Ethnobotany of The Mountain Regions of Africa 2021

Ethnobotany of Mountain Regions
Series Editors:
R. W. Bussmann · N. Y. Paniagua-Zambrana
Rainer W. Bussmann Editor
Ethnobotany of
the Mountain
Regions of Africa
Ethnobotany of Mountain Regions
Series Editors
Rainer W. Bussmann
Department of Ethnobotany
Institute of Botany and Bakuriani Alpine Botanical Garden
Ilia State University
Tbilisi, Georgia
Saving Knowledge
La Paz, Bolivia
Narel Y. Paniagua-Zambrana
Department of Ethnobotany
Institute of Botany and Bakuriani Alpine Botanical Garden
Tbilisi, Georgia
Saving Knowledge
La Paz, Bolivia
Herbario Nacionál de Bolivia
Universidad Mayor de San Andrés
La Paz, Bolivia
Ethnobotanical research in recent years has increasingly shifted into applied aspects
of the discipline, including climate change research, conservation, and sustainable
development. It has by now widely been recognized that “traditional” knowledge is
always in flux and adapting to a quickly changing environment. Trends of global-
ization, especially the globalization of plant markets, have greatly influenced how
plant resources are managed nowadays. While ethnobotanical studies are now
available from many regions of the world, no comprehensive encyclopedic series
focusing on the worlds mountain regions is available in the market. Scholars in plant
sciences worldwide will be interested in this website and its dynamic content.
The field (and thus the market) of ethnobotany and ethnopharmacology has grown
considerably in recent years. Student interest is on the rise, attendance at professional
conferences has grown steadily, and the number of professionals calling themselves
ethnobotanists has increased significantly (the various societies—Society for
Economic Botany, International Society of Ethnopharmacology, Society of Ethnobi-
ology, International Society for Ethnobiology, and many regional and national
societies in the field currently have thousands of members). Growth has been most
robust in BRIC countries.
The objective of this new series on Ethnobotany of Mountain Regions is to take
advantage of the increasing international interest and scholarship in the field of
mountain research. We anticipate including the best and latest research on a full
range of descriptive, methodological, theoretical, and applied research on the most
important plants for each region. Each contribution will be scientifically rigorous and
contribute to the overall field of study.
More information about this series at http://www.springer.com/series/15885

Rainer W. Bussmann
Editor
Ethnobotany of the
Mountain Regions of
Africa
With 666 Figures

Editor
Rainer W. Bussmann
Department of Ethnobotany, Institute of
Botany and Bakuriani Alpine Botanical
Garden
Tbilisi, Georgia
Saving Knowledge
La Paz, Bolivia
ISSN 2523-7489 ISSN 2523-7497 (electronic)

ISBN 978-3-030-38385-5 ISBN 978-3-030-38386-2 (eBook)
ISBN 978-3-030-38387-9 (print and electronic bundle)
https://doi.org/10.1007/978-3-030-38386-2
© Springer Nature Switzerland AG 2021
This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the
material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms or in any other physical way, and transmission or information
storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology
now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors
or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims
in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
This volume is dedicated to three great
“African” botanists:
Prof. Dr. Dr. h.c. Erwin Beck, who never
hesitated to give an enthusiastic student,
including this editor, the opportunity for
botanical work in Africa.
Prof. Dr. John O. Kokwaro, 1940–2019,
leading scholar in East African plant
taxonomy and ethnobotany, whose Medicinal
Plants of East Africa was the first
encyclopedia on the topic.
Mr. Simon G. Mathenge, ca. 1938–2017,
para-taxonomist at the University of Nairobi
Herbarium, without whose help the fieldwork
and plant identification of none of many a
thesis, including of this editor, would have
been done.
Preface
The use of plants by humans stretches back to the very beginning of human
evolution, and Africa, as the cradle of humankind, is certainly a hotspot of human-
plant interactions.
While much of the African continent shows some mountain regions and high
plateaus, the East African Rift and Ethiopian plateau are without doubt the moun-
tainous areas of largest extension of the continent. In many ways, they are also the
most populated and ethnically diverse systems of Africa. However, many peoples in
mountain areas arrived relatively late, especially during the large Bantu migration in
the seventeenth and eighteenth centuries.
Interestingly, in contrast to many other areas where leaves are used as medicine,
plant use in Africa often focuses on barks and tubers, raising the issue of sustain-
ability. Many species are used to produce strengthening soups, often important for
the digestion of meat consumed during rituals, and many medicinal species are still
used to purge disease-causing agents from patients’ bodies.
Plant use in Africa has been documented especially since the 1970s, and the
above-cited Medicinal Plants of East Africa of Prof. John Kokwaro is a striking
example of these efforts. In the last two decades, lots of local ethnobotanical studies
have been published, with Ethiopian scientists leading the way into modern ethno-
botanical research, while the documentation – and recuperation – of traditional
knowledge has gained renewed importance in the whole of Africa more recently.
With the present volume, we aim to provide an in-depth introduction to the plant
use knowledge of the peoples of the main tropical African mountain areas. Follow-
ing an introduction to Africa. and the wider region, over 170 plant chapters feature a
modern overview on taxonomy, local names, and information on the ecology and
distribution of all species. The ethnobotanical information provides both an over-
view on historic uses as well as data from the most recent scientific studies of plant
use in the region and contains the most up-to-date literature sources.
We hope that the present volume will give both interested laypeople as well as
professionals an opportunity to learn about the fascinating biodiversity and plant use
culture of the African mountains, and will spark interest in its further documentation,
sustainable use, and conservation.
vii
Contents
Volume 1
Part I Regions ........................................... 1

Ethnobotany of Mountain Regions: Africa . . . . . . . . . . . . . . . . . . . . . . 3
Rainer W. Bussmann, Narel Y. Paniagua-Zambrana, and Grace N. Njoroge
Ethnobotany of Mountain Regions: Eastern Africa . . . . . . . . . . . . . . . . 43
Part II Plants ............................................ 67
Abutilon sp. Abutilon angulatum (Guill. & Perr.) Mast. Abutilon

grandiflorum (Will.) Sweet. Abutilon longicuspe Hochst. ex A. Rich.
Abutilon mauritianum (Jacq.) Medik. Hibiscus sp. Hibiscus
calyphyllus Cav. Hibiscus vitifolius L. MALVACEAE . . . . . . . . . . . . . . . . . 69
Acacia drepanolobium Harmes ex Sjostedt Acacia nilotica (L.) Willd.
ex Delile Acacia senegal (L.) Willd. Acacia seyal Delille Acacia
tortilis (Forssk.) Hayne FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Acanthus eminens C.B. Clarke Acanthus sennii Chiov.
ACANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Achyranthes aspera L. AMARANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 101
Acokanthera schimperi (A. DC.) Benth. & Hook. f. ex Schweinf.
EUPHOPRBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Adansonia digitata L. MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
ix
x Contents
Aerva lanata (L.) Juss. ex Schult. AMARANTHACEAE . . . . . . . . . . . . . . . . . 119

Afrocanthium keniense (Bullock) Lantz Afrocanthium lactescens
(Hiern) Lantz Canthium glaucum Hiern. RUBIACEAE . . . . . . . . . . . . . . . 123
Ageratum conyzioides L. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Albizia anthelminthica Brogn. Albizia gummifera (J.F. Gmel.)
C.A. Sm. FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Alchemilla argyrophylla Oliv. Alchemilla johnstonii Oliv. ROSACEAE . . . 141
Aloe sp. Aloe vera (L.) Burm. f. ASPHODELACEAE . . . . . . . . . . . . . . . . . . . 147
Amaranthus hybridus L. Amaranthus spinosus L. AMARANTHACEAE . . . . 153
Anthocleista zambesiaca Baker GENTIANACEAE . . . . . . . . . . . . . . . . . . . . 161
Artemisia absinthium L. Artemisia afra Jacq. ex Willd. ASTERACEAE . . . 165
Asparagus africanus Lam. Asparagus falcatus L. ASPARAGACEAE . . . . . . 173
Asystasia gangetica (L.) T. Anderson ACANTHACEAE . . . . . . . . . . . . . . . . 181
Azadirachta indica A. Juss. MELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Balanites aegyptiaca (L.) Delile Balanites pedicellaris Mildbr. &
Schltr. Balanites rotundifolia (Tiegh.) Blatt. ZYGOPHYLLACEAE . . . . . . . . 189
Basella alba L. BASELLACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
Berberis holstii Engl. BERBERIDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Bersama abyssinica Fresen. FRANCOACEAE . . . . . . . . . . . . . . . . . . . . . . . 211
Bidens pilosa L. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
Contents xi
Blighia unijugata Bak. SAPINDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221

