincinni are rarely one-flowered, 

and when they are, they occur in dense thyrses. 

Leaf-opposed inflorescences do not occur in Aneilema, 

although, as noted above, each individual flower is bracteoleopposed as in all Commelinaceae that possess bracteoles. On 

this basis the entire plant of A. calundrinoides might be 

considered equivalent to a single cincinnus of Aneilema, with 

the leaves homologous to the bracteoles and the internodes to 

the cincinnus axis. Because flowers appear to be borne opposed 

to every leaf in A. culundrinoides, this model has some appeal. 

The major drawbacks with it are (1) in order to be homologous 

to a cincinnus, the leaves and flowers in A. culundrinoides 

should be two-ranked, but, unfortunately, none of the shoots is 

long enough to determine this character, and (2) every 

internode of A. culundrinoides is subtended by a prophyll, 

which is characteristic of lateral shoots, but prophylls do not 

occur in cincinni. The cincinnus homology hypothesis therefore is probably incorrect, and it is likely that each flower 

represents a reduced cincinnus. 

I believe that, because of its Aneilemu floral structure, unique 

inflorescence, and unusual growth habit, A. culundrinoides 

must be treated either as a very aberrant species of Aneilema or 

as a separate genus. Although I once suggested the latter course 

(Faden, 1975), I now prefer to await more material before 

making a final decision. Whatever its generic position, A. 

culundrinoides clearly must have been derived from Aneilema 

section Aneilema. The decumbent habit and linear leaves of this 

species suggest a marsh habitat. None of the Australo-Oceanic 

Aneilema species occurs in such situations. Possibly the 

adaptation to an aquatic habitat by some ancestral Aneilema 

was accompanied by the radical changes in habit and 

inflorescence structure that produced A. culundrinoides. 

Aneilema brasiliense C.B. Clarke 

Aneilema brasiliense C.B. Clarke in De Candolle, Monogr. Phan., 3:225,1881. 

[Syntypes: Brazil, Piauhy [Piaui] Province, between Cacumbino and Retiro, 
NUMBER 76 149 

Mar 1829, Gurdner 2333 (K (ex Herbarium Hookerianum), lectotype; FHO, 

K (ex Herbarium Benthamianum), isolectotypes); Bahia, Blunchet 873 

(G-2 sheets, S).] 

Erect, unbranched or occasionally sparsely branched, annual 

herbs to -50 cm tall. Roots fibrous. Lower internodes elongate 

(to 10 cm long), uppermost greatly reduced so that the upper 

leaves are clustered below the inflorescence, all internodes 

puberulous with hook-hairs or the lower glabrescent. Leaves 

spirally arranged, sheaths 5-15 mm long, puberulous with 

hook-hairs, ciliate at the apex; laminae petiolate (except the 

uppermost), elliptic to lanceolate-elliptic, 6-15 cm long, 2-4.5 

cm wide, apex acuminate, base cuneate to broadly cuneate, 

margins planar, scabrid with prickle-hairs for their entire 

length, both surfaces sparsely to densely puberulous with 

hook-hairs, the adaxial scabrid with prickle hairs towards the 

margin, also occasionally with uniseriate hairs, midrib very 

prominent on the abaxial surface. 

Inflorescence a terminal (and frequently also axillary from 

the uppermost leaves), lax, obovoid (rarely ovoid) thyrse, 5-13 

cm long, 4-12 cm wide, with 1-5 widely spaced, alternate, 

erect to ascending (rarely patent) cincinni. Peduncles to -5 cm 

long, densely puberulous with hook-hairs. Inflorescence axis 

puberulous with hook-hairs. Cincinni to 9 cm long and 

10-flowered. Cincinnus bracts herbaceous or membranous, 

narrowly lanceolate, 3.5-lo(-20) mm long, longest in the 

lowermost cincinnus, decreasing in size upwards, subglabrous 

to puberulous with hook-hairs, generally with uniseriate hairs 

as well, particularly towards their apices, persistent. Cincinnus 

peduncles exceeding the cincinnus bracts, longest in the 

lowermost cincinnus, decreasing upwards, (1-)lS-3.5(-5) 

cm long, puberulous with hook-hairs. Cincinnus axes puberulous with hook-hairs, also with uniseriate hairs at the nodes. 

Bracteoles attached (0.8-)3-lo(-15) mm apart, herbaceous, 

lanceolate-ovate to ovate-elliptic, amplexicaul but not perfoliate, 2.5-3.5 mm long, sparsely pilose-puberulous with uniseriate hairs and hook-hairs, glandular(?) near the apex, persistent. 

Pedicels 5-7 mm long in flower, to 11 mm long in fruit, 

ascending in flower, erect in fruit, puberulous with hook-hairs. 

Sepals sepaloid, free, subequal, 4-7 mm long, sparsely 

pilose-puberulous with a mixture of hook-hairs and uniseriate 

hairs, glandular(?) near the apex; medial sepal lanceolate to 

lanceolate-elliptic, strongly hooded at the apex; lateral sepals 

narrowly elliptic to oblong, not strongly hooded. Paired petals 

-3-5 mm long (or longer?), apparently not to shortly clawed, 

limb ovate, blue to violet. Medial petal oblong, -3 mm long x 

0.6 mm wide, white(?). Filament bases free. Staminodes 

absent, or when present, subequal and opposite the outer sepal 

and paired petals, filaments very slender, glabrous, 1-2 mm 

long, antherodes unlobed to bilobed with sessile lobes, 

0.15-0.5 mm long, 0.2-0.7 mm wide (Figure 57). Lateral 

stamens opposite the medial petal and lateral sepals, with 

filaments k horizontal, straight, then sharply recurved near the 

apex, 2-6 mm long, glabrous, anthers ovate to ovate-elliptic, 

0.7-1.5 mm long, 0.6 mm wide. Medial stamen with filament 

il 

a b 

ROUM 57.-Antherodes of Aneilemu brusiliense C.B. Clarke: a, medial 

antherode; b, lateral antherode (from Gluziou 12253-P) (Bar = 0.5 mm.) 

arcuate-ascending, -3 mm long, glabrous, anther broadly 

elliptic, 0.4-1.3 mm long, 0.45-0.7 mm wide. Ovary sessile, 

obovate to oblong-elliptic, 1-1.2 mm long, glabrous, apex 

truncate, dorsal locule humpbacked towards the ovary apex, 

l-ovulate, ventral locules 2-owlate; style 2.5-6 mm long, 

gently arcuate-decurved for most of its length, then sharply 

recurved near the apex, stigma very large, capitate. 

Capsules sessile, oblong-elliptic to obovate-elliptic, dehiscent, bivalved, trilocular, 5.5-6.5 mm long, 3-3.7 mm wide, 

tan, dull, glabrous, apex emarginate, valves slightly spreading, 

persistent, dorsal locule usually (always?) empty, ventral 

locules 2-seeded, cells of the capsule wall isodiametric, 

arranged in longitudinal ranks. Seeds C rectangular, & truncate 

at both ends, -1.7 mm long x 2 mm wide, hilum linear, 

prominent, raised in a shallow groove, slightly extended onto 

apical and basal surfaces, embryotega lateral, testa rugose, tan, 

spotted with dark brown, especially on the bumps, with a very 

low, fine, irregular, colorless reticulum on the surface. 

DIsTRIBunON.-Venezue~a and Brazil. 

SPECIMENSEEN.-VENEZUELA. BOLIVAR: San Felix, 

Aristeguieta 5807 (K, NY, VEN); La Camilera, 40 km E of El 

Manteco, Delascio & Liesner 6989 (US) & 7031 (US); Ias 

Aceites, NE of El Manteco, Delascio & Liesner 7413 (MO). 

MONAGAS: Jusepin-Rio Amana, Aristeguieta & Virrueta 7543 

BRAZIL. BAHIA: Bahia, Blunchet 873 (G, S) & I263 (G). 

CEARA: Ceara, coastal region, Bollard 5 (K); Serra de Araripe, 

Novo Eu, yon Lutzelburg 26129 (M). MINAS GERMS: 15 km W 

of Januhia on road to Serra da Araras, Anderson 9256) (F, US). 

Lagoa Santa, Warming s.n. Cp). PERNAMBUCO: Tapera, Pickel 

3062 (US). PIAUI: Between Cacumbino and Retiro, Gardner 

2333 (FHO, K). RIO DE JANEIRO: Environs de Rio [de] Janeiro, 

Larangeiras, chez les Princes, Glariou 12253 (B, K, P). 

WN). 

DISCUSSION 

The most outstanding characters of A. brasiliense are its 

annual habit, very lax inflorescences with few cincinni, 

non-perfoliate bracteoles, sepals covered with a mixture of 

hook-hairs and uniseriate hairs, oblong medial petal, lack of 
150 SMITHSOMAN CONTRIBUTIONS TO BOTANY 

staminodes or, when present, their antherodes sometimes not 

bilobed, horizontal lateral stamens, glabrous ovary and capsule, 

and isodiametric cells of the capsule wall. The combination of 

these characters and the exclusive South American distribution 

of A. brasiliense readily separates it from all other species of 

Aneilema. The question remains as to whether A. brasiliense is 

sufficiently similar to other Aneilema species to merit retention 

in the genus. 

Aneilema brasiliense occupies the same position relative to 

the rest of Aneilema as does the genus Rhopalephora: it has 

very few characters that are otherwise unknown in Aneilema, 

e.g., oblong medial petal and complete lack of staminodes, but 

it combines a number of unusual ones so that associating A. 

brasiliense with any section of Aneilema is difficult. 

Overall, Aneilema brasiliense appears to be closest to 

species in section Aneilema or section Amelina. From the 

former it differs in its annual habit, bracteole form and 

pubescence, sepal pubescence type, reduced medial petal, 

shape of the cells of the capsule wall, and much longer hilum. 

Aneilema brasiliense differs from section Amelina, itself a 

rather heterogeneous group of species, in habit (only A. 

ephemerum of section Amelina is annual), cincinnus arrangement (A. longirrhizum has mostly alternate cincinni), bracteole 

form and pubescence (bracteoles not perfoliate in A. gillettii), 

medial petal and antherode form, and glabrous ovary and 

capsule (capsule glabrous in A. johnstonii, nearly glabrous in A. 

gillettii). 

Besides A. brasiliense, the only neotmpical Aneilema is A. 

umbrosum subsp. ovato-oblongum, a West African taxon that 

presumably has been introduced into the neotropics (Faden, 

1978~). It belongs to section Brevibarbata and is clearly 

unrelated to A. brasiliense. 