Bothriocline longipes (Oliv. & Hiern.) N.E. Br. ASTERACEAE . . . . . . . . . 223
Brachylaena huillensis O. Hoffm. ASTERACEAE . . . . . . . . . . . . . . . . . . . . 227
Brucea antidysenterica J.F. Mill. SIMAROUBACEAE . . . . . . . . . . . . . . . . . . 231
Buddleja polystachya Fresen. SCROPHULARIACEAE . . . . . . . . . . . . . . . . . . 237
Caesalpinia volkensii Harms FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 243
Calodendrum capense (L. f.) Thunb. RUTACEAE . . . . . . . . . . . . . . . . . . . 249
Capparis tomentosa Lam. CAPPARIDACEAE Gynandropsis gynandra
(L.) Briq. CLEOMACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Carissa spinarum L. APOCYNACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
Casearia battiscombei R.E. Fr. SALICACEAE . . . . . . . . . . . . . . . . . . . . . . . 267
Cassipourea malosana (Baker) Alston RHIZOPHORACEAE . . . . . . . . . . . . . 269
Catha edulis Forssk. CELASTRACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Celtis africana Burm. f. CANNABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Cissampelos pareira L. MENISPERMACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 285
Clausena anisata (Willd.) Hook. f. ex Benth. RUTACEAE . . . . . . . . . . . . . 289
Clematis brachiata Thunb. RANUNCULACEAE . . . . . . . . . . . . . . . . . . . . . . 293
Clerodendrum myricoides (Hochst.) R. Br. ex Vatke LAMIACEAE . . . . . . . 301
xii Contents
Clutia abyssinica Jaub. & Spach. PERACEAE . . . . . . . . . . . . . . . . . . . . . . 305

Coccinia grandis (L.) Voigt CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . 309
Coffea arabica L. RUBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
Cola greenwayi Brenan MALVACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319
Commelina africana L. Commelina benghalensis L. COMMELINACEAE . . . 321
Commiphora africana (A. Rich.) Engl. Commiphora rostrata Engl.
Commiphora schimperi (O. Bern.) Engl. BURSERACEAE . . . . . . . . . . . . . . 331
Conyza volkensii O. Hoffm. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 337
Cordia africana Lam. Cordia monoica Roxb. Cordia sinensis Lam.
BORAGINACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
Crotalaria agatiflora Schweinf. Crotalaria fascicularis Polhill
Crotalaria incana L. Crotalaria natalitia Meisn. FABACEAE . . . . . . . . . . . 351
Croton dichogamus Pax Croton macrostachyus Hochst ex Delile
Croton megalocarpus Hutch. EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . 357
Cucumis dipsaceus Ehrenb. ex Spach CUCURBITACEAE . . . . . . . . . . . . . . 367
Cupressus lusitanica Mill. CUPRESSACEAE . . . . . . . . . . . . . . . . . . . . . . . . 371
Cussonia holstii Harms ex Engl. ARALIACEAE . . . . . . . . . . . . . . . . . . . . . 375
Cyathea manniana Hook. CYATHEACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 379
Cyathula cylindrica Moq. Cyathula polycephala Bak.
AMARANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Cynodon dactylon (L.) Pers. POACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
Contents xiii
Cyphostemma maranguense (Gilg.) Desc. VITACEAE . . . . . . . . . . . . . . . . 399

Dactyloctenium aegyticum (L.) Willd. Dactyloctenium giganteum
B.S. Fischer & Schweick. POACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
Dalbergia lactea Vatke FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407
Datura stramonium L. SOLANACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
Dioscorea dodecaneura Vell. Dioscorea dumetorum (Kunth) Pax
Dioscorea minutiflora Engl. DIOSCOREACEAE . . . . . . . . . . . . . . . . . . . . . . 421
Diospyros abyssinica (Hiern.) F. White Diospyros mespiliformis
Hochst. ex A. DC. EBENACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
Dodonaea viscosa Jacq. SAPINDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
Dombeya torrida (J.F. Gmel.) Bamps Dombeya sp. MALVACEAE . . . . . . . 441
Dovyalis abyssinica (A. Rich.) Warb. Dovyalis caffra (Hook. f. &
Harv.) Warb. Dovyalis macrocalyx (Oliv.) Warb. SALICACEAE . . . . . . . . 453
Dracaena afromontana Mildbr. Dracaena steudneri Engl.
ASPARAGACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 461
Drypetes gerrardii Hutch. PUNTRANJIVACEAE . . . . . . . . . . . . . . . . . . . . . . 467
Ehretia cymosa Thonn. EHRETIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 469
Ekebergia capensis Sparrm. MELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 473
Englerina woodfordioides (Schweinf.) Balle ex M.G. Gilbert
LORANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
Ensete ventricosum (Welw.) Cheesm. MUSACEAE . . . . . . . . . . . . . . . . . . . 485
Rainer W. Bussmann, Narel Y. Paniagua-Zambrana, Grace N. Njoroge, and
John Grimshaw
xiv Contents
Eragrostis tef (Zuccagni) Trotter POACEAE . . . . . . . . . . . . . . . . . . . . . . . 489

Erica arborea L. ERICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493
Erythrina abyssinica DC FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501
Erythrococca bongensis Pax EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . 507
Eucalyptus globulus Labill. MYRTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 511
Euclea divinorum Hiern. EBENACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519
Euphorbia candelabrum Welw. Euphorbia tirucalli
L. EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
Fagaropsis angolensis (Engl.) Dale RUTACEAE . . . . . . . . . . . . . . . . . . . . . 533
Faurea saligna Harv. PROTEACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 535
Ficus sycomorus L. Ficus thonningii Blume MORACEAE . . . . . . . . . . . . . 537
Flacourtia indica (Burm. f.) Merr. SALICACEAE . . . . . . . . . . . . . . . . . . . . 549
Galinsoga parviflora Cav. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 553
Garcinia volkensii Engl. CLUSIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 557
Gnidia glauca (Fresen.) Gilg. THYMELAEACEAE . . . . . . . . . . . . . . . . . . . . 561
Volume 2
Grewia bicolor Juss. Grewia tembensis Fresen. Grewia tenax

(Forssk.) Fiori Grewia villosa Willd. MALVACEAE . . . . . . . . . . . . . . . . . . 567
Hagenia abyssinica J.F. Gmel. ROSACEAE . . . . . . . . . . . . . . . . . . . . . . . . 577
Contents xv
Harungana madagascariensis Poir. HYPERICACEAE . . . . . . . . . . . . . . . . . 591

Hypericum revolutum Vahl HYPERICACEAE . . . . . . . . . . . . . . . . . . . . . . . 593
John Grimshaw
Hyphaene compressa H. Wendl. Hyphaene coriacea Gaertn.
Hyphaene thebaica (L.) Mart. ARECACEAE . . . . . . . . . . . . . . . . . . . . . . . 599
Hypoestes aristata (Vahl) Sol. ex Roem. & Schult. Hypoestes
forsskaolii (Vahl.) R. Br. ACANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 611
Ilex mitis (L.) Radkl. AQUIFOLIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 615
Juniperus procera Hochst. ex Endl. CUPRESSACEAE . . . . . . . . . . . . . . . . . 619
Kalanchoe densiflora Rolfe Kalanchoe petitiana A. Rich.
CRASSULACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 633
Kigelia africana (Lam.) Benth. BIGNONIACEAE . . . . . . . . . . . . . . . . . . . . . 641
Kyllinga bulbosa P. Beauv. CYPERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . 649
Landolphia buchananii (Hallier f.) Stapf Landolphia kirkii DyIer
APOCYNACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653
Lantana camara L. VERBENACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
Lasianthus kilimandscharicus K. Schum. RUBIACEAE . . . . . . . . . . . . . . . 663
Lippia javanica (Burm. f.) Spreng. VERBENACEAE . . . . . . . . . . . . . . . . . . 667
Lobelia giberroa Hemsl. CAMPANULACEAE . . . . . . . . . . . . . . . . . . . . . . . . 673
Macaranga kilimandscharica Pax EUPHORBIACEAE . . . . . . . . . . . . . . . . . 677
Maerua angolensis DC. Maerua triphylla A. Rich. CAPPARACEAE . . . . . . 685
xvi Contents
Mangifera indica L. ANACARDIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691

Markhamia lutea (Benth.) K. Schum. BIGNONIACEAE . . . . . . . . . . . . . . . 697
Maytenus senegalensis (Lam.) Exell Maytenus undata (Thunb.)
Blakelock CELASTRACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 699
Moringa oleifera Lam. MORINGACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 711
Myrianthus holstii Engl. URTICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 717
Myrsine africana L. PRIMULACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 721
Neoboutonia macrocalyx Pax EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . 727
Nicotiana rustica L. Nicotiana tabacum L. SOLANACEAE . . . . . . . . . . . . . 731
Nuxia congesta R. Br. ex Fresen. LOGANIACEAE . . . . . . . . . . . . . . . . . . . 739
Ocimum gratissimum Lam. Ocimum kilimandscharicum Guerke
LAMIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 745
Ocotea usambarensis Engl. LAURACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . 753
Oldeania alpina (K. Schum.) Stapleton POACEAE . . . . . . . . . . . . . . . . . . 761
Olea capensis L. Olea europaea subsp. africana (Mill.) P.S. Greene
OLEACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777
Osyris lanceolata Hochst. & Steud. SANTALACEAE . . . . . . . . . . . . . . . . . . 787
Pavonia patens (Andrews) Chiov. Pavonia urens Cav. MALVACEAE . . . . . 791
Phoenix reclinata Jacq. ARECACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 795
Physalis peruviana L. SOLANACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 801
Contents xvii
Phytolacca dodecandra L’Hér. PHYTOLACCACEAE . . . . . . . . . . . . . . . . . . . 807

Pinus patula Schltdl. & Cham. Pinus radiata D. Don. PINACEAE . . . . . . 815
Piper capense L. PIPERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 825

Pistacia aethiopica Kokwaro ANACARDIACEAE . . . . . . . . . . . . . . . . . . . . . 831

Podocarpus milanjianus Rendle Afrocarpus falcatus (Thunb.)

C.N. Page Afrocarpus gracilior (Pilg.) C.N. Page PODOCARPACEAE . . . . . 835
John Grimshaw
Portulaca oleracea L. PORTULACACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 845

Pouteria adolfi-friederici (Engl.) A. Meeuse Pouteria altissima

(A. Chev.) Baehni SAPOTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 851
Prunus africana (Hook. f.) Kalkman ROSACEAE . . . . . . . . . . . . . . . . . . . 857

Psidium guajava L. MYRTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 863

Psychotria orophila E.A.A. Petit Psychotria sp. RUBIACEAE . . . . . . . . . . . 869

Pterolobium stellatum (Forssk.) Brenan FABACEAE . . . . . . . . . . . . . . . . . 873

Rapanea melanophloeos (L.) Mez. PRIMULACEAE . . . . . . . . . . . . . . . . . . . 879

John Grimshaw
Rhamnus prinoides L’Hér. Rhamnus staddo A. Rich. RHAMNACEAE . . . . 885

Rhus natalensis Bernh. ex C. Krauss Rhus ruspolii Engl. Rhus

tenuinervis Engl. Rhus vulgaris Meikle ANACARDIACEAE . . . . . . . . . . . . . 893
Ricinus communis L. EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 903

xviii Contents
Rubus apetalus Poir. Rubus rosifolius Sm. Rubus volkensii Engl.

ROSACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 911
Rumex abyssinicus Jacq. Rumex usambarensis (Dammer) Dammer
POLYGONACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 919
Salvadora persica L. SALVADORACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 927
Sambucus africana Engl. CAPRIFOLIACEAE . . . . . . . . . . . . . . . . . . . . . . . . 931
Schefflera volkensii (Harms) Harms ARALIACEAE . . . . . . . . . . . . . . . . . . 943
Schkuhria pinnata (Lam.) Kuntze ex Thell. ASTERACEAE . . . . . . . . . . . . 949
Scutia myrtina (Burm. f.) Kurz RHAMNACEAE . . . . . . . . . . . . . . . . . . . . . 953
Solanecio mannii (Hook. f.) C. Jeffrey ASTERACEAE . . . . . . . . . . . . . . . . 959
Solanum aculeastrum Dunal Solanum anguivi Lam. Solanum
incanum L. Solanum nigrum L. SOLANACEAE . . . . . . . . . . . . . . . . . . . . . 963
Sonchus asper (L.) Hill Sonchus oleraceus L. ASTERACEAE . . . . . . . . . . . 975
Spilanthes mauritiana (Rich. ex Pers.) DC. ASTERACEAE . . . . . . . . . . . . . 983
Stephania abyssinica (Quart.-Dill. & A. Rich.) Walp.
MENISPERMACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 985
Stoebe kilimandscharica O. Hoffm ASTERACEAE . . . . . . . . . . . . . . . . . . . 989
Strychnos henningsii Gilg. Strychnos spinosa Lam. Strychnos
usambarensis Gilg. LOGANIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 991
Syzygium cordatum Hochst. Syzygium guineense (Willd.) DC.
MYRTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 997
Contents xix
Tabernaemontana stapfiana Brizten APOCYNACEAE . . . . . . . . . . . . . . . . . 1005

Tagetes minuta L. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1009
Tamarindus indica L. FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1013
Tarchonanthus camphoratus L. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . 1017
Teclea simplicifolia Engl. Vepris dainelii (Pic. Serm.) Mziray
Vepris nobilis Delile. RUTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1021
Themeda triandra Forssk. POACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1031
Thunbergia alata Sims ACANTHACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1033
Toddalia asiatica (L.) Lam. RUTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1037
Trichilia emetica Vahl MELIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041
Typha domingensis Pers. Typha latifolia L. TYPHACEAE . . . . . . . . . . . . . 1045
Urtica maasaica Mildbr. URTICACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1053
Uvaria acuminata Oliv. Uvaria scheffleri L.L. Zhou, Y.C.F. Su &
R.M.K Saunders ANNONACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1059
Vangueria apiculata K. Schum. Vangueria infausta Burch.
Vangueria madagascariensis J.F. Gmel. Vangueria volkensii K.
Schum. RUBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1063
Vernonia amygdalina Deille Vernonia auriculifera Hiern Vernonia
galamensis (Cass.) Less. ASTERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 1069
Vigna friesiorum Harms Vigna membranacea A. Rich. Vigna
subterranea (L.) Verdc. Vigna unguiculata (L.) Walp.
FABACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1079
xx Contents
Vitex keniensis Turrill LAMIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1085

Warburgia salutaris (Bertol.) Chiov. CANELLACEAE . . . . . . . . . . . . . . . . . 1089
Withania somnifera (L.) Dunal SOLANACEAE . . . . . . . . . . . . . . . . . . . . . . 1093
Ximenia americana L. XIMENIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1099
Zanthoxylum chalybeum Engl. Zanthoxylum gilletii (De Wild)
Waterm. Zanthoxylum usambarense (Engl.) Kokwaro RUTACEAE . . . . . 1103
Zehneria scabra (L.f.) Sond. CUCURBITACEAE . . . . . . . . . . . . . . . . . . . . . . 1111
Ziziphus jujuba Mill. Ziziphus mauritiana Lam. Ziziphus mucronata
Willd. RHAMNACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1117
Rainer W. Bussmann, Narel Y. Paniagua-Zambrana, Grace N. Njoroge,
Maroof Ali Turi, and Hammad Ahmad Jan
About the Editor
Dr. Rainer W. Bussmann earned his M.Sc. (Diploma)

in Biology at the University of Tübingen, Germany, in
1993 and his doctorate at the University of Bayreuth,
Germany, in 1994. He is an ethnobotanist and vegetation
ecologist. He is currently co-director of Saving Knowl-
edge, La Paz, Bolivia, and Principal Scientist at the
Department of Ethnobotany, Institute of Botany, Ilia
State University, both of which he co-founded. Before,
Dr. Bussmann was director of the William L. Brown
Center (WLBC) at Missouri Botanical Garden, William
L. Brown Curator of Economic Botany, and Senior
Curator. Before accepting the directorship of WLBC,
he held academic appointments as Research Fellow in
Geography and the Environment at the University of
Texas at Austin from 2006 to 2007, as Associate Pro-
fessor of Botany and Scientific Director of Harold Lyon
Arboretum at the University of Hawaii from 2003 to
2006, and as Assistant Professor at the University of
Bayreuth from 1997 to 2003, following a postdoc at the
same institution from 1994 to 1997. He holds affiliate
faculty appointments at Washington University,
St. Louis, USA; University of Missouri–St. Louis,
USA; Florida Atlantic University, Boca Raton, USA;
Federal University of Paraíba, Brazil; National Univer-
sity of San Marcos, Peru; and Ilia State University,
Republic of Georgia, and serves as external thesis advi-
sor at multiple other universities worldwide. His work
focuses on ethnobotanical research and the preservation
of traditional knowledge in Bolivia, Peru, Madagascar,
the Caucasus, and the Himalayas. To date,
Dr. Bussmann has authored over 270 peer-reviewed
papers, over 1200 book chapters, and authored or edited
over 35 books.
xxi
xxii About the Editor
Dr. Bussmann is a past President of the Society for

Economic Botany and has served as board/council
member of the International Society for Ethnophar-
macology, Society of Ethnobiology, Botanical Society
of America, and International Society of Ethnobiology.
See more of his work on his website (https://www.
cejaandina.org/rainer-w-bussmann/) and download pub-
lications from ResearchGate (https://www.researchgate.
net/profile/Rainer_Bussmann).
Contributors
Rainer W. Bussmann Department of Ethnobotany, Institute of Botany and

Bakuriani Alpine Botanical Garden, Ilia State University, Tbilisi, Georgia
Saving Knowledge, La Paz, Bolivia
John Grimshaw The Yorkshire Arboretum, Castle Howard, York, UK
Hammad Ahmad Jan Islamia College Peshawar Jamrod Road, University
Campus Peshawar, Peshawar, Pakistan
Grace N. Njoroge Department of Botany, Jomo Kenyatta University of Agriculture
and Technology, Nairobi, Kenya
Narel Y. Paniagua-Zambrana Department of Ethnobotany, Institute of Botany
and Bakuriani Alpine Botanical Garden, Ilia State University, Tbilisi, Georgia
Herbario Nacionál de Bolivia, Universidad Mayor de San Andrés, La Paz, Bolivia
Maroof Ali Turi College of Life Science, Anhui Normal University, Wuhu, China
xxiii
Part I
Regions
Ethnobotany of Mountain Regions: Africa
Rainer W. Bussmann, Narel Y. Paniagua-Zambrana, and

Grace N. Njoroge
Introduction
Geology, Landforms and Soil Conditions
Africa is an old continent. Nowhere else is so much of the surface covered by

basement rocks, and the larger part of the geology of Africa is indeed the geology of
pre-cambrian material (Petters 1991). Large areas, so-called cratons, have remained
essentially unchanged since the early Proterozoic (~2000 MYA). The mobile belts,
i.e., the huge basins and swells between these cratons, are composed of equally old
rocks, but have been subject to deformation and partly to metamorphoses, mainly in
the Pan-African orogenesis (700–500 MYA). Today, relief differences are relatively
small in these areas, so the continent can be described as a large, uneven plateau,
which is tilted to the northwest. Extensive erosion surfaces covered by ancient,
heavily leached soils are more widespread in Africa than on any other continent, and
the huge relatively flat Savannahs are well known. Inselbergs are very characteristic
remnants of the older surfaces in these plains, but far too low to merit treatment here.
R. W. Bussmann (*)
Department of Ethnobotany, Institute of Botany and Bakuriani Alpine Botanical Garden, Ilia State
University, Tbilisi, Georgia
e-mail: rainer.bussmann@iliauni.edu.ge
N. Y. Paniagua-Zambrana
G. N. Njoroge
Department of Botany, Jomo Kenyatta University of Agriculture and Technology, Nairobi, Kenya
© Springer Nature Switzerland AG 2021 3