I believe that there are only two reasonable alternatives for 

the taxonomic treatment of A. brasiliense: (1) inclusion in 

Aneilema as a distinct section, or (2) exclusion and treatment as 

a separate, monotypic genus as was suggested by Clarke 

(1881a.b). A final decision must await more conclusive 

evidence obtainable only from living material. 
Appendix I 

Rejected Generic Synonyms of Aneilema 

The genus Aneilemu has been used in a much broader sense in the past than it is here. As 

a result, a number of generic names, formerly treated as synonyms of it, are now 

considered valid genera themselves or synonyms of other genera. These rejected generic 

synonyms of Aneilemu, along with their present status, are as follows: 

Aphylax Salisbury, Trans. Hort. Soc. London, 1:271, 1812 [nom. nud. = Murdannia 

Dichaespermum Wight [“Dichoespermum”], Icon. PI. Ind. Orient., 6:31, 1853 

Dicfyospermum Wight, Icon. PI. Ind. Orient., 6:29, 1853 [= distinct genus]. 

Dilasia Rafhesque, Fl. Tellur., 4:122, 1838 [= Murdannia]. 

Difelesia Rafinesque, F1. Rllur., 3:69, 1837 [= Murdannia]. 

Murdannia Royle, Illus. Bot. Himalayan ME., 403, tab. 95, fig. 3, 1840 [nom. cons. = 

Pilefocarpus Hasskarl, Flora, 49:212, 1866 [in clavi]; Commel. Ind., 14, 1870 

Prionosfachys Hasskarl, Flora, 49:212, 1866 [in clavil [= Murdannia]. 

Rhopalephora Hasskarl, Bot. Zeitung (Berlin), 186458, 1864 [= distinct genus]. 

Strepfylis Rafinesque, Fl. Rllur., 4:122, 1838 [= Murdannia]. 

Talipula Rafinesque, F1. Tcllur., 2:17, 1837 [= Murdannia]. 

Royle] . 

[= Murdannia]. 

distinct genus]. 

[= Rhopalephora Hasskarl]. 

151 
Appendix I1 

Index to Specific Epithets in Aneilema 

Epithets referable to a variety of genera have been included in Aneilema at one time or 

another. Their present status, when known, is given below. Correct names for species in 

Aneilema are in roman type. Specific names used in the text above but not yet published 

are listed as “Faden, nom. nud.” 

acuminatum R. Brown 

acutifolium Lauterbach & K. Schumann = Murdannia acutifolia (Lauterbach & K. Schumann) Faden, new combination 

(Basionym: Aneilema acutifolia Lauterbach & K. Schumann in Schumann and Lauterbach, Fl. deutsch. Schutzgebiete Sudsee, 217, 1900) 

adhaerans Kunth = A. aequinoctiale 

aequinoctiale (P. de Beauvois) G. Don 

aflne R. Brown = Murdannia graminea (R. Brown) Bruckner 

@ne De Wildeman, nom. illeg. (non A. affine R. Brown) = A. 

africanum P. de Beauvois = Floscopa africana (P. de 

aguense (Standley & Steyermark) Standley & Steyermark = 

allardii De Wildeman = Murdannia allardii (De Wildeman) 

ambiguum (P. de Beauvois) G. Don = Palisota ambigua (P. de 

angolense C.B. Clarke 

angustifolium N.E. Brown = Murdannia 

angustifolium De Wildeman, nom. illeg. (non A. angustifolium 

N.E. Brown) = A. macrorrhizum 

anthericoides R. Brown = Murdannia graminea (R. Brown) 

Bruckner 

aparine Perrier de la Bathie 

arenicola Faden 

asperum Buchanan-Hamilton ex Wallich, nom. nud. = Floarureum Merrill= Murdannia spectabilis (Kurz) Faden 

silvaticum 

Beauvois) C.B. Clarke 

Gibasis trij7ora (Martens & Galeotti) D.R. Hunt? 

Brenan 

Beauvois) C.B. Clarke 

scopa scandens Loureiro 

benadirense Chiovenda 

beniniense (P. de Beauvois) Kunth 

biflorum R. Brown 

blumei (Hasskarl) Bakhuizen van den Brink f. = Murdannia 

bodinieri Leveille & Vaniot = Murdannia hookeri (C.B. 

bracteatum (C.B. Clarke) Kuntze = Murdannia bracteata (C.B. 

blumei (Hasskarl) Brenan 

Clarke) Briickner 

Clarke) J.K. Morton ex Hong 

bracteolatum Martius = A. umbrosum 

brasiliense C.B. Clarke, generic status uncertain (see p. 150) 

brenanianum Faden 

brunneospermum Faden 

buaricum Mildbraed ex J.K. Morton, nom. nud. pro syn. = A. 

lanceolatum 

calandrinoides F. Mueller, generic status uncertain (see p. 148) 

calceolus Brenan 

canaliculatum Dalzell = Murdannia spirata (L.) Bruckner 

cavaleriei Leveille & Vaniot = Murdannia simplex (Vahl) 

chihuahuense (Standley) Woodson = Gibasis chihuahuensis 

chrysanthum K. Schumann = A. petersii 

chrysopogon Brenan 

clandestinum Ridley = Murdannia clandestina (Ridley) Faden, 

new combination (Basionym: Aneilema clandestinum Ridley in J. Roy. Asian Soc.Straits, 57:108, 1911) 

Brenan, new synonymy 

(Standley) Rohweder 

clarkei Rendle 

collinum (Brandegee) Matuda = Gibasis karwinskyana (J.A. & 

compressum Dalzell = Murdannia nudiflora (L.) Brenan 

conspicuum (Blume) Kunth = Dictyospermum conspicuum 

cordatum Buchanan-Hamilton ex Wallich, nom. nud. = 

coreanum Leveille & Vaniot = Murdannia keisak (Hasskarl) 

crispatum R. Brown = Pollia crispata (R. Brown) Bentham 

croceum Griffith = Murdannia crocea (Griffith) Faden 

cymosum (Blume) Kunth = Floscopa scandens Loureiro 

J.H. Schultes) Rohweder 

(Blume) Hasskarl 

Murdannia 

Handel-Mazzetti 

debile Wallich, nom. nud. = Murdannia nudiflora (L.) Brenan 

delicatulum C.B. Clarke ex Jex-Blake, nom. subnud. = 

densiforum (Blume) Kunth = Floscopa scandens bureiro 

densum Th. Fries, jr. = A. welwitschii 

diandrum Buchanan-Hamilton ex Wallich, nom. nud. = 

Murdannia nudifora (L.) Brenan 

Murdannia clarkeana Brenan 

152 
NUMBER 76 153 

didymum Buchanan-Hamilton ex Wallich, nom. nud. = Pollia 

didymum Seemann, nom. nud. = Pollia 

dimorphum Dalzell = Murdannia dimorpha (Dalzell) Bruckner 

discretum Craib = Murdannia 

dispermum Brenan 

divergens C.B. Clarke = Murdannia divergens (C.B. Clarke) 

diversifolium Hasskarl = Murdannia nudiflora (L.) Brenan 

dregeanum Kunth 

Briickner 

ecuadoriense Steyermark = Dichorisandra 

ehrenbergii (Hasskarl) C.B. Clarke = A. forskalii 

elatum (Vahl) Kunth = Murdannia japonica (Thunberg) Faden 

elatum (Vahl) Dalzell = Murdannia japonica (Thunberg) 

ensifolium Wight = Murdannia gigantea (Vahl) Bruckner 

ephemerum Faden, nom. nud. 

erectum De Wildeman = A. welwitschii 

esculentum Wallich ex C.B. Clarke = Murdannia esculenta 

(Wallich ex C.B.Clarke) Raizada ex Rolla Rao & Kammathy 

Faden 

fasciatum Warburg ex Lauterbach = Murdannia 

filiforme Buchanan-Hamilton ex Wallich, nom. nud. = Murfilipes Martius = Gibasis geniculata (Jacquin) Rohweder 

florentii De Wildeman = A. welwitschii 

floribundum (Kunth) Hooker & Arnott = Tripogandra multifoliosum Hasskarl = Murdannia nudiflora (L.) Brenan? 

formosanum N.E. Brown = Murdannia edulis (Stokes) Faden 

forskalii Kunth 

dunnia vaginata (L.) Briickner 

flora (Swartz) Rafinesque 

gardneri Seubert = Murdannia gardneri (Seubert) Bruckner 

geniculatum (Jacquin) Woodson = Gibasis geniculata (Jac - 

quin) Rohweder 

giganteum (Vahl) R. Brown = Murdannia gigantea (Vahl) 

Bruckner 

gillettii Brenan 

glanduliferum Joseph & Rolla Rao = Tricarpelema glanduliferum (Joseph & Rolla Rao) Faden, new combination 

(Basionym: Aneilema glanduliferum Joseph & Rolla Rao, J. 

Ind. Bot. Soc., 47:367, 1968) 

glaucum Thwaites ex C.B. Clarke = Murdannia glauca 

(Thwaites ex C.B. Clarke) Briickner 

gracile (Kotschy & Peyritsch) C.B. Clarke = A. lanceolatum 

gracile (Kunth) Steyermark, nom. illeg. (non A. gracile 

(Kotschy & Peyritsch) C.B. Clarke) = Callisia gracilis 

(Kunth) D.R. Hunt 

gramineum R. Brown = Murdannia graminea (R. Brown) 

Bruckner 

grandibracteolatum Faden 

greenmanii Woodson = Thyrsanthemum macrophyllum 

(Greenman) Rohweder 

hallbergii Blatter = Dictyospermum ovalifolium Wight 

hamiltonianum Wallich ex C.B. Clarke = Murdannia blumei 

(Hasskarl) Brenan 

herbaceum (Roxburgh) Wallich = Murdannia japonica (Thunberg) Faden 

heterophyllum (Brandegee) Matuda = Gibasis venustula 

(Kunth) D.R. Hunt subsp. peninsulae D.R. Hunt 

hirticeps C.B. Clarke ex J.K. Morton, nom. nud. pro syn. = A. 

welwitschii 

hirtum A. Richard 

hispidum D. Don = Floscopa scandens Loureiro 

hockii De Wildeman 

holosericeum (Kunth) Woodson = Thyrsanthemum floribun - 

homblei De Wildeman 

hookeri C.B. Clarke = Murdannia hookeri (C.B. Clarke) 

Bruckner 

humile Warburg = Dictyospermum humile (Warburg) J.K. 