R. W. Bussmann (ed.), Ethnobotany of the Mountain Regions of Africa, Ethnobotany of
Mountain Regions, https://doi.org/10.1007/978-3-030-38386-2_1
4 R. W. Bussmann et al.
Fig. 1 Mawenzi (left) and Kilimanjaro (right), from Amboseli National Park, Kenya. (Photo
R.W. Bussmann)
Equally well known as the flat surfaces are, however, some much more recent
geological features, namely, the huge, isolated volcanoes of the tropical part of the
country, and the high fault mountains in the Atlas region and in South Africa. In these
three regions, true high-altitude environments are found, with several ranges reaching
well above 4000 m. These are almost exclusively located in the tropical parts, with
Mt. Kilimanjaro (5985 m, Fig. 1), Mt. Kenya (5198 m, Fig. 2), and Ruwenzori (5109 m)
being the highest sites. Several other mountains reach above the tree line, with the most
extensive areas being in Ethiopia. However, the High Atlas in Morocco is also high
enough to support truly alpine vegetation (4165 m).
Morocco occupies an intermediate position between the tectonic domains of
southern Europe and Africa. The evolution is complex, and the atlas region was
formed in several successive uplifting events. The south-westernmost part, the Anti-
Atlas contains Precambrian material, but was largely formed in the late Proterozoic.
Uplift occurred in Hercynian times in close correspondence to some mountains in
Europe, and again mainly in late Cretaceous tectonic movements that formed the
several chains of the High Atlas and the Saharan Atlas. The northern Riff and the Tell
Atlas in northern Algeria are the southernmost outposts of the Alpine orogen, but
recent Tertiary uplift affected also the older ranges, notably the Saharan Atlas. Most
of the material is acidic, but several ranges display calcareous rocks, like the central
High Atlas, part of the Saharan Atlas, and the Rif. Only the High Atlas reaches above
3000 m, while the neighboring ranges hardly climb above 2000 m.
The central Saharan mountains Hoggar and Tibesti rest on the relatively stable
West African craton. Part of the Tibesti is of ancient, Proterozoic origin, but there has
Ethnobotany of Mountain Regions: Africa 5
Fig. 2 Peak region of Mt. Kenya, with dense Dendrosenecio keniodendron, Central Province,
Kenya. (Photo R.W. Bussmann)
been extensive recent volcanism. Both ranges are part of a chain of volcanoes of
Cenozoic age that extends southwards to the central African Mt. Cameroon and
eastwards to the East African Rift Valley System.
Mt. Cameroon is a huge single structure of largely Tertiary origin but has been active
ever since with more than 10 major events in the twentieth century, some as recent as
1982 and 2000 (Ndam et al. 2001). The series of volcanoes starting at Mt. Cameroon has
been described to continue up to the Jebel Marra in central Sudan. This is a massif rather
than a single volcano, which has been active from the Miocene to the early Holocene
(Miehe 1988). Lavas are predominantly basaltic in all these ranges.
Further eastwards, the huge volcanic plateau of Ethiopia rests on ancient material
but was largely formed in the tectonic processes during the formation of the East
African Rift Valley system. The EARS is the prime example for a continental rift,
and its huge structure extends over more than 6000 km from northern Mozambique
along all of eastern Africa and through the Red Sea. Rifting started in the Cretaceous
but has intensified in the Tertiary and is most likely a consequence of tensional forces
driving the two escarpments apart. Volcanism has been intensive and complex,
following the two main branches, the Albertine Rift in the west, and the Gregory
Rift in the east (Schlüter 1997).
The Ethiopian Rift is an extension of the latter, mainly formed by basaltic
magmas in the Oligocene to Miocene, and reaching above 4000 m at several sites,
notably in Simen Mts. (Fig. 3) in the northern part of Ethiopia and the Bale Mts.
(Fig. 4) in the southwestern part. Relatively deep and dry basins from the isolated
volcanoes in east Africa proper separate this group of well-connected ranges.
Fig. 3 Simien Mountains landscape, Ethiopia. (Photo R.W. Bussmann)
Fig. 4 Bale Mountains, Southern Ethiopia. (Photo R.W. Bussmann)

Most of these are relatively young and of Pliocene to Quaternary age. This refers
to mountains such as the Aberdares, which were formed from 3 MYA onwards;
Mt. Kenya was formed between 2 and 3 MYA (Heinrichs 2001); Mt. Kilimanjaro is
even younger at 1–2 and Mt. Meru appears to be of similar age as are the Aberdare
Mts. Lavas on the huge shields are predominantly alkaline or basaltic, but interme-
diate lavas formed several stratovolcanoes.
The mineral composition is similar for Mt. Elgon, which is however an exception
with respect to its age. Mt. Elgon lies at the border of Uganda and Kenya and is thus not
strictly located in the Rift Valley. Here volcanism commenced some 20 Mio years ago
and lasted for about 10 Mio years. The neighboring Cherangani range is an ancient
block of largely metamorphic rocks but has been lifted to its present height in the
Miocene. These two ranges have been high mountains, long before their neighbors in
the eastern part of Africa, and also those in central Africa had attained high altitudes.
The Ruwenzori is a huge pre-Cambrian block, but has been lifted to alpine heights
later than 2 MYA. The Kahuzi-Biéga west of Lake Kivu has apparently experienced a
similar evolution (Fischer 1996). The other high mountains along the Albertine Rift
Valley are exclusively of volcanic origin, with volcanism commencing as early as
some 13 MYA in the Virungas. However, most of the activity occurred in the last
2 Mio. years and continued to the present, as was so strikingly demonstrated at the city
of Goma, which was largely destroyed during a catastrophic lava outpour in 2001.
These young mountains are in striking contrast to the truly ancient Eastern Arc
Mountains in Tanzania and northern Kenya. They are composed of granulite gneiss
complexes and have been mountains during for over 25 Mio years. However, they
barely reach 3000 m and lack an alpine belt typical for the other mountains discussed
here.
Faulting during the formation of the Rift Valley has formed much of the present
tropical part of African. Beyond Mozambique, mountains in the southernmost part
of the continent are fault mountains like the northwestern Atlas system
The lower slopes of the volcanic ranges have been subject to intensive weathering
and leaching, resulting in deep reddish soil profiles, mainly Acri- and Nitisols,
occasionally Ferralsols. The dominant soil types of the upper montane belt are,
however, well-developed Andosols. The Andosols have moderately acidic soil pH
values and are exceedingly fertile and explain the intensive agriculture on the tropical
mountains. The alpine regions are partly covered by Andosols, notably on the younger
volcanoes, partly by various weakly developed soils on glacial deposits such as
Regosols and Rankers. Very widespread are various Histosols that result from slow
decomposition of organic matter due to coldness and stagnant water. They are
characterized by low pH values (<4 to 5.5) and generally low nutrient availability.
Vegetation Belts in African Mountains
Atlas
With respect to their geologic origin, mountains in northwest Africa are closely
related to European mountains like the Alps. Correspondingly, climate, flora, and
vegetation display strong similarities to Mediterranean mountains like the Sierra

Nevada in Spain and mediate to the arid sites of central northern Africa.
There are three principal complexes, namely, the High Atlas and its neighbors
Anti-Atlas and Central Atlas; the Rif and Tell Atlas in northern Morocco and
Algeria; and the Saharan Atlas in Algeria and Tunisia. These display large-scale
climatic gradients, with conditions getting progressively drier with distance to the
coast, so the Rif and Tell Atlas get most precipitation with annual totals above
750 mm in the northern foot zone. In the Moroccan Atlas, conditions get drier from
west to east. These differences among ranges are small in comparison to the large
differences between the northern and the south (-eastern) slopes within the mountain
ranges. The northern lowlands receive well above 300 mm, while the southern slopes
border the Saharan basin, where precipitation quickly drops to less than 100 mm in
the lowlands (Weischet and Endlicher 2000).
Montane regions clearly receive more than lowlands with a precipitation maxi-
mum at 2000–3000 m elevation. This is around 750 mm in the High Atlas, and above
500 mm in all regions except for parts of the Saharan Atlas. The Rif and the Tell
Atlas receive annual totals of above 1000 mm, which rise occasionally up to
2000 mm. Rainfalls occur almost exclusively in winter, so the summit regions are
at least seasonally covered by snow, which might last until the following year in
moist conditions.
Thus, moisture availability and the usual altitudinal temperature lapse rate control
most of the vegetation patterns, but these are modified by edaphic components. Like
in the Alps, calcareous and acidic bedrocks intersect, so vicariating plant communi-
ties are found depending on the geological background. This analogy is however
limited, since the extremely harsh climatic conditions with a pronounced seasonality
and the general dryness of the summit region induce excessive weathering. Thus, the
uppermost regions are covered with scree, which move in the frequent freeze-thaw
cycles. Soils are weakly developed, and conditions resemble those of dry ranges in
the Sahara or even Central Asia, with an open and rather sparse vegetation cover
completely different from the Alps.
Human impact has been tremendous in the entire region. Agriculture and livestock
keeping were introduced some 5000 BP, and, like the entire Mediterranean region, most
of the natural vegetation has been replaced by various secondary plant communities,
rendering inferences of the potential character of the landscapes often different.
The Moroccan Atlas