Morton 

humile Memll, nom. illeg. (non A. hum’le Warburg) = 

Tricarpelema philippense (Panigrahi) Faden, new combination (Basionym: Dictyospermum philippense Panigrahi, 

Phytologia, 29: 338, 1975) 

dum (Martens & Galeotti) Pichon 

imberbe Ridley = Dictyospermum conspicuum (Blume) Hassimbricatum Warburg = Pollia 

indehiscens Faden 

irregularibracteatum Kew Bulletin, sphalm. = Senecio irregukarl 

laribracteatum De Wildeman 

japonicum (Thunberg) Kunth = Murdannia japonica (Thunjaponicum Maximowicz = Pollia secundifora (Blume) Bakjohnstonii K. Schumann 

junghuhnianum Miquel = Murdannia nudifora (L.) Brenan 

berg) Faden 

huizen van den Brink f.? 

kainanense Masamune = Murdannia kainanensis (Masamune) 

karwinskyanum (J.A. & J.H. Schultes) Woodson = Gibasis 

katangense De Wildeman = A. welwitschii 

keisak Hasskarl = Murdannia keisak (Hasskarl) Handelkeyense Warburg = Rhopalephora vitiensis (Seemann) Faden 

koenigii Wallich ex C.B. Clarke = Murdannia lanceolata 

(Wight) Kammathy 

kuntzei C.B. Clarke ex Kuntze, nom. nud. pro syn. = 

Murdannia bracteata (C.B. Clarke) J.K. Morton ex Hong 

Hong 

kanvinskyana (J.A. & J.H. Schultes) Rohweder 

Mazzetti 

lamuense Faden 

lanceolatum Bentham 

lancifolium Griffith = Murdannia nudiflora (L.) Brenan 
154 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

lanufolium Griffith, sphalm. = praec. 

lanuginosum Wallich ex C.B. Clarke = Murdannia lanuginosa 

latifolium Wight = Murdannia japonica (Thunberg) Faden 

laxiflorum Bentham ex C.B. Clarke, nom. nud. pro syn. = A. 

laxum R. Brown = A. acuminatum 

leiocaule K. Schumann 

leptospermum K. Schumann ex Faden, nom. nud. pro syn. = A. 

lineare (Bentham) Woodson = Gibasis linearis (Bentham) 

lineolatum (Blume) Kunth = Murdannia juponica (Thunberg) 

longicapsa Faden 

longifolium Hooker = Murdannia simplex (Vahl) Brenan 

longifolium Wallich, nom. nud. = Murdannia 

longirrhizum Faden 

loriforme Hasskarl = Murdannia loriformis (Hasskarl) Rolla 

loureirii Hance = Murdannia edulis (Stokes) Faden 

loureiroi Index Kewensis = praec. 

lujai De Wildeman & Th. Durand = A. lanceolatum 

(Wallich ex C. B. Clarke) Bruckner 

umbrosum 

petersii 

Rohweder 

Faden 

Rao & Kammathy 

macrophyllum R. Brown = Pollia macrophylla (R. Brown) 

macrorrhizum Th. Fries, jr. 

malabaricum (L.) Merrill= Murdannia nudij7ora (L.) Brenan 

medicum (Loureiro) Kostelesky = Murdannia medica (Loumelanostictum Hance = Murdannia spirata (L.) Bruckner 

micranthum (Vahl) Kunth = Rhopalephora micrantha (Vahl) 

minutiflorum Faden, nom. nud. 

minutum (Blume) Kunth = Murdannia nudij7ora (L.) Brenan? 

monadelphum (Blume) Kunth = Rhopalephora micrantha 

montanum (Wight) Wallich ex C.B. Clarke = Dictyospermum 

mortehanii De Wildeman = A. bcniniense 

mortonii Brenan 

multiscaposum Lauterbach = Murdannia edulis (Stokes) Faden 

Bentham 

reiro) Hong 

Faden 

(Vahl) Faden 

montanum Wight 

nanum (Roxburgh) Kunth = Murdannia spirata (L.) Bruckner 

neocaledonicum Schlechter 

nicholsonii C.B. Clarke 

nigritianum (C.B. Clarke) Hutchinson ex Hutchinson & Dalziel 

nudicaule (N.L. Burmann) G. Don = Murdannia nudiflora (L.) 

nudiflorum (L.) Sweet = Murdannia nudij7ora (L.) Brenan 

nummularium Miquel = Murdannia spirata (L.) Bruckner 

nutans Leveille = Murdannia triquetra (Wallich ex C.B. 

Clarke) Bruickner 

nyasense C.B. Clarke 

= A. umbrosum subsp. umbrosum 

Brenan 

obbiadense Chiovenda 

ochraceum Dalzell = Murdannia crocea (Griffith) Faden 

octospermum C.B. Clarke ex Brenan, nom. nud. pro syn. = A. 

oliganthum Franchet & Savatier = Murdannia keisak (Hassovalifolium (Wight) Hooker f. ex C.B. Clarke = Dictyosperovato-oblongum P. de Beauvois = A. umbrosum subsp. 

ovatum (Hasskarl) Wallich ex C.B. Clarke = Dictyospermum 

subsp. ochracea (Dalzell) Faden 

rendlei 

karl) Handel-Mazzetti 

mum ovalifolium Wight 

ovato-oblongum (I? de Beauvois) J.K. Morton 

ovatum Hasskarl 

paludosum A. Chevalier 

paniculatum Wight = Murdannia dimorpha (Dalzell) Briickner 

paniculatum Wight ex C.B. Clarke, nom. illeg. (non A. 

paniculatum Wight) = Murdannia semiteres (Dalzell) 

Santapau 

papuanum Warburg = A. acuminatum 

paraguayense C.B. Clarke ex Chodat = Murdannia paraguaypauciforum Dalzell = Murdannia vaginata (L.) Bruckner 

pauciforum Wight, nom. illeg. (non A. paucij'lorum Dalzell) = 

paucij'lorum (Urban & Ekman) Alain, nom. illeg. (non A. 

paucifolium N.E. Brown = Murdannia medica (Loureiro) Hong 

pedunculosum C.B. Clarke 

petersii (Hasskarl) C.B. Clarke 

pilosum Wallich, nom. nud. = Murdannia crocea (Griffith) 

pinetorum (Greene) Matuda = Tradescantia pinetorum Greene 

plagiocapsa K. Schumann 

platyphyllum Menill= Murdannia edulis (Stokes) Faden 

poaeoides Seubert = Callisia filiformis (Martens & Galeotti) 

pomeridianum Stanfield & Brenan 

protensum (Wight) Wallich ex C.B. Clarke = Rhopalephora 

scaberrima (Blume) Faden 

puberulum Hasskarl, sphalm. = A. lanceolatum 

pulchellum (Kunth) Woodson = Gibasis pulchella (Kunth) 

pulneyense Fyson = Murdannia esculenta (Wallich ex C.B. 

pusillum Chiovenda 

ensis (C.B. Clarke ex Chodat) Bruckner 

Murdannia paucij'lora Bruckner 

pauciforum Dalzell) = Gibasis 

Faden 

D.R. Hunt 

Rafinesque 

Clarke) Raizada ex Rao Rolla & Kammathy? 

radicans D. Don = Murdannia nudij7ora (L.) Brenan 

recurvatum Faden 

rendlei C.B. Clarke 

reniforme Buchanan-Hamilton ex Wallich, nom. nud. = Pollia 

rhodospermum K. Schumann ex C.B. Clarke, nom. nud. pro 

richardsiae Brenan 

rigidum Blatter = Murdannia simplex (Vahl) Brenan 

syn. = A. silvaticum 
NUMBER 76 155 

ringoetii De Wildeman = A. hirtum 

rivulare A. Richard = Floscopa glomerata (Willdenow ex J.A. 

rugosum Pemer de la Bithie = Rhopalephora rugosa (Femer 

russeggeri (Fenzl) C.B. Clarke (“russegeri” sphalm.) = A. 

& J.H. Schultes) Hasskarl 

de la Bathie) Faden 

lanceolatum 

sacleuxii Hua = A. petersii 

scaberrimum (Blume) Kunth = Rhopalephora scaberrima 

scapijlorum (Roxburgh) Kostelesky = Murdannia edulis 

schlechteri K. Schumann 

schomburgkianum Kunth = Murdannia schomburgkiana 

(Kunth) Bruckner 

schweinfurthii Baker ex C.B. Clarke = A. lanceolatum 

sclerocarpum F. Mueller 

sebitense Faden 

secundijlorum (Blume) Kunth = Pollia secundiflora (Blume) 

secundum Wight = Murdannia simplex (Vahl) Brenan 

semifoliatum C.B. Clarke = Murdannia semifoliata (C.B. 

semiteres Dalzell = Murdannia semiteres (Dalzell) Santapau 

sepalosum C.B. Clarke = Anthericopsis sepalosa (C.B. Clarke) 

serotinum D. Don ex C.B. Clarke, nom. nud. pro syn. = 

serrulatum (Vahl) G. Don = Tripogandra serrulata (Vahl) 

sessilifolium (Bentham) J.K. Morton, pro syn. = A. beniniense 

setiferum A. Chevalier 

siamense Craib = Pollia secundifora (Blume) Bakhuizen van 

den Brink f. 

sienneum Blatter = Murdannia lanuginosa (Wallich ex C.B. 

Clarke) Briickner 

siliculosum R. Brown 

silvaticum Brenan 

simplex (Vahl) Kunth = Murdannia simplex (Vahl) Brenan 

sinicum Ker-Gawler = praec. 

smithii C.B. Clarke = A. somaliense 

somaliense C.B. Clarke 

soudanicum C.B. Clarke = A. lanceolatum 

spectabile Kurz = Murdannia spectabilis (Kurz) Faden 

spekei C.B. Clarke 

spicatum Wallich, nom. nud. = Murdannia spectabilis (Kurz) 

spiratum (I.,.) Sweet = Murdannia spirata (L.) Briickner 

stenothyrsum Diels = Murdannia stenothyrsa (Diels) Handel- 

(Blume) Faden 

(Stokes) Faden 

Bakhuizen van den Brink f. 

Clarke) Bruckner 

Engler 

Murdannia edulis (Stokes) Faden 

Handlos 

subsp. sessifolium (Bentham) J.K. Morton 

Faden 

Mazzetti 

stolrii Mildbraed ex J.K. Morton, nom. nud. pro syn. = A. 

subnudum A. Chevalier = A. lanceolatum subsp. subnudum (A. 

subovatum Ridley = Dictyospermum ovatum Hasskarl 

succulentum Faden 

hirtum 

Chevalier) J.K. Morton 

tacazzeanum Hochstetter ex A. Richard = A. forskalii 

tacarreanum Hochstetter ex C.B. Clarke = A. forskalii 

tanaense Faden 

taquetii Leveille = Murdannia keisak (Hasskarl) HandelMazzetti 

taylorii C.B. Clarke 

tenerum Baker = Pseudoparis tenera (Baker) Faden, new 

combination (Basionym: Aneilema tenerum Baker in J. 