Botanical exploration the area started equally late as in the seemingly much more
remote tropical parts of Africa and started in the 1870s. It has, however, been intense,
and the general patterns are known since the middle of the last century (Rauh 1952).
The High Atlas displays the typical situation. The northern foot zones are covered by
open shrublands with Zizyphus ssp., Lotus and notably Acacia gummifera up to
900 m, where the dwarf palm Chamaerops humilis demarcates the transition to forest
vegetation. Tetraclinis articulata forests are the natural community up to 1400 m,
from where they are replaced by various communities dominated by Quercus ilex.
Stands are dense and relatively shady with few companions as Cistus laurifolius on
acidic, and Buxus sempervirens on calcareous bedrocks. Much of these forests have
been replaced by various scrub communities with Pistacia lentiscus and Juniperus
phoenicea in the lower regions, and by J. thurifera in the upper montane belt.
Impressive Cedar (Cedrus atlantica) stands built a special forest community
restricted to the moistest parts of the Atlas, but are more extensive in the Rif and
in the Tell Atlas. Above 2800 m, J. thurifera forests dominate up to the tree line in
some 3100 m. Unlike in other areas of Africa, progressively diminishing and finally
shrubby growth of the dominant species do apparently not characterize the tree line
ecotone. Instead, Junipers retain their single-stemmed growth up to the tree line, with
stands opening progressively and distance between trees increasing.
Thorny cushion plants characterize the lower alpine region up to 3600 m. Cush-
ions grow 0.5–1.0 m in height and have thick, often several meters long root. The
dominant species are the widespread Alyssum spinosum, Bupleurum spinosum, joint
by the flat thorny cushions of Arenaria pungens in upper elevations. In contrast,
Erinacea pungens and Vellea mairei are restricted to calcareous bedrocks. Cushions
prefer weakly inclined to level slopes where they can build almost closed stands, but
have also benefited from anthropo-zoogenic human impact on the upper montane
Juniper forest, where they from secondary stands on disturbed sites.
Moist sites in the lower alpine zone near small brooks are the habitat of communities
called "Pozzines". These are meadows, physiognomically similar to alpine meadows in
Europe, and thus colorfully flowering in summer. However, they are of very limited
spatial extent, and are restricted to soils with impeded drainage over acidic bedrocks.
Substrate movement in the upper alpine region becomes increasingly strong
above 3300–3600. The cushions become more and more restricted to the few level
sites, and plants truly adapted to scree slopes and rock crevices take over. Again, we
find vicariant taxa on acidic and calcareous substrates, like Viola dyris var. orientalis
on calcareous rocks and Viola dyris var. typica on acidic bedrocks. The latter is an
endemic taxon of the Atlas, other examples are Linaria lurida, L. heterophylla ssp.
galioides and Vicia glauca var. rerayenssis, all growing on scree slopes. The relative
importance of endemic species increases with altitude in the Atlas, and is around
30% in the lower alpine zone, while it reaches some 75% in the upper alpine and
summit regions. There is no truly nival belt and plants occur up to the summit region.
The zonation on the southern slopes is somewhat different because of the greater
aridity. The foot zone is covered by succulent Euphorbia species or open shrublands
like on the northern slopes up to 1500. Juniperus phoenicea partly replaces Tetra-
clinis articulata and the belt of Quercus ilex forests is smaller and widely replaced by
open Juniper scrub. Cedrus forests are entirely absent. The timberline is formed by
J. thurifera as on the northern slopes, and the alpine vegetation is similar as well.
The Rif and the Tell Atlas

The Rif reaches barely 2500 m and the Tell Atlas is even lower, but affinities to the
Moroccan Atlas are strong in the montane belt (Knapp 1973), so the description is
kept short. The most widespread montane forest communities are also built by
Quercus ilex, which forms dense and rather shady stands. Quercus suber forms
extensive forests at the relatively moist coast of Algeria and Tunisia up to some
1000 m asl., but small stands of the Oak are also found in Morocco. Like Quercus
coccifera, it has rather high moisture demands and is restricted to the coastal regions
below 1200 m. Much of the areas have been cleared for agriculture, especially Olea
europea plantations. The olive forms (semi-)natural stands with Pistacia lentiscus in
drier parts of the lowlands, which might however be secondary in nature. Aleppo
Pine (Pinus halepenis), often mixed with Quercus ilex, dominates much of the lower
montane zone at 500–1500 m in the Rif and Tell Atlas, but is also occasionally found
in the Moroccan Atlas. Cedrus atlantica forest is common in the moister ranges
above 1500 m. However, the moistest sites there are covered by deciduous oak
forests (e.g., Q. faginea) and coniferous forests with Abies maroccana and
A. numidica. Juniperus phoenicea forests in the drier parts of the montane belt
have suffered heavily from anthropo-zoogenic disturbance but occur up to 2400 m
under natural conditions. Replacement communities include mixed scrub with
various Cistaceae, Fabaceae and Ericaceae species in the lower montane belt, and
shrublands with the dwarf palm Chamaerops humilis and Rosmarinus officinalis in
the montane belt.
There is no truly alpine vegetation in the Rif and in the Tell Atlas simply because
the ranges are too low.
The Saharan Atlas in Northern Algeria
The Saharan Atlas forms a second chain leeward of the Tell Atlas and is thus
relatively dry. Much of the natural vegetation has been replaced by widespread
steppes with Stipa tenacissima, that is an important species of secondary grasslands
in the western Mediterranean region. It is accompanied by Lygeum spartum or
Artemisia herbae-alba, mostly on dense or compacted soils. The natural montane
vegetation closely resembles the situation in the Moroccan Atlas described above,
notably in the western part of the country near the inter-state boundary. Communities
with higher moisture demands are almost exclusively absent, and forest with
Quercus ilex and occasionally Pinus halepensis would be the zonal vegetation of
much of the northern slopes. The southern slopes are influenced by the arid Saharan
climate and dry vegetation types with succulents climb up well into the montane belt.
Where the mountains are high enough, alpine vegetation types occur, which are also
similar to the High Atlas. Thorny cushions are relatively common, as are scree
communities, but communities of moist rock crevices are impoverished and
Pozzines are entirely absent. The alpine vegetation shows the typical differentiation
of vicariating species depending on the presence of acidic or calcareous bedrocks as
was described for the High Atlas.
Dry Mountains in Central Sudan: The Jebel Marra
The Jebel Marra is a highly isolated volcanic massif in the center of Africa, more
than 1500 km away from the nearest coast and more than 1000 km distant to the next
mountain, with the Simen Mts. in northern Ethiopia being the most “closely” related
range. The Jebel Marra emerges from a pedi-plain in 1000–1200 m, with a semiarid
climate with down to 280 mm of annual precipitation (Miehe 1988). The mountain
slopes receive more rain, and the southwestern side is clearly moister than the
eastern. The belt of maximum precipitation is situated at around 2000 m asl.,
where totals can be as high as 1800 mm in extreme years, but 800 mm appear to
reflect the average conditions better. For the summit region (max. 2976 m Times
Atlas 3071) 600 mm have been estimated.
The vegetation shows correspondingly clear altitudinal gradients. Drought decid-
uous woodlands cover the piedmont and Acacia scrub, interspersed with lowland
gallery forests along watercourses and in moist ravines and valleys, mainly in the
southwest. These can be regarded as remote outposts of tropical evergreen forest.
Typical species include evergreen Ficus ingens, F. thonningii, F. sur, Syzygium
guineense and Trema orientalis, indicating affinities to tropical montane forest
described below.
The dominant vegetation type in the submontane belt are, however, communities
with Combretum molle and Boswellia papyrifera, which are replaced by wooded
grasslands with Entada abyssinica and Acacia albida as a transitional belt to the
montane communities.
The transition from these submontane lowlands to the montane, sparsely wooded
grasslands occurs at some 2000 m. The principal tree species is Olea laperrinei, which
forms riverine forests above 1900 m with Ficus palmata. The olive is also found in the
Saharan mountains and indicates the relationships to Hoggar and Tibesti. However,
the dominant vegetation are grasslands with Andropogon-Hyparrhenia, and Themeda
triandra-Conyza hochstetteri. Scattered trees of O. laperrinei can be found on scree
slopes and in the grasslands, indicating that at least parts of these grasslands are
suitable for tree growth. The upper montane grasslands above 2800 m are composed
of Pentaschistis species and Festuca abyssinica. Occasionally, remnants of ericaceous
vegetation, notably E. arborea, occur in the upper montane region. Ericaceous
vegetation dominates the timberline ecotone in much of tropical Africa, but the
Jebel Marra is not high enough for a true alpine belt.
Thus, with respect to the temperature and moisture availability most of the
montane belt should be suitable for dry Afromontane forests dominated by
O. laperrinei, which have presumably been cleared by large-scale woodcutting,
grazing, and fire. This would clearly indicate parallels to the tropical mountains
described in the following chapter.
Mount Kenya: A Typical Example for the Altitudinal Zonation