Linn. Soc., Bot., 22530, 1887) 

tenuissimum (A. Chevalier) A. Chevalier ex Hutchinson & 

Dalziel = Murdannia tenuissima (A. Chevalier) Brenan 

terminale Wight = Murdannia loriformis (Hasskarl) Rolla Rao 

& Kammathy 

terminale (Blume) Boldingh ex Haines, nom. illeg. (non A. 

terminale Wight), probably = Cyanotis moluccana (Roxburgh) Merrill 

termitarium Faden, nom. nud. 

tetraspermum K. Schumann = A. petersii 

thomsonii (C.B. Clarke) C.B. Clarke = Tricarpelema gigantrichocoleum Schauer = Murdannia nudijlora (L.) Brenan 

triquetrum Wallich ex C.B. Clarke = Murdannia triquetra 

tuberosum Buchanan-Hamilton ex Wallich, nom. nud. = 

teum (Hasskarl) Hara 

(Wallich ex C.B. Clarke) Briickner 

Murdannia edulis (Stokes) Faden 

umbrosum (Vahl) Kunth 

urbinanum (Greenman) Matuda = Gibasis 

usambarense Faden 

vaginatum (L.) Wallich = Murdannia vaginata (L.) Bruckner 

vankerckhovenii De Wildeman = A. nyasense 

versicolor Dalzell = Murdannia versicolor (Dalzell) Bruckner 

violaceum F. Mueller ex C.B. Clarke, nom. nud. pro syn. = 

vitiense Seemann = Rhopalephora vitiensis (Seemann) Faden 

Murdannia graminea (R. Brown) Bruckner 

welwitschii C.B. Clarke 

whytei C.B. Clarke = A. hirtum 

wildii Merxmiiller = A. hockii 

woodii Faden 

zebrinurn Chiovenda 

reylanicum C.B. Clarke = Murdannia zeylanica (C.B. Clarke) 

Bruckner 
Appendix I11 

Specimens Cited 

All collections of Aneilema taxa belonging to sections Rendlei, Somaliensia, and 

Lamprodithyros, but none belonging to other sections, are listed. The first number in 

parentheses refers to the section, the second to the species within the section. A letter 

following the species number refers to a subspecies or hybrid (when applicable). A full list 

of the taxa and their corresponding numbers is given in the table of contents. 

Adamson, J. 42 (4-1); 599 (4-1). 

Allen 94 (5-la). 

Ament & Magogo 263 (5-lb). 

Andrews A3589 (5-5); in EA15070 (5-8); s.n. (5-la). 

Archbold 713 (3-3); 1408 (3-1); 1491 @-la); 1897 (3-3). 

Aylmer 182 (5-5). 

Ash 823 (4-1). 

Baldrati 933 (5-5); 1387 (5-5); 1388 (5-5). 

Bally B8534 (3-3); B10133 (4-1); B12145 (5-13); B12193 

(5-13); B12348 (5-13); B12692 (3-3); B13592 (3-3); 

B13931 (5-13). 

Bally & Smith B14320 (5-13); B14357 (5-la). 

Barbosa 655 (5-2b). 

Barbosa & Lemos 843 1 (5-2b). 

Batty 162 (5-la). 

Bax 368 (5-lb). 

Bayliss BS7216 (5-2b). 

Beckett 1022 (4-5c). 

Birnie 9 (5-2c). 

Boaler 626 (5-la). 

Boivin s.n. (5-la). 

Borle 364 (5-2b). 

Botany Students in DSM1351 (5-9). 

Botta s.n. (5-5). 

Braun 1377 (5-la); 3624 @-la). 

Bredenkamp 1795 (5-2b). 

Brenan 14232 (5-2b); 14243 (5-13). 

Buchanan s.n. (3-3). 

Busse 2281 (5-la); 3054 (5-la). 

Carter & Stannard 652 (4-1); 692 (4-1). 

Carvalho s.n. (5-la). 

Ciferri 73 (5-7). 

Compton 28610 (5-2b). 

Corradi 2154 (5-13); 2155 (5-13); 2159 (3-3); 2160 (3-3); 

Cufodontis 319 (4-1); 705 (5-la). 

Codd 689 1 (5-2b). 

2163 (5-13); 2196 (4-1). 

Davey D79 (4-1). 

Deflers 968 (5-5). 

de Wade & de Wilde Duyfjes 10529 (5-5). 

Donaldson Smith 346 (5-4); s.n. (3-3); s.n. (4-1); s.n. (4-1). 

Drummond & Hemsley 2337 (5-2a); 2928 (3-3); 3561 (5-9); 

Dummer 5020 (3-3). 

DEW 114 (5-5); 287 (5-5); 300B (5-5). 

3610 @-la). 

Ehrenberg 413 (5-5). 

Engler 3416 (3-1). 

Evans & Maikweki 51 (5-10); 54 (5-12). 

Everard in EA 11456 (4-1). 

Faden 69/1300 (5-3); 69/1300A (5-2c); 69/1608 (5-3); 

69/1642 (5-3); 69/1699 (5-3); 69/1802 (5-3); 69/1933 

(5-3). 

Faden & Beentje 85/69 (5-10). 

Faden & Evans 701782 (5-12); 70/788 (3-3); 70/819 @-la); 

71/498 (3-4); 71/513 (3-4); 71/620 (5-12); 71/714 

(5-10); 711726 (5-10). 

Faden & Faden 71/788 (5-10); 71/809 (5-10); 72/30 (5-12); 

72/72 (5-2a); 72/88 (5-la); 72/91 (5-12); 72/92 (5-13); 

72/166 (3-4); 72/226A (5-10); 72/231 (5-lb); 72/231A 

(5-lb); 72/240 (3-3); 72/284 (5-lb); 74/202 (5-2b); 

74/208 (5-2b); 74/232 (3-3); 74/237 (5-lb); 74/26 

(5-lb); 74/274 (5-lb); 74/281 (3-3); 74/291 (5-la); 

74/308 (5-la); 74/324 (5-9); 74/333 (5-9); 74/371 (3-1); 

74/380 (5-2a); 74/482 (5-13); 74/489 (5-2a); 74/503 

(3-3); 74/524 (5-12); 74/529 (5-lb); 74/532 (5-2a); 

74/533 (5-13); 74/536 (5-2a); 741751 (5-lb); 74/753 

(5-lb); 74/923 (4-1); 74/939 (4-1); 74/951 (4-1); 

74/1052 (5-13); 74/1053 (5-8); 74/1064 (5-8); 74/1066 

(5-2a); 74/1069 (5-9); 74/1083 (5-1 1); 74/1142 (5-la); 

74/1152 (5-12); 74/1171 (5-8); 74/1173 (5-2a); 74/1176 

(5-13); 74/1179 (5-8); 74/1183 (5-la); 74/1184 (5-2a); 

(5-la); 74/1281 (5-12); 74/1283 (5-13); 74/1284 (5-2a); 

74/1185 (5-8); 74/1186 (5-12); 74/1215 (5-10); 74/1237 

74/1285 (3-3); 77/310 (5-lb); 77/311 (5-13); 77/342 

156 
NUMBER 76 157 

(5-lb); 77/364 (5-13); 77/378 (5-9); 77/484 (5-10); 

77/565 (5-9); 77/574 (5-12); 77/582 (5-8); 77/585 

(5-13); 77/611 (3-1); 77/629 (5-10); 77/738 (5-8); 

77/749 (5-10); 77/775 (5-10); 77/785 (5-3); 77/786 

(5-3); 77/788 (5-2~); 771803 (5-4); 77/803A (5-4). 

Faden & Napper 69/238 (3-4); 69/29 (3-4). 

Faden et al. 69/318 (5-2a); 69/322 (3-3); 69/395 (5-lb); 

69/403B (3-3); 69/453 (5-la); 69/617 (5-lb); 69/1066 

70/158 (5-2a); 70/183 (5-13); 70/686A (5-la); 70/888 

(5-3); 69/1068 (5-lb); 69/2069 (3-4); 70/157 (3-3); 

(5-13); 70/894 (5-3); 70/894A (5-2~); 70/937 (5-12); 

70/966 (5-13); 70/969 (5-12); 71/633 (5-10); 72/38 

(5-la); 72/39 (5-12); 72/227 (5-12); 72/228 (5-la); 

74/205 (5-13); 74/326 (5-la); 74/330 (5-13); 77/523 

(5-10). 

Faden, A. 3/79 (5-24; 16/85 (3-3); 27/85 (3-3). 

Faulkner 838 (5-la); 1160 (5-2a); 3670 (5-la); 4045 (5-9); 

4471 (5-2a); s.n. (5-la). 

Faulkner in K712 @-la). 

Fiori 874 (5-5); 875 (5-5). 

Forsskg in Herb. Forskalii 31 (5-5); 32 (5-5); 33 (5-5). 

Friis et al. 2670 (4-1). 

Gagal556 (4-1). 

Geilinger 133 (5-la); 155 (5-la); 642 (5-la); 4783 (3-3). 

Gilbert 5018 (4-1); s.n. (3-3). 

Gilbert & Gilbert 1617 (5-3); 2186 (5-5). 

Gilbert & Jefford 4559 (4-1). 

Gilbert & Rankin 4834 (5-8). 

Gilbert & Thulin 266 (5-4); 318 (5-3); 969 (5-5); 1452 

Gilbert et al. 5303 (4-1). 

Gillett 12651 (4-1); 12702 (4-1); 13132 (5-lb); 14032 (4-1); 

14091 (5-la); 17204 (5-lb); 17271 (5-lb); 18604 (5-lb); 

19528 (5-8); 19532 (5-8); 19562 (5-2a); 19973 (5-8). 

(4-1); 1619A (4-1); 1730 (5-lb); 4659 (4-5b). 

Gillett & Beckett 23289 (4-4). 

Gillett & Gachathi 20488 (5-lb). 

Gillett & Mathew 19068 (5-lb); 19073 (5-lb); 19090 (5-lb). 

Gillett et al. 22229 (4-3); 22399 (4-5b); 22597 (4-5a); 22609 

Glover & Gilliland 408 (4-2); 1031 (4-5c). 

Gomes & Sousa 3441 (5-2b); s.n. (5-2b). 

Graham 1848 (5-la); 1903 (5-la); 3019 (5-3). 

Greenway 1668 (3-1); 2193 (3-3); 5236 (5-la); 6058 (3-2). 

Greenway & Kanuri 11871 (5-lb). 

Greenway & Rawlins 9344 (5-10). 

Gregory s.n. (5-8); s.n. (5-10). 

Grenfell s.n. (3-3). 

(4-4). 

Haarer 447 (3-3); 1445 (3-3). 

Harris 1484 (5-la); 2045 (5-la); BJH2848 (5-la); BJH3069 

Harris & Procter 1018 (5-9). 

Harris & Tadros BJH5721 (5-la). 

(5-la); BJH3413 (5-la); BJH5266 (5-la). 

Harris et al. BJH4377 (3-3). 

Heriz Smith s.n. (3-4). 