of Tropical African Mountain Vegetation
Mt. Kenya (Fig. 5a, b) is intermediate among the mountains in tropical Africa in
terms of geographical position and climate. Research there has started early in the
last century, so the available data are relatively comprehensive. Hence, we will
discuss the vegetation zonation of Mt. Kenya in some detail, while descriptions for
the other mountains are kept shorter and focus on the differences to the typical
situation in central Kenya.
Mount Kenya, located about 180 km north of Nairobi on the eastern side of the
Great Rift Valley, is a widely isolated volcano. The base diameter is about 120 km;
the northern foothills reach the equator. With its main peaks Batian (5198 m) and
Nelion (5188 m), Mt. Kenya is, after Kilimanjaro (Kibo 5899 m), the second highest
mountain in Africa. It is broadly cone-shaped with deeply incised, in the upper parts
U-shaped valleys, indicating extensive former glaciations. Most parts of Mt. Kenya
are still covered with forests up to approximately 3400 m in the South and 3000 m in
the North, where a sharp boundary separates the forest from the lower alpine zone.
Shape and location of the upper tree line has been severely influenced by fires. The
present lower timberline is a result of extensive forest clearance and agriculture with
cultivation reaching up to 1800 m on the southern, up to 2400 m on the eastern and
western, and nearly up to 2900 m on the northern slopes. Without human impact, the
mountain would be almost completely surrounded by dense forests.
As an important water catchment, Mt. Kenya contributes mainly to the Ewaso
Nyiro and Tana River drainage systems and is therefore of outmost importance for
the water supply of about 50% of the Kenyan population. Moreover, since other
Fig. 5 (continued)
Fig. 5 (a) Vegetation zonation on Mt. Kenya – transect from W-E. (Bussmann 2006). (b)
Vegetation zonation on Mt. Kenya – transect from NW-SE. (Bussmann 2006)
power stations did not fulfill the expectations, hydropower from these rivers con-
tinues to generate more than half of the country's production of electrical energy
(Berger 1989; Decurtins 1985, 1992; Leibundgut 1986). Densely populated farming
areas surround the whole mountain, and large parts of the forest suffered heavily
from encroachment in the last decades. The vertical zonation of the mountain has
already been described in the early twentieth century (Engler 1895; Allaud and
Jeannel 1914), but the first comprehensive accounts are from the 1950s (Hedberg
1951).
Supratropical Evergreen Montane Forests
The lowermost areas on the eastern and northeastern slopes from 1200 to 1500 m are
covered with evergreen mountain forests, which closely resemble forest types of
central and western tropical Africa. Newtonia buchananii, Lovoa swynnertonii and
Fig. 6 Dense crowns of

supratropical evergreen
montane forests, Mt. Kenya,
Kenya. (Photo
R.W. Bussmann)
Chrysophyllum gorungosanum are prominent tree species. These have survived only
in comparatively small remnants, while the principal forest formation are typical
Camphor forests, with Ocotea usambarensis, Aningeria adolfi-friederici and
Syzygium guineensis being the most important canopy species. These evergreen
submontane forests (Bussmann 1994, 2001; Bussmann and Beck 1995a, b, 1999;
Fig. 6) are two-storied, with huge specimens of the African Camphor tree, Ocotea
usambarensis in the upper canopy, and Xymalos monospora, Lasianthus
kilimandscharicus, Pauridiantha holstii, and Psychotria orophila in the lower tree
layer and the often-dense shrub stratum. Further characteristic species in the upper
tree layer are Strombosia scheffleri and Apodytes dimidiata in the upper tree layer,
while Tabernaemontana stapfiana, Ochna insculpta, Macaranga kilimandscharica,
and Peddiea fischeri form low trees and shrubs. Asplenium sandersonii, A. elliottii,
and Panicum calvum occur in the herb layer, together with Piper capense,
Oplismenus hirtellus, Plectranthus luteus, Begonia meyeri-johannis, and the ferns
Dryopteris kilemensis and Blotiella stipitata. Cyphostemma kilimandscharicum is a
common climber, and abundant fern epiphytes include Elaphoglossum lastii, Tri-
chomanes borbonica, Asplenium theciferum, and Oleandra distenta.
On the lowermost eastern slopes of Mt. Kenya, forests with only one closed tree
layer are found, dominated by Newtonia buchananii and many, often very tall palms
(Phoenix reclinata), which give these areas an appearance like an Andean cloud
forest Fig. 7). Once, these forests covered large tracts of land especially on the
Fig. 7 Moist submontane forests, Mt. Kenya, Kenya. (Photo R.W. Bussmann)
eastern slopes of Mt. Kenya and formed the lowermost submontane forest belt in
transition to the savannah lowlands, but these forests have long ago been cleared for
cultivation.
On the western side of Mt. Kenya, evergreen submontane semi-deciduous forests,
where drought resistant species like Calodendron capense or Croton megalocarpus
are common, follow woodlands dominated by Acacia drepanolobium from 2000 to
2300 m.
On the northern part of Mt. Kenya, the woodlands lead directly into very drought-
resistant xeromorphic forests, which are almost entirely dominated by Pencil Cedar
(Juniperus procera) and Wild Olive (Olea europaea subsp. africana). A Bamboo
belt is absent there, and even the narrow cloud forest belt is heavily interspersed with
Cedar. Several authors mentioned a so-called forest gap on the drier northern slopes
of the mountain (e.g., Hutchins 1909; White 1950). Whether this gap is natural or
man-made has been a matter of debate. According to the statements of long-term
residents of the area, clearing the forest for farming purposes has widened the gap. In
the late 1970s, the Kenyan government for farmland excised some areas of the
Mt. Kenya Forest Reserve. In many smallholder fields (shambas) of this area, forest
trees or remnants of trees are still present, indicating clearly the former extension of
the forest belt. Climatic conditions are less probable to be responsible for the gap
since in an even drier area exactly north of the gap on Mount Kenya, the Ngare
Ndare forest with extensive stands of Juniperus procera is found. Therefore, there
are no reasons to assume a natural reason for the existence of the forest gap. Most
probably, a formerly closed forest belt must be concluded.
Riverine forests and a variety of very different forests can be also found,
indicating affinities to the mountain forest communities of Central and Western
Africa. Lovoa swynnertonii, a very tall and dominant canopy tree is a typical species,
growing together with the shrubs Rawsonia lucida, and with Heinsenia
diervilleoides and Rinorea convallarioides. In many places, Uvariodendron
anisatum, with its beautiful and very fragrant white leaves, grows abundantly in
the lower tree stratum and in the shrub layer.
Pure Camphor forests are found mainly on the very wet southeastern and southern
slopes of Mt. Kenya on altitudes between 1550 and 2550 m. They grow on humic
Niti- and Acrisols and receive an annual rainfall of 1500–2500 mm. Moist forests in
the lower and middle submontane region of southeastern Mt. Kenya are dominated
by the evergreen species Syzygium guineense (Myrtaceae) and Aningeria adolfi-
friederici (Sapotaceae), reaching a height of up to 50 m. Furthermore, the shrubs
Drypetes gerrardii (Euphorbiaceae) and Allophyllus cuneatus (Sapindaceae), as well
as the lianas Adenia gummifera (Passifloraceae) and Jaundea pinnata
(Connaraceae) are characteristic species. The number of epiphytic mosses is limited,
but a wide variety of epiphyllous and epiphytic liverworts and lichens occur.
Most stands suffered from heavy exploitation, and have been replaced by sec-
ondary vegetation types, leaving untouched evergreen forests only in steep ravines
or in remote areas, which are difficult to access. Macaranga kilimandscharica
dominates the canopy of these secondary forests, and often forms dense pure stands.
It is a very fast-growing species suppressing the regeneration of other trees.
Orotropical Montane Forest: Evergreen Broad-Leaved and Evergreen

Xeromorphic Montane Forests
The cedar forests (Juniperus procera) grow on humic Acrisols (Speck 1986) at
rainfall totals between 700 and 1500 mm (Fig. 8). They represent the typical
vegetation of the altitudinal range between 2500 and 2950 m of the drier exposures
(Bussmann 1994, 2001, Bussmann and Beck 1995a, c, 1999; Bussmann and Lange
1999). Evergreen-broadleaved forests cover the moister southern and southeastern
slopes, at lower altitudes between 2150 and 2650 m. Juniperus procera itself is rare
or completely absent, due to heavy logging and suppressed regeneration in the
closed stands. The dominant tree species in these two-storied stands are Cassipourea
malosana (Rhizophoraceae) and Olea capensis subsp. hochstetteri, the East African
Olive; both forming the upper canopy. Lepidotrichilia volkensii in the lower tree and
shrub strata, and Ilex mitis, the African Holly are further important woody species.
Cedar forest, in contrast, is rather open and one or two-storied, with J. procera
growing about 50 m tall under favorable conditions. Other important tree species,
mainly of the lower canopy, are the Wild Olive (Olea europaea subsp. africana) and
Podocarpus latifolius. Mosaics of dense grass layers with Stipa dregeana and
Brachypodium flexum, interspersed with herbs (Sanicula elata, Isoglossa gregorii,
and Achyranthes aspera), and low shrubs (e.g., Berberis holstii) are also character-
istic of these open cedar forests.
Fig. 8 Dry Juniperus-Olea

forests. Note fire scar in
middle. Mt. Kenya, Kenya.
(Photo R.W. Bussmann)
Orotropical Bamboo Forest
The current scientific name of East African Bamboo is Oldeania alpina. However,
for reasons of comparability, the much better-known synonym "Sinarundinaria
alpina" is used here. Dense forests dominated by the African Bamboo
(Sinarundinaria alpina), which are found on various East African mountains,
constitute a very distinct formation (Bussmann 1994, 1997, 2001; Bussmann and
Beck 1995a, 1999, Bytebier and Bussmann 2000). Sinarundinaria alpina is the
overwhelmingly dominant species in all strata, but Impatiens hoehnelii, Pseudo-
carum eminii as well as Selaginella kraussiana, Cyperus dereilema, and Anthriscus
sylvestris are common companions. Bamboo forests are restricted to a relatively
narrow ecological range, mainly depending on soil temperatures of 10–15 °C and the
presence of very deep volcanic soils, namely, humic Andosols (Fig. 9). On
Mt. Kenya, they are found in an almost closed belt around the entire mountain,
interrupted only on the dry northern slopes. Stands are somewhat more extensive on
the very wet southern parts.
On steep slopes of the western, northwestern, and eastern side of Mt. Kenya, in
areas with lower rainfall and higher soil temperatures, often huge individual trees
protrude from the dense Bamboo layer in some 10–15 m height. The dominant
species of the tree stratum is always Podocarpus latifolius, which is also common in
the surrounding montane fores (Fig. 10). Towards the hygric limits of the Bamboo,
Fig. 9 Sinarundinaria alpina

forming dense forest,
Mt. Kenya, Kenya. (Photo
R.W. Bussmann)
Fig. 10 Mixed bamboo-

Podocarpus latifolius forest,
Mt. Kenya. (Photo
R.W. Bussmann)
mainly on the dry northwestern parts of the mountain, Juniperus procera and Olea
europaea subsp. africana are found as characteristic members of the tree stratum.
Trails and heavily trampled resting places of big game occur everywhere in these
forests and seem to facilitate regeneration of trees in the otherwise extremely shady
stands of Sinarundinaria alpina.
Orotropical Cloud Forest

Two forest types, namely, tall and malacophyllous forest with Hagenia abyssinica
occupy this vegetation zone, and lower stands built by various species with
sclerophyllous, “ericoid” leaves. The so-called ericaceous belt forms the transitions
zone towards truly afroalpine vegetation within the timberline ecotone. In contrast,
the malacophyllous forests extend from 2900 to 3300 m a.s.l., where low clouds and
mist are frequent. The dominant tree is the Kosso tree, Hagenia abyssinica
(Rosaceae), often accompanied by the Giant St. John’s Worth, Hypericum
revolutum. Bothriocline fusca is a common shrub, while Polygonum afromontanum,
Stephania abyssinica, Cinereria deltoidea, and Carduus afromontanus are common
herbs. Pure Hagenia-Hypericum forests with a low, dense shrub layer of Hypericum
are characteristic for the wetter southeastern to western slopes of Mt. Kenya. Here,
important herbaceous companions are Luzula johnstonii, the large Lobelia
bambuseti, Rubus frieseorum, Agrostis schimperiana, Poa schimperiana,
Helictotrichon milanjeanum, and one of the few succulents, Uebelinia rotundifolia.
In the drier northern part of Mt. Kenya, the Hagenia-dominated forests are
replaced by mixed forests with Afrocarpus gracilior and broad-leaved species,
clearly differentiated by the high frequency of Juniperus procera, Olea europaea,
Nuxia congesta, and especially Rapanea melanophloeos. Hagenia abyssinica is only
a codominant canopy species and Hypericum revolutum is mostly absent.
From some 3300 m upwards, open ericaceous communities gradually replace
Hagenia forests. They grow at the upper limit of forests, but the timberline has been
lowered by several hundred meters due to fires and undulates now at 3100–3300 m. True
Erica excelsa forests are poorly developed on Mt. Kenya, and most often the ericaceous
belt is formed by remnant stands of ericaceous scrub with Erica trimera (formerly
Philippia, Oliver 1987), E. arborea, Conyza vernonioides, and the astonishingly “eri-
coid” Asteraceae Stoebe kilimandscharica (Rehder et al. 1981, 1988). The latter,
together with Protea kilimanscharica often indicates regular disturbance by high altitude
fires. Fires are usually discontinuous, so the scrub is interspersed with various grassland
types that are already afroalpine. Thus, ericaceous and afroalpine communities generally
form patchy mosaics and transition communities rather than clear altitudinal belts.
Altotropical Grasslands and Moorlands

In tropical Africa, the altitudinal belt above the timberline has been termed “afroalpine”
(Hauman 1955). Communities intersect with ericaceous vegetation and are mainly
dominated by tussock grasses and stands of Giant Rosette Plants. Fortunately, a
vegetation map is available, so the spatial distribution of communities is well known
(Rehder et al. 1988; Beck et al. 1990). “Open Moorlands” are particularly extensive on
the western slopes of Mt. Kenya at altitudes from 3400 to 3800 m, often found at
clearings of ericaceous vegetation. The dominant species are grasses, namely, the large
sedge Carex monostachya, interspersed with tussocks of Festuca pilgeri in drier places.
Most characteristic are the large, cabbage-like ground rosettes of Dendrosenecio bras-
sica (Fig. 11) and Lobelia keniensis Fig. 12). The Giant Rosette "tree" Dendrosenecio
johnstonii subsp. battiscombei (Fig. 13) is common in the lower moorlands as well as in
remnant ericaceous stands.
Fig. 11 Dendrosenecio brassica grasslands on Mt. Kenya, Kenya. (Photo R.W. Bussmann)
Fig. 12 Lobelia keniensis grasslands on Mt. Kenya, Kenya. (Photo R.W. Bussmann)
Fig. 13 Dendrosenecio
johnstonii subsp. battiscombei
grasslands on Mt. Kenya,
Kenya. (Photo
R.W. Bussmann)
The “Upper Moorland Zone,” extending from about 3800 m upwards as high as
4550 m comprises two very characteristic vegetation units. Where moisture avail-
ability is sufficient, “Upper Alpine Wetlands” are formed by Dendrosenecio brassica
and Lobelia deckenii ssp. keniensis. In contrast to lower altitudes, sclerophyllous
dwarf shrubs like Alchemilla johnstonii, Alchemilla argyrophylla subsp.
argyrophylla and Helichrysum brownei often replace grasses. Sedges are less impor-
tant than in the lower moorlands.
The typical plant community above 4300 m is the “Dendrosenecio woodland,” in
which D. brassica is replaced by the Giant Groundsel, Dendrosenecio keniodendron
(Fig. 14), growing together with Lobelia telekii. From 4500 m onwards, the Giant
Groundsels recede, and an open layer of Festuca pilgeri, interspersed with Lobelia
telekii (Fig. 15), Festuca abyssinica, Carduus chamaecephalus, and Senecio
keniophytum extends to the subnival zone. Occasional specimens of vascular plants
where found up to the summit region in above 5000 m (Rehder et al. 1988).
In contrast to the situation on Kilimanjaro, where low precipitation creates a real
alpine desert, closed vegetation on Mt. Kenya is more limited due to freezing and
continuous solifluction, although moisture availability probably has some influence
(Beck 1994).
Fig. 14 Dendrosenecio keniodendron on Mt. Kenya, Kenya. (Photo R.W. Bussmann)
Fig. 15 Lobelia telekii on

Mt. Kenya, Kenya. (Photo
R.W. Bussmann)
Mt. Kenya’s Neighbors: Mt. Kilimanjaro, Mt. Meru, Aberdare Mts.,

and Mt. Elgon
Although they are separated by hundreds of kilometers of lowlands, the neighboring

mountains show a vegetation zonation very similar to Mt. Kenya. These striking
similarities have long been known to phytogeographers (Hedberg 1951, 1964) and
have let to the general classification scheme that provided the base for the previous
chapter. The vegetation belts are very distinct, as are differences between sides of the
mountains, where drought-resistant forest types on their northern escarpments
replace very humid elements on their southern and southwestern slopes.
Exposure differences are particularly pronounced on Mt. Kilimanjaro (Fig. 16).
The southern slopes are unusually moist with an annual precipitation of around
900 mm in the southern foothills, around 2000 mm at 1500 m asl. and well above
3000 mm between 2000 and 2300 m. The summit region is, in sharp contrast, an
alpine desert with well below 200 mm annual precipitation (Hemp 2002). Although
no exact measurements are available, the northern slopes are clearly drier as
Fig. 16 Vegetation zonation on Mt. Kilimanjaro (Bussmann 2006)