Herlocker 757 (4-1). 

Hindorf 806 (5-3). 

Hucks 221 (5-lb); 669 (3-3); 669 (5-lb); 690 (5-lb). 

Hucks & Hucks 15 (5-lb). 

Jackson 2804 (5-5). 

James & Thrupp s.n. (4-1); s.n. (4-5b). 

Jeffrey K80 (5-la): K315 (5-10). 

Junod 493 (5-2b). 

Kabuye & Evans 163 (3-3). 

Kirk s.n. (5-2b). 

Kloetzli et al. 144 (5-la). 

Lavranos 11466 (5-5); 17063 (5-2c). 

Lawn 1289 (5-2b). 

Lswis 5999 (5-3). 

Lucas et al. 256 (5-la). 

Lye et al. 3981 (5-3). 

Lock 81/275 (5-lb). 

Mabberley & McCall88 (5-4); 88a (5-4). 

Maffey 51 (5-5). 

Magogo in EA14203 (3-1). 

Magogo & Glover 906 (5-la). 

Mandaville 6860 (5-5); 6886 (5-5). 

Mathew & Hanid 6061 (5-lb). 

McKeag [or McCraig] in CM9187 (5-la). 

Mgaza 781 (5-la). 

Mhoro & Wingfield BM2411 (5-13). 

Milne-Redhead & Taylor 7285 (5-9); 7437 (5-la). 

Mistry M-7 (5-5); M-11A (5-3); M-27 (5-3). 

Mogg 27469 (5-2b); 28886 (5-2b). 

Moll 4152 (5-2b). 

Moomaw 951 (5-la). 

Moura et al. 399 (5-2b). 

Msafiri 256 (5-lb); 307A (3-3). 

Napier 914 (3-3); 973 (5-2a). 

Napier & Fox 3431 (5-3). 

Napper & Kanuri 2087 (5-lb). 

Napper et al. 1999 (3-4). 

Newbould 2957 (4-1); 3184 (3-3). 

Oatley C6 (5-2b). 

Organ in EA14970 (5-9); in EA15079 (3-1). 

Paoli 1085 (4-1); 1277 (4-5a). 

Pappi 451 1 (5-5). 

Peters s.n. (5-la). 

Polhill & Paul0 629 (5-12); 796 (5-10). 

Pedro 109 (5-2b). 
158 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Poole 1255 (5-13). 

Pooley 1399 (5-2b). 

Puccioni & Stefanini 579 (4-4); 592 (4-3). 

Quartin Dillon 449 (5-5); s.n. (5-5); s.n. (5-5). 

Quartin Dillon & Petit 449 (5-5). 

Quintas 64 (5-2b). 

Rawlins 239 (5-la); 454 (5-12); 863 (5-12); in EA11280 

Richards 21988 (5-la). 

Robertson, D.C. in TRU781 (5-la). 

Robertson, S.A. 529 (5-la). 

Ruspoli & Riva 457 (4-5b). 

@-la). 

Saboureau 259 (5-5). 

Sacleux 1142 (5-la). 

Saunders in EA11211 (5-la). 

Schimper 1660 (5-5); 1729 (5-5). 

Schlieben 5942 (5-la); 12121 (5-11). 

Schweinfurth 289 (5-5); 289 (5-6); 944 (5-5); 944 (5-6); 

Schweinfurth & Riva 427 (5-5); 565 (5-5); 1672 (5-5). 

Scott Elliot 6179 (3-3). 

Spjut 3812 @-la). 

Strey 4752 (5-13); 10326 (5-2b). 

1416 (5-5). 

Shbetd F1507 (5-5). 

Seid 2406 (5-lb). 

Tackholm s. n. (5-5).

Tanner 401 1 (5-3). 

Taylor s.n. (3-1). 

Thomas 1061b (3-3). 

Thulin 5673 (4-5d). 

Thulin & Warfa 4570 (4-5d); 4614 (4-4); 4659 (4-5b); 5280 

(5-lb); 5320 (4-5d); 5341 (4-4); 5395A (4-5~); 5423 

(4 - 5c). 

Thulin et al. 3539 (4-1). 

Timberlake 156a (5-lb); 344 (5-lb). 

Torre & Correia 142% (5-la). 

Tweedie 1202 (5-12); 1624 (5-la); 3183 (5-la); 3364 

(5-12); 3365 (5-la); 4036 (3-3). 

Vahrmeijer 693 (5-2b). 

Vahrmeijer & Tolken 192 (5-2b). 

Vaughan 621 (5-10); 1851 ‘(5-2a); 2064 (5-10); 2106 (5-la); 

Verdcourt 161 (3-1); 1921 (3-1); 2410 (3-1); 2531 (3-4); 

Verdcourt & Polhill 2708 (5-lb). 

Verdcourt et al. 2667 (3-4). 

Vesey-FitzGerald 1236/8 (5-5). 

Volkens 175 (5-la). 

Vollesen 1964 (5-lb); 3350 @-la). 

2395 (5-10). 

3892 (5-13); 3901 (3-3); 5234 (5-lb). 

Wallace 810 (5-la). 

Warnecke 323 (3-1). 

Welch 243 (5-la). 

Whyte s.n. (5-la); s.n. (5-la); s.n. (5-la). 

Wingfield 2899 (3-3). 

Winkler 3638 (3-1). 

Wood 1892 (5-5); 1902 (5-5); 1997 (5-5); 2126 (5-6a); 2128 

(5-6a); 2173 (5-5); 2395 (5-5); 2580 (5-5); 2616A 

(5-6a); 26T6B (5-6); 2959 (5-5). 

Zimmerman 8231 (3-1). 
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Index 

This index includes all scientific names of Commelinaceae that are mentioned in the text and appendices. 

Synonyms and page number of the principal reference for a taxon that is described in the text are in italics. 

Numbers in brackets refer to page numbers of figures and maps. 

Amelk C.B. Clarke, 55,57,61,83 

wdlichii C.B. Clarke, 57,61 

Aneilema R. Brown 

subgenus Dicorpeltorio C.B. Clarke, 55,5758 

subgenus Tricorpellario C.B. Clarke, 58 

section Amelina (C.B. Clarke) C.B. Clarke, 1,2,7,8, 10,23, 30,35, 36, 

section Aneilema, 1, 2, 7, 8, 10, 22, 30, 35, 36, 43, 47-49, 59, 60, 150, 

section Brevibarbata Faden, v, 1.7, 8, 10, 23.28, 30.36, 43.47-50,54, 

section Dickespermwn (Wight) C.B. Clarke, 58 

section Dictyospermum (Wight) C.B. Clarke, 58 

section Euaneilemo C.B. Clarke, 58 

section Lamprodithyros (Hasskarl) C.B. Clarke, 1,2.7,8,20,22.23, (241, 

73,90,93, [931,99,100,110,117,121,125,128,130.133,142,143, 

section Pedunculosa Faden, v, 1,2.7,8, 10,12.13,28,30,31,36,38,43, 

section Pseudo-axillores C.B. Clarke, 58,90.92 

section Rendlei Faden, v, 1,2.7.8,23,28,30,34-36.43,47-51.54,58, 

section Somaliensia Faden, v, 1,2,7,8,28,30, 35,36,43.47-49,51,54, 

acuminatum R. Brown, 1, 2, 7, 9. 10, 16, 20, 26, [30], 33, 35, 36, 42, 

47-49,51.58,60. 113, 152. 154, Plate 1 

ocutifoliwn Lauterbach & K. Schumann, 152 

odhoerons Kunth. 152 

aequinwtiale (P. de Beauvois) G. Don, 1,2,7-10, 16, 17, [17], 23,26, 

38,43,47-50,54,58,59,61,62, 142, 143. 150, Plate 1 

Plate 1 

59.62,81.143,144-146.150, Plate 4 

L251.26, [271,28. [281, [291,30,35,36,38,42,43,47-51,54,58,59, 

Plates 2-4 

47-49.54,59,146, 147, Plate 4 

62, [63], 64, 81,82, Plates 1, 2 

59,77, [78], 82, 85, 86, 143, Plate 2 

30, [30]. 31,32,36, [38], 42,43, [44], [45], [46], 47-49,57,61.62, 

113. 116, 152, Plate 1 

p minor C.B. Clarke, 116 

offie R. Brown, 152 

offie De Wildanan, 152 

ofriconwn P. de Beauvois, 152 

oguense (Standley & Steyermark) Standley & Styennark, 152 

ollordii De Wildeman, 152 

mbiguwn (P. de Beauvois) G. Don, 152 

angolense C.B. Clarke, 7,23,144-146.152 

ongrcstifoliwn N.E. Brown, 152 

ongustifoIiwn De Wildeman. 152 

onthericoides R. Brown, 152 

aparine Perrier de la Bathie, 7, 10,53,60, 152 

arenicola Faden, 1.7-9, 16,36,50, 144-146, 152, Plate 7 

asperwn Buchanan-Hamilton ex Wallich, 152 

ozwewn Merrill, 152 

benadirense Chiovenda,2,7,22,23,91,93,110,122, [123], [124], 125, 

128,152, Plate 6 

beniniense (P. de Beauvois) Kunth, 1,7,9, 10, 16, 17.23. 31, 32. [33], 

42,43, [441, [451,47-49, 143,145, 146, 152, 154 

subsp. beniniense, 32,35, Plate 4 

subsp. sessifolium (Bentham) J.K. Morton, 155 

biflo- R. Brown, 1,7,9, 14.16,20,22.31, [MI. 47-49,55,58,60, 

152 

blwnei (Hasskarl) Bakhuizen van den Brink f., 152 

bodinieri Liveill6 & Vaniot, 152 

brocteotwn (C.B. Clarke) Kuntze, 152 

brocteolotwn Martius. 152 

brasiliense C.B. Clarke, 56,148, 149, [149], 150, 152 

brenanianum Faden, v, 1.7, 8, 10, 14, 16, 17,23, 36, 50, 64, [67], 73, 

brunneospermum Faden, 1,7-9, 16, 17,36, [45], 46,47-49,144-146, 

buaricum Mildbraed ex J.K. Morton, 152 

calandrinoides F. Mueller, 56,147, 148, [148]. 152 

calceolus Bman, 1,7-10,16,23,[241. [261. [281,32. [371,38. [40], 42. 

49.50,92,93, [124], 125,128, 130, 133, 143,152, Plates 3,6 

conolicrclotum Dalzell, 152 

covoleriei Leveille & Vaniot, 152 

chihuohuense (Standley) Woodson, 152 

chrysonthwn K. Schumann, 94,98, 152 

chrysopogon Brenan, 1,7,16,22, 147,152, Plate 4 

clondestinwn Ridley, 152 

clarkei Rendle, 1,7,8, 10, [lo], 11, 13, 16,20. [211,23, [25], [29], 32, 

[74]. 76,77,81, 85, 152, Plates 2.5 

152, Plates 4,7 

1371, [46], 47-49,90,92,93, 108, 125. 128,130, [132], 133, 136, 

138, 140, 143, 152, Plates 3,4,6 

collinwn (Brandegee) Matuda. 152 

compresswn DalzelI, 152 

conspicuum (Blume) Kunth, 152 

cordotum Buchanan-Hamilton ex Wallich, 152 

coreonum Leveille & Vaniot, 152 

crispotwn R. Brown, 55, 152 

crocewn Griffith, 152 

cymoswn (Blume) Kunth, 152 

debile Wallich, 152 

delicatulum C.B. Clarke ex Jex-Blake, 152 

densifIorwn (Blume) Kunth, 152 

densum Th. Fries jr.. 152 

diandrwn Buchanan-Hamilton ex Wallich, 152 

didymwn Buchanan-Hamilton ex Wallich. 153 

dkiymwn Seemann, 153 

dimorphwn Dalzell, 153 

dircretum Craib, 153 

dispermum Brenan, 1,7,16,42, 144-146,153 

divergens C.B. Clarke, 153 

dversifoliwn Hasskarl, 153 

dregeanumKunth, 1,7,16,36,42, 100,105, 145,146,153 

subsp. dregeanum, 17.31 

subsp. mossambicense Faden, 42 

ecdoriense Steyermark, 153 

ehrenbergii (Hasskarl) C.B. Clarke, 113, 153 

elotwn (Vahl) Kunth, 153 

elotwn (Vahl) Dalzell, 153 

162 
NUMBER 76 163 

ensifoliwn Wight, 153 

ephemerum Faden, 1,7.9,10,16,23,31,36, [44], [46], 47-49,62,150, 

erectwn De Wildeman, 153 

esculentwn Wallich ex C.B. Clarke, 153 

fasciatwn Warburg ex Lauterbach, 153 

filiform Buchanan-Hamilton ex Wallich, 153 

filipes Martius. 153 

florentii De Wildeman, 153 

floribundwn (Kunth) Hooker & Amott, 153 

folioswn Hasskarl, 153 

formosonwn N.E. Brown, 153 

forskalii Kunth, 1, 3,7, 12, 13, 16, [27], 36,38. 90.92, 93, 106. 110, 

gordneri Seubert. 153 

geniculotwn (Jaquin) Woodson, 153 

gigontewn (Vahl) R. Brown, 55, 153 

gillettii Brenan, 1,7, 10, 16, 17, 22,30,31,36,43, [45], 47-49,61, 83, 

glonduliferwn Joseph & Rolla Rao, 153 

gloucwn Thwaites ex C.B. Clarke, 153 

grocile (Kotschy & Peyritsch) C.B. Clarke, 153 

gracile (Kunth) Steyermark, 153 

grom'ncwn R. Brown, 55, 153 

grandibracteolatum Faden, v. 2, 7, 36, 38, 59. 77, 78, [go], 82, 83. 85. 

86, 153, Plate 5 

greenmonii Woodson, 153 

hallbergii Blatter. 153 

homiltonionwn Wallich ex C.B. Clarke, 153 

herbocewn (Roxburgh) Wallich, 153 

heterophyllwn (Brandegee) Matuda, 153 

hirticeps C.B. Clarke ex J.K. Morton, 153 

hirtum A. Richard, 1, 7-10, 16, 22. [44], [45]. 1461, 47-49, 147, 153, 

155, Plates 4,7 

hispidwn D. Don, 153 

hockii De Wildeman, 1,7-9, 16. [16L 17,1.181, 19, [19], [20], 22, 23, 

153, Plate 5 

113, [115], 116-118,120-122,125,153, 155, Plates 3.6 

143, 150,153, Plate 1 

26, [30], 31, [311, 32, 35,36. 38, [381, [391, 42, 1441, [461, 47-50, 

[56], 58,62,76, 105, 142, 153, 155, Plates 1, 5 

holosericeum (Kunth) Woodson, 153 

homblei De Wildeman, 7,23, 144-146,153 

hookeri C.B. Clarke, 153 

humile Warburg, 153 

hwnile Merrill, 153 

imberbe Ridley, 153 

imbricotwn Warburg, 153 

indehiscens Faden, v, 1. 7-9. 11, 16, 36, 38. 49, 50, 91, 93, 98, 100, 

subsp. indehiscens, [241, [271, 36, 50, 100, 11011, 102, [103], 

subsp. keniense Faden, v. 1,102, [103], 104, 105, 106,110, Plate 3 

subsp. lilacinum Faden, 26, 100, [loll, 102, [103], 104, 105, 106, 

[loll, 102.105, 106, 110. 125, 143, 153 

104-106, Plate 3 

Plate 6 

irreguloribrocteotum Kew Bulletin, 153 

joponicwn munberg) Kunth, 153 

joponicum Maximowicz, 153 

johnstonii K. Schumann, 1,7,9, 10. 16, 17,22,23,31,36, [46], 47-50, 

jwrghuhnionwn Miquel, 153 

koinonense Masamune. 153 

korwinskyonwn (J.A. & J.H. Schultes) Woodson. 153 

kotongense De Wildeman, 153 

keisok Hasskarl, 153 

keyense Warburg, 153 

koenigii Wallich ex C.B. Clarke. 153 

61,62,73, 150, 153, Plates 1,7 

kuntzei C.B. Clarke ex Kuntze, 153 

lamuense Faden, v, 1,7-11, 13, 16, 17, [251.26. [291,32, 49, 92,93. 

130, [132], 133, 134, [135], 136, 138, 140, 143, 153. Plates 4,6 

lanceolatum Bentham, 7,s. 32,42,81,145,146,152-155 

subsp. lanceolatum, 1, 16,32 

subsp. subnudum (A. Chevalier) J.K. Morton, 8, 16, 155 

loncifoliwn Griffith, 153 

kanyfoliwn Griffith, 154 

lonuginoswn Wallich ex C.B. Clarke, 154 

lotifoliwn Wight, 154 

Iariforwn Bentham ex C.B. Clarke, 154 

lmwn R. Brown, 154 

1eiocauleK. Schumann, 1,7,16, 147,154, Plate4 

leptospermwn K. Schumann ex Faden, 94,98,154 

lineore (Bentham) Woodson, 154 

lineolotwn (Blume) Kunth, [56], 154 

longicapsa Faden, v, 1, 7, 16, 17, 36, 38, 76-78, [80], 83, 85, 86, 87. 

longifoliwn Hooker, 154 

longifoliwn Wallich, 154 

lcngirrhizum Faden, 1,7-9, 16, 17,20, [21], 22, 36, 42,58,59,61,62, 

loriforme Hasskarl, 154 

loureirii Hance, 154 

lozueiroi Index Kewensis, 154 

lujoi De Wildeman & Th. Durand, 154 

mocrophyllwn R. Brown, 55, 154 

rnacrorrhh'h. Fries, jr., 1,7,16,23, 144-146.152.154 

maloboricwn (L.) Memll, 154 

medicwn (Laureiro) Kostelesky. 154 

melonostictwn Hance. 154 

micronthwn (Vahl) Kunth, 154 

minutiflonun Faden, 1,7-10,16,22,23,38. 147,154, Plate 7 

minutwn (Blume) Kunth, 154 

monodelphwn (Blume) Kunth, 154 

montonwn (Wight) Wallich ex C.B. Clarke, 154 

mortehanii De Wildeman, 154 

mononii Brenan, 7,36,59, 145,146,154 

multhcoposwn Lauterbach, 154 

nonwn (Roxburgh) Kunth, 154 

neocsledonicum Schlechter, 1,2.7-9,14, 16.20, [21], 22.26, [30], 31, 

nicholsonii C.B. Clarke, 1.7. 8, 16, 17, 31, 38, 49, 147, 154, Plates 4, 

nigritknwn (C.B. Clarke) Hutchinson ex Hutchinson & Dalziel, 154 

nudicoule (N.L. Bumann) G. Don, 154 

nudiflorwn &) Sweet. 154 

nwnmuloriwn Miquel, 154 

nutons Leveille,, 154 

nyasense C.B. Clarke, 1, 7, 8, 20, [211, 23, 30, 31, 36, 43, [44], [451, 

154, Plate 2 

76, 150. 154, Plates 1.5 

1311.36, [38]. [41], 47-49.51.60, 148, 154. Plate 1 

7 

[461,47-49,62,92.98.154.155 

var. brevisepalum Brenan, 16, Plate 1 

var. ongustifoliwn Chiovenda, 85, 86 

obbiadense Chiovenda. 2,7.8, 16,36,76-78,83, [84], 85, 86, 154 

ochrocewn Dalzell, 154 

octospermum C.B. Clarke ex Brenan, 71,73. 154 

oligonthwn Franchet & Savatier, 154 

ovalifohm (Wight) Hooker f. ex C.B. Clarke, 154 

ovoto-oblongwn P. de Beauvois, 154 

ovotwn (Hasskarl) Wallich ex C.B. Clarke, 154 

paludosum A. Chevalier, 7.36.38.42.59, 145, 146, 154 

subsp. paludosum, 7 

subsp. pseudolanceolatum J.K. Morton, 7.53 

subsp. pauciflorum J.K. Morton, 7 
164 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

panicdatum Wight, 154 

panicdatum Wight ex C.B. Clarke. 154 

papuanwn Warburg, 36,53,60,154 

plraguayense C.B. Clarke ex Chodat, 154 

pauciflorwn DalzelI, 154 

pluciflorwn Wight, 154 

pauciforwn (Urban & Ekman) Main, 154 

paucifdiwn N.E. Brown. 154 

pedunculosum C.B. Clarke, 1,7-11,16,36,49,58,92,146,147,154 

petersii (Hasskarl) C.B. Clarke, 1,7-9,16,20, [21], 23,26,31,36,38, 

42,49,50,57,90, 91,93, 95,96, 98-100, 105. 106, 110, 116. 120, 

125,133,152, 154,155 

95, [961,97,98-100, 125. 142, Plates 2.6 

Plate 2 

subsp. pallidiflorum Faden, v, 2,3,78, 10, 11, [331,35,49, [56]. 72, 

subsp. petersii, 7, [241, [261, [271,[301, [371,50,73,95, [%I, 98,99, 

pilmwn Wallich, 154 

pinetorum (Greene) Matuda, 154 

plagiocapsa K. Schumann, 1,7,36,61, 62, 154 

platyphyllwn Med, 154 

poaeoides Seubert, 154 

pomeridianumStanfield& Brenan, 1,7,8,10,16,30,32,145,146,154, 

protenswn (Wight) Wallich ex C.B. Clarke, 154 

puberulwn Hasskarl, 154 

pulchellwn (Kunth) Woodson, 154 

pulneynse Fyson. 154 

pusillum Chiovenda, 7-10,16,36,38,50,75 78,87,88,89,154 

Plate 4 

subsp. gypsophilum Faden, v, 1,16, [84]. 87.88,89 

subsp. pusillum, 1,23, [841,88, 89, Plates 2,s 

subsp. thulinii Faden, v, 1, [84], 87,88,89, Plate 2 

subsp. variabile Faden, v, [84], 88, 89 

radicans D. Don, 154 

recuwatum Faden, v, 1.7. 16, [241, [271, [301,35, 38, [461,47-50,73, 

90, 93. 105, 106, [1071, 108, [1091, 110. 113, 116, 118, 125, 133, 

154, Plates 3.6 

rend1eiC.B. Clarke, 1.7.9. 16, 17, [171, [181,20, [21], 22,23,26, [30], 

31,32,35, [461,47-50. [%I, 62.64, [651,66, [67], [68], 69,70, 71, 

73.76.7781, 98, 106, 110, 113, 154, Plates 1.2.5 

rendlei x taylorii, [as], [68] 

reniforme Buchanan-Hamilton ex Wallich. 154 

rhodosperm K. Schumann ex C.B. Clarke, 154 

richardsiae Brenan, 1,7,38, 146,147, 154, Plate 7 

rigidwn Blatter, 154 

ringoetii De Wildeman, 155 

rivulare A. Richard, 155 

rugmwn Pemer de la Bathie, 155 

rrcsseggeri (Fend) C.B. Clarke, 155 

saclewu’i Hua, 94,98, 155 

scaberrimwn (Blume) Kunth, 155 

scapiflorwn (Roxburgh) Kostelesky, 155 

schlechteri K. Schumann, v, 7,16,36,98,144-146,155 

schombwgkianum Kunth, 155 

schweinfurthii Baker ex C.B. Clarke, 155 

sclerocarpum F. Mueller, 7,36.60, 155 

sebitense Faden, v, 1, 3, 7, 16, 73, 90. 93, 1961, 110, [lll], 113, 117, 

secundiflonun (Blume) Kunth, 155 

secundwn Wight, 155 

semifoliatwn C.B. Clarke, 155 

semiteres Dalzell, 155 

sepaloswn C.B. Clarke, 155 

serotinwn D. Don ex C.B. Clarke, 155 

serrulatwn (Vahl) Loudon. 155 

118, 125. 155. Plate 3 

sessilifolium (Bentham) J.K. Morton, 155 

setifenun A. Chevalier, 7,16.23,32,42,145,146, 155 

sinmense Craib, 155 

siennewn Blatter, 155 

siliculosum R. Brown, 7,58,60, 155 

silvaticum Brenan, 1,7,145, 146, 152, 154, 155 

simplex (Vahl) Kunth, 155 

sinicwn Ker-Gawler, 155 

smithii C.B. Clarke, 79, 81, 155 

somaliense C.B. Clarke, 1,2,7, 16, 17,22,26, [301,[33], [341,35, 36, 

soudanicwn C.B. Clarke, 155 

species aff. spekei C.B. Clarke, 50 

species C, 106 

species nov. aff. umbroswn (Vahl) Kunth, 64 

spectabile Kun, 155 

spekeiC.B. Clarke, 1,2,7-11,16,49,116,146.147,155 

spicafwn Wallich, 155 

spiratwn (L.) Sweet, 30, 155 

stenothyrswn Eels, 155 

stolzii Mildbraed ex J.K. Morton, 155 

subnudwn A. Chevalier, 42, 155 

subovatwn Ridley. 155 

succulentum Faden. v, 1, 2, 7-9, 11, 13, 16, 17, [21], 22, [25], [26]. 

[291,42,49,50.92,93,130,[1321,136, 138, [139], 140,143,155, 

Plates 4.6 

tacazzeanum Hochstetter ex A. Richard, 79.92.94, 106, 113, 116, 117, 

155 

tacazzeanum Hochstetter ex C.B. Clarke, 113, 155 

tanaense Faden, v, 1,7-10,16,20,[211,22,23, [241, [XI, [281,32,35, 

43, [46]. 47-49,76,77 78, [SO], 81.82. 116, 143, 155, Plates 2,5 

38, 49-51.91-93, [1241,125, [126], 128, 130, 133, 142, 143, 155, 

Plates 3,6 

taquetii Leveille, 155 

tay1oriiC.B. Clarke, 1,2,7,9, 16, 17, 19,20.26,31, [31], 32.38, [46], 

47-49, 64, [651,66, [671, [681,69-71,73, 76, 92. 130, 155, Plates 

1.5 

tenerwn Baker, 155 

tenuirsimwn (A. Chevalier) A. Chevalier ex Hutchinson & Dalziel, 155 

terminale Wight, 155 

terminale (Blume) Boldingh ex Haines, 155 

termitarium Faden, 1,7-10,16.23.47-49, 147. 155 

tetraspermwn K. Schumann, 93.98, 155 

thomronii (C.B. Clarke) C.B. Clarke, 155 

trichocolewn Schauer, 155 

triquetrum Wallich ex C.B. Clarke, 155 

tuberoswn Buchanan-Hamilton ex Wallich, 155 

umbrosum (Vahl) Kunth. 1.7,9, 11, 16,20, [21], 23,32,38,42,43,50, 

subsp. ovato-oblmgum (p. de Beauvois) J.K. Morton, 8, [9]. 10, [56], 

subsp. umbrosum. 19.32.50,154 

wbinanwn (Greenman) Matuda, 155 

usambarenseFaden, v. 2.7,16,17,64, [671,69, [69],70,71,73.76,155 

vaginarum (L.) Wallich. 155 

vankerckhovenii De Wildeman. 155 

versicolor Dalzell, 155 

vioiacewn F. Mueller ex C.B. Clarke, 155 

vitiense Seemann. 155 

w4witschiiC.B. Clarke, 1,7,9,10,16,20, [211,23,26, [30],31,36,43, 

whytei C.B. Clarke, 155 

wildii Merxmuller, 155 

woodii Faden. v, 1.7,16,91,93,118, [1191,120, 121,[121], 122, 125, 

51,57,64,66,144-146,152, 154,155 

66,150,154. Plate 7 

49,81, 144-146,152, 153, 155, Plates 4,7 

155, Plate 3 
NUMB= 76 165 

woodii x forskalii, 1,120, [121], Plate 3 

zebrinum Chiovenda, 1,2,7-9, 11, 13, 16,22, [251, [26],[28].36,38, 

42, [44], [46]. 47-50.57,90.92,93,100, 133, 138.140, 142, [142], 

143,155, Plates 4.7 

zeylanicwn C.B. Clarke, 155 

sepalosa (C.B. Clarke) Engler, 155 

Athericopsis Engler, 58 

AphylaxSalisbury, 151 

Ballya Brenan, 57,90,92,142, 143 

Bauschia Seubert, 5557 

Buforrestia C.B. Clarke, 54 

zebrina (Chiovenda) Brenan, 42,57,90.140 

bracteolata (Martius) Seubert, 57 

Callisia hfling 

fiiliformis (Martens & Galeotti) D.R. Hunt, 154 

gracilis (Kunth) D.R. Hunt, 153 

Cochliostemma Lemaire, 23 

Colemype C.B. Clarke, 23 

Commelina Plumier ex L., 1,8, 13,23,26,36,50.54,55,57, 116 

aequinoctialis P. de Beauvois. 62 

beniniemis P. de Beauvois, 145 

ebracteata Ehrenberg, 113 

erecta L., 49 

hberbis Ehrenberg ex Hasskarl, 42 

panicdata VaN. 113, 116 

tuberosa Forsskal, 113.116 

wnbroswn VaN, 145 

Commeheae, 57 

Cyanotis D. Don. 9,143 

moluccana (Roxburgh) Merrill. 155 

Dichaespermum Wight, 151 

Dichoespermwn Wight, 151 

Dichorisandra Mikan, 153 

Dictyospemum Wight, 55, 56,58, 151 

section Dictyospermum, 56 

section Tricarpelema, 56 

conspimum (Blume) Hasskarl, 152, 153 

humile (Warburg) J.K. Morton, 153 

montanum Wight, 154 

ovalifolium Wight, 153,154 

ovatum Hasskarl, 154, 155 

phillipeme Panigrahi, 153 

Dilasia Rafiiesque. 15 1 

Ditelesia Rafiiesque, 151 

Floscopa Loureiro, 23,42,54,55 

africana (P. de Beauvois) C.B. Clarke, 152 

glomerata (Willdenow ex J.A. & J.H. Schultes) Hasskarl, 55, 155 

scandens Loureiro, 152.153 

yunnanensis Hong, 56 

Geogenanthus Ule, 23 

Gibasis Rafmesque, 154,155 

chihuahumsis (Standley) Rohweder, 152 

geniculata (Jacquin) Rohweder, 153 

karwinskyana (J.A. & J.H. Schultes) Rohweder, 152, 153 

linearis (Bentham) Rohweder, 154 

pulchella (Kunth) Rafiiesque, 154 

triflora (Martens & Galeotti) D.R. Hunt, 152 

venustula (Kunth) D.R. Hunt 

subsp. peninsulae D.R. Hunt, 153 

Lamprodithyros Hasskarl, 8.55.57.92 

beniniemw (P. de Beauvois) Hasskarl, 92 

dregeanw (Kunth) Hasskarl, 92 

ehrenbergii Hasskarl, 113,116 

petersii Hasskarl. 57.90,92,93, 96 

radicam, 92 

schomburgkianus (Kunth) Hasskarl, 92 

stenorrachys Hasskarl ex C.B. Clarke, 92 

tacazzeanw (Hochsteter ex A. Richard) Hasskarl, 92, 113, 116 

MurdanniaRoyle, 9,23,30,54-56,58, 151, 152, 153, 154 

acutifolia (Lauterbach & K. Schumann) Faden, 152 

allardii (De Wildeman) Brenan, 152 

blumei (Hasskarl) Brenan. 152, 153 

bracteata (C.B. Clarke) J.K. Morton ex Hong, 152,153 

clandestina (Ridley) Faden, 152 

clarkeana Brenan. [56], 152 

crOcea (Griffith) Faden, 152,154 

subsp. ochracea (Dalzell) Faden, 154 

dimorpha (Dalzell) Bruckner. 153,154 

divergens (C.B. Clarke) Bruckner, 153 

edulis (Stokes) Faden, 153-155 

esculenta (Wallich ex C.B. Clarke) Raizada ex Rolla Rao & Kammathy, 

gardneri (Seubert) Bruckner, 153 

gigantea (Vahl) Bruckner, 153 

glauca (Thwaites ex C.B. Clarke) Bruckner, 153 

graminea (R. Brown) Bruckner, 152, 153, 155 

hookeri (C.B. Clarke) Bruckner, 152, 153 

japonica (Thunberg) Faden, [56]. 153, 154 

kainanensis (Masumune) Hong, 153 

keisak (Hasskarl) Handel-Mazzetti, 152-155 

lanceolata (Wight) Kammathy, 153 

lanuginosa (Wallich ex C.B. Clarke) Bruckner, 154, 155 

loriformis (Hasskarl) Rolla Rao & Kammathy, 154, 155 

medica (Loureiro) Hong, 154 

nudiflora (L.) Brenan, 152-155 

paraguayensis (C.B. Clarke ex Chodat) Bruckner, 154 

pauciflora Bruckner, 154 

schomburgkiana (Kunth) Bruckner, 155 

semifoliata (C.B. Clarke) Bruckner, 155 

semiteres (Dalzell) Santapau, 154,155 

simplex (Vahl) Brenan, 23, [56], 152, 154, 155 

spectabilis (Kun) Faden. 152, 155 

spirata (L.) Bruckner, 30, 152, 154, 155 

stenothyrsa (Diels) Handel-Mazzetti, 155 

tenuissima (A. Chevalier) Brenan, 155 

triquetra (Wallich ex C.B. Clarke) Bruckner, 154, 155 

vaginata (L.) Bruckner, 153-155 

versicolor (Dalzell) Bruckner, 155 

zeylanica (C.B. Clarke) Bruckner, 36, [56], 155 

153, 154 

Palisota Reichenbach ex Endlicher, 32 

Permanthero Fend, 57 

Phaeneilema Bruckner, 55 

Piletocarpus Hasskarl, 55, 151 

Pollia Thunberg, 54-56,153, 154 

ambigua (P. de Beauvois) C.B. Clarke, 152 

crispata (R. Brown) Bentham. 152 

macrophylla (R. Brown) Bentham, 154 

secundiflora (Blume) Bakhuizen van den Brink f., 153,155 

Prionostachys Hasskarl, 55, 151 

Pseudoparis Pemer de la Bathie 

tenera (Baker) Faden, 155 
166 

Rhopalephora Hasskad, 43,5556, 150,151 

micrantha (Vahl) Faden. v, 154 

monadeipha (Biume) Faden, v 

rugosa (Pemer de la Bathiee) Faden. 155 

scabenha (Blume) Faden, 154, 155 

vitiensis (Seemam) Faden, 153,155 

Strepfylis Rafinesque. 151 

Talipula Rafinesque, 151 

'Ihyrsanthemum Pichon 

SMITHSOMAN CONTRIBUTIONS TO BOTANY 

floribundum (Martens & Galeotti) F'ichon, 153 

macrophyllum (Greenman) Rohweder, 153 

Tinantia Scheidweiler, 8 

Tradescantia L, 1,9 

pinetorum Greene. 154 

Tricarpelema J.K. Morton, 55.56,58 

giganteurn (Hasskari) Ham, 155 

glandulifenun (Joseph & Rolla Rao) Faden. 153 

philippense (Panigrahi) Faden, 153 

multiflora (Swartz) Rafinesque, 153 

Tripogandra Rafinesque, 32,42 
Plates 1-7
PLATE l.-Flowers of Aneilema species. Sections Aneilema, Amelina, and Rendlei (in part). Section Aneilema: 

(a,b) A. acwninatwn R. Brown: a, perfect flower, front view; b, perfect flower, top view, showing reflexed medial 

petal (from Blaxell 1091); c, A. neocaledonicwn Schlecter, perfect flower, front view (from Bierhorst in Kew 

Accession No. 000-69.12046). Section Amelina: d, A. johnstonii K. Schumann, perfect flower, top view (from 

Faden & Faden 741490); (e,J A. gillettii Brenan: e, perfect flower, front view; f, perfect flower, side view (from 

Friis 1044); (g,h,l) A. hockii De Wildeman: g, inflorescence, showing perfect and staminate flowers; h, perfect 

flower, top view; I, pistillate flower, side view (g from Faden & Faden 7411 182, h from Faden & Faden 7411065, 

1 from Faden et al. 72/234); i, A. aequinoctiale (P. de Beauvois) G. Don, perfect flower, top view (from Faden 

et al. 71/922); j, A. nyasense C.B. Clarke var. brevisepala Brenan, perfect flower, side view (from Faden et al. 

701369); k, A. longirrhizwn Faden, perfect flower, front view (from Faden & Faden 741217). Section Rendlei: 

(ma) A. taylorii C.B. Clarke: m, perfect flower, front view; n, perfect flower, side view (from Faden & Faden 

74/371); (op) A. rendlei C.B. Clarke: 0, perfect flower, front view; p, perfect flower, side view (from Kabuye & 

Evans 163). 

PLATE 2.-Flowers of Aneilema species. Sections Rendlei (in part), Somaliemia, and Lurnprodithyros (in part). 

Section Rendlei: (ax) A. brenanianum Faden: u, habit, showing lower portions of stems covered by long, 

overlapping sheaths; b, perfect and staminate flowers, side views; c, inflorescence (from Faden & Fuden 72/166); 

d, A. rendlei C.B. Clarke, inflorescence, showing perfect and staminate flowers (from Faden et al. 69/322). 

Section Somuliensia: (e-g) A. somalieme C.B. Clarke: e, inflorescence;f, pistillate flower, front view; g, perfect 

flower, front view (from Faden & Faden 74923); (h,l) A. longicapsa Faden: h, perfect flower, front view; I, 

perfect flower, side view (from Gilletf & Becktt 23289); (ij) A. pusiNwn Chiovenda subsp. purillum): i, habit; 

j, perfect flower, front view (from Gillett et ul. 22597); k, A. pusillum Chiovenda subsp. thulinii Faden, perfect 

flower, front view (from Thulin & Warfa 5320). Section Lumprodithyros: (m,n) A. petersii (Hasskarl) C.B. Clarke 

subsp. pefersii: m, inflorescence; n, perfect (lower) and staminate (upper) flowers, front view (m from Faden et 

al. 691453; n from Andrews 8.n.); (09) A. pefersii (Hasskarl) C.B. Clarke subsp. pallidiflorum Faden: 0, 

inflorescence, showing erect inflorescence bract serving as a background for the flowers; p, perfect flower, front 

view (0 from Faden & Faden 741246; p from Faden & Faden 741237). 

PIATE 3.-Flowers of Aneilema species. Section Lumprodithyros (in part): a, A. indehiccens Faden subsp. 

indehircens, perfect flower, front view (from Faden & Faden 72172); b, A. indehiscenr Faden subsp. kenienre 

Faden, perfect flower, front view (from Faden & Faden 771788); (c,d) A. recurvatum Faden: c, perfect flower, 

showing lateral stamens retained in lower petal, side view; d, perfect flower, stamens released, side view (from 

Faden et al. 6911066); e, A. sebitense Faden, perfect flower, front view (from Gilbert & Thulin 266);J A. forskalii 

Kunth, perfect flower, front view (from Gilbert & Thulin 969); g, A. woodii Faden x forskalii Kunth, perfect 

flower, front view (from Wood 2616A); h, A. woodii Faden, perfect flower, front view (from Wood 26168); (id) 

A. tanaense Faden: i, habit; j, terminal inflorescence on primary shoot; k, perfect flower showing lateral stamens 

retained in lower petal, front view, I, perfect flower, stamens released, front view (i,k,l from Faden & Faden 

7411053; j from Faden & Faden 7411185); (ma) A. calceolus Brenan: m, mat-forming habit; n, perfect flower, 

front view; 0, perfect flower, side view (m from Faden & Faden 7411069; n and o from Organ in EA14970); p, 

A. clarkei Rendle, shoot with two inflorescences, each with one perfect flower (from Faden & Faden 711809). 



PLATE 5.Seeds of Aneilema species: a, A. taylorii C.B. Clarke (from Faden & Faden 741371); b, A. 

brenanianwn Faden (from Faden et al. 6912069); c, A. grandibracteolatum Faden (from Glover & GilIiland408); 

d, A. ephemerwn Faden (from Faden C? Faden 741794); e. A. hockii De Wildeman (fm Faden & Faden 741215); 

fi A. rendlei C.B. Clarke (fran Faden et al. 691322); g. A. somaliense C.B. Clarke (from Faden & Faden 741923); 

h, A. pusillwn Chiwenda subsp. pusillum (from Paoli 1277); i, A. longirrhizwn Faden (from Faden & Faden 

741217); j, A. hockii De Wildeman (from Faden & Faden 7411065). (a = apical ventral lode seed, b = basal 

vend locule seed, m = medial ventral lode seed, o = ventral locule seed from one-seeded locule, u = 

unifferentiated ventral locule seed.) (All to the same scale.) 

PLATE 6.4eeds of Aneilema species: a, A. petersii (Hasskarl) C.B. Clarke subsp. pallidiforum Faden (from 

Faden & Faden 741753); b, A. indehiscens Faden subsp. lilacinum Faden (from Faden & Faden 741202); c, A. 

recurvatwn Faden (from Faden 6911300); d, A. forskalii Kunth (from Maffey 51); e, A. benadirense Chiovenda 

(from Ciferri 73); h A. calceolus Brenan (from Faden & Faden 741333); g, A. tanaense Faden (from Gillett 

19973); h, A. clorkei Rendle (from Faden et al. 711633); i, A. lamuense Faden (from Faden & Faden 7411083); 

j, A. succulentum Faden (from Faden et al. 701937). (a = apical ventral locule seed, b = basal ventral locule seed, 

m = medial ventral locule seed, o = ventral locule seed from one-seeded locule, d = dorsal locule seed.) (All to 

the same scale.) 

PLATE 7.Seeds of Aneilema species: a, A. zebrinum Chiovenda (from Faden & Faden 7411052); b, A. 

nicholsonii C.B. Clarke (from Faden et al. 741192); c, A. minufif[orum Faden (from Pogef-Wilkes A); d, A. hirtum 

A. Richard (from Milne-Redhead & Toylor 10242); e, A. richardsiae Brenan (from Richards 9960A); f, A. 

johnsfonii K. Schumann (from Faden & Faden 7411 10); g, A. wnbrosum (Vahl) Kunth subsp. ovato-oblongum (P. 

de Beauvois) J.K. Morton (frm Lye & Katende 6235); h, A. welwitschii C.B. Clarke (from Faden & Faden 

741157); i, A. brwtneospermum Faden (from Faden et 01. 741209); j, A. arenicola Faden (from Faden & Faden 

74/21 1). (a = apical ventral locule seed, b =basal ventral locule seed, m = medial ventral locule seed, o =ventral 

locule seed from one-seeded locule, d = dorsal locule seed.) (All to the same scale.)