indicated on the available vegetation map and the accompanying descriptions (Hemp
2001). Below 1000 m, dry savannah surrounds the entire mountain, but on the
northern slopes, savannahs with Acacia drepanolobium and forest plantations
climb above 2000 m altitude, followed by a small belt of xeromorphic forests with
Cedar and Olive up to 2800 m.
In the moist south between 1000 and 1800 m, evergreen submontane deciduous
forests has largely been replaced by dense agriculture. Remnants of the lowermost
submontane forests are characterized by Olea europaea and at drier sites by
O. capensis, but from 1700 m upwards, lower montane evergreen Camphor forests
and other broadleaved forests dominated by Cassipourea malosana replace these. In
proper montane forests above 2000 m, Podocarpus latifolius becomes increasingly
common. The most striking feature of these forests is the abundance of epiphytes, in
particular Hymenophyllaceae and other fern groups, which give the stands the
appearance of a true elfin forest. On all sides of Kilimanjaro, dense Erica excelsa
forests occupy the belt between 2700–3100 m (South) and 2800–3400 m (North).
Lower down, in the central montane belt, such forests form replacement communi-
ties at disturbed sites (Hemp and Beck 2001). Cloud forests with Hagenia abyssinica
and Hypericum revolutum are restricted to special sites such as boulder streams and
moist valleys. Above 3100 m, Erica forests are gradually replaced by ericaceous
scrub mixed with altotropical moorlands, where subspecies of Dendrosenecio
johnstonii and Lobelia deckenii grow as vicariant taxa to Mt. Kenya (Knox 1993).
Above 3900 m, subnival Helichrysum scrub takes over up to some 4600 m, where it
becomes too dry for growth of vascular plants. The summit region is truly nival and
devote of vegetation.
The northern flanks of the neighboring Mt. Meru (Fig. 17) are covered by decid-
uous open woodlands, which are followed by a small band of cloud forest with
Hagenia abyssinica, Hypericum revolutum, and Gnidia glauca between 2500 and
3000 m (Hedberg 1955, 1964). Higher up, these are replaced by evergreen ericaceous
scrub up to 3700 m, which forms the transition to altotropical moorlands, where Carex
monostachya, Lobelia deckenii, and Dendrosenecio johnstonii dominate. On the
southern slopes of this mountain, however, evergreen montane xeromorphic forest
with Podocarpus latifolius, Cassipourea malosana, and Olea capensis grows at
altitudes from 1700 to 2300 m. These merge gradually into a distinct Bamboo belt
that is replaced by typical cloud forest at 2550–2700 m. The zonation of the uppermost
regions shows the same features as the northern escarpment.
The Aberdare Mts. form the next neighboring mountain range west of Mt. Kenya.
Climate and vegetation were comprehensively described by Schmitt 1991, who also
provides a vegetation map. Climatic conditions are similar to Mt. Kenya, with highest
precipitation in the southeast and a maximum in the montane belt of around 2200 mm.
The vegetation zonation is also largely similar, with Cassipurea malosana and Olea
capensis in the supratropical mountain forest. Above 2500 to 2700 m, forests with
Cedar, Podocarpus latifolius and riverine forests with Afrocrania volkensii dominate.
An almost closed belt of Bamboo, from which specimens of P. latifolius and Nuxia
congesta emerge, follows these. Above 2900 m, cloud forests with Hagenia abyssinica,
Hypericum revolutum, and Rapanea melanophloeos take over. Erica excelsa becomes
Fig. 17 Vegetation zonation on Mt. Meru (Bussmann 2006)
important towards the upper limit of montane forests on the main plateau in some
3100 m altitude. Thus, most of the upper region of the Aberdare Mts. is well in the
altitudinal range of ericaceous forests and scrub, so that grasslands are largely secondary
and promoted by fires as well as grazing by wild ungulates. The vegetation gives a very
patchy impression, and altitudinal belts can hardly be inferred except towards the
summits from 3600 m upwards. Ericaceous vegetation includes well-developed cloud
forest of E. excelsa, and scrub with E. trimera, Stoebe kilimanscharica, and Cliffortia
nitidula. The altotropical grasslands are dominated by Festuca pilgeri, Koeleria
capensis, the C4 grass Andropogon lima, and at moist sites sedges (Carex monostachya,
Cyperus kerstenii). Again, vicariant subspecies of the Dendrosenecio johnstonii and of
the Lobelia deckenii groups grow in the afroalpine zone.
Near the boundary to Uganda, Mt. Elgon and the Cherangani Mts. form the
easternmost part of these semi-humid mountain ranges. The Cheranganis are heavily
cultivated and grazed on all elevations, so there is hardly any natural montane forest left.
With some 3600 m altitude, they are not high enough for extensive natural grasslands, so
much of the present altotropical grasslands appear to be secondary (Mabberley 1975).
In contrast, Mt. Elgon is the largest solitary volcano with a full set of altitudinal
belts comparable to Mt. Kenya (Wesche 2002, vegetation maps provided by van
Heist 1994a; van Heist 1994b). The western and southwestern slopes are clearly
moister than the eastern part of the mountain, with a maximum of precipitation
between 2300 and 2700 m asl. (1500–2000 mm on the western slopes, 1200–1500
on the eastern). Hardly any natural vegetation has survived the intense agriculture
below 2200 and 2500 m, but higher up, large supratropical forests with scattered
Camphor, Aningeria adolfi-friederici, Podocarpus latifolius, and riverine Afrocrania
forests survived. The eastern side is covered by dry orotropical forest with
P. gracilior, Olea capensis, and scattered Cedars. The Bamboo belt is almost closed
except in the northernmost part. Cloud forests with Kosso and tree heather have
mainly survived in the western part. Above the present timberline, mosaics of
ericaceous vegetation with E. trimera and Stoebe kilimandscharica, Helichrysum
and Alchemilla dwarf scrub, and altotropical Festuca pilgeri grassland form the
vegetation up to some 3700 m. Above this, afroalpine grasslands, Helichrysum scrub
and Dendrosencio woodlands extend up to the summits. Mt. Elgon carries a separate
species within the Dendrosenecio johnstonii complex, with one subspecies occur-
ring up to 4000 and a second one between 4000 and 4300 m, resembling
D. keniodendron on Mt. Kenya.
The Interlacustrine Highlands: Ruwenzori, Virunga Volcanoes,

and Kahuzi-Biéga
The mountains at the western branch of the Rift Valley form a separate group due to
their perhumid rather than semi-humid climate. They border the huge rainforests of
the Congo basin, so their lower slopes are covered by luxurious tropical evergreen
forest with a high richness in species. Comprehensive vegetation surveys have been
produced for Kahuzi-Biéga (Fischer 1996, including land cover map), the eastern
half of the Ruwenzori range (Schmitt 1992; Osmaston 1996, land cover map by van
Heist 1999, glaciological map by Osmaston and Kaser 2001), and for parts of the
Virunga Mts. (Snowden 1933; Biedinger 1995; Karlowski 1995).
Although the eastern side is generally drier, the vegetation of Ruwenzori
(Fig. 18), as of Mt. Cameroon, shows no clear distinction between the humid
Western and drier Eastern side of the mountain, whereas on the Virunga Volcanoes
(Karisimbi Fig. 19 and Muhavura Fig. 20) exposition differences are pronounced.
The evergreen submontane forest belt of Ruwenzori stretches from 1000 to 1600 m.
Common and typical tree species are Ocotea usambarensis, Aningeria adolfi-
friederici, and Syzygium guineense. On Muhavura and Karisimbi, submontane forest
occupies both western and eastern slopes from 1300 to 2000m. At altitudes between
2000 and 2300 m evergreen supratropical mountain forests with Olea capensis,
Cassipourea malosana and Podocarpus latifolius replace lowland species in all
three regions. In striking contrast to East Africa, no Cedar (Juniperus procera) can
be found on any mountain in central Africa. Bamboo (Sinarundinaria alpina),
mostly in stands interspersed with P. latifolius, is a prominent feature on Ruwenzori
(from 2300 to 3000 m) and on the Virungas (2400–2600 m on Karisimbi and
stretching up as high as 2900 m on western Muhavura).
On Ruwenzori, evergreen Erica forest follows immediately after the Bamboo
belt and forms extensive stands at 3000–3900 m. The situation on the Virunga
Volcanoes is different. On Western Karisimbi (Fig. 5), a small fringe of evergreen
cloud forest with Hagenia abyssinica and Hypericum revolutum separates Bam-
boo and ericaceous belt between 2600 and 2700 m, from which ericaceous forests
Fig. 18 Vegetation zonation on Mt. Ruwenzori (Bussmann 2006)
Fig. 19 Vegetation zonation on Mt. Karisimbi, Virunga Volcanoes (Bussmann 2006)

Fig. 20 Vegetation zonation on Mt. Muhavura, Virunga Volcanoes (Bussmann 2006)
can be found up to 3800 m. The cloud forest belt is better developed on the humid
eastern slope, where it extends from 2600 to 3500 m, followed by ericaceous
scrub up to 3800 m. The drier Muhavura (Fig. 6) shows an exactly inverted cloud
forest distribution: on the more humid western slope, Hagenia prevails from
2900–3300 m, followed by the ericaceous vegetation up to 3700 m, whereas on
the dry eastern escarpment the cloud forest zone is completely replaced by
afroalpine grassland (Karlowski 1995).
Extensive altotropical moorlands always occupy the alpine zone of the Central
African Mountains. Tussocks of Festuca abyssinica and Carex runssoroensis, as
well as the giant rosette plants Lobelia wollastonii and Dendrosenecio johnstonii
occur on all three mountains, whereas Lobelia stuhlmannii grows only on the
Virungas. Due to its much higher elevation, upper alpine dwarf scrub is only
found on the uppermost parts of Ruwenzori, where Helichrysum stuhlmanii
forms isolated thickets. The Kahuzi-Biéga range is not high enough for a true
afroalpine zone, so the summit region is covered by ericaceous scrub and
Deschampsia flexuosa grassland. The latter indicates the regular presence of fires.
The Westernmost Outpost: Mt. Cameroon
The highlands in Cameroon form a widely isolated archipelago in western central

Africa. The highest peak is Mt. Cameroon (4095 m, Fig. 21), an active volcano that
had its most recent eruption in 1982. The climate is extraordinary moist, with up to
10000 mm annual precipitation in the foot zone and below 2000 mm in the summit
Fig. 21 Vegetation zonation on Mt. Cameroon (Bussmann 2006)
region (Cable and Cheek 1998, including vegetation map). Mean air temperature is
around 20°C at 900 m and estimated to be around 4°C in the summit region. Snow is
rare, even on top of Mt. Kilimanjaro. Soils in the mountains are mostly young and
therefore fertile, so large areas have been cleared for agriculture. About a third of
Mt. Cameroon is now under legal protection, and here natural vegetation has
survived partly down to the sea level (Cable and Cheek 1998). The small remnants
of lowland forests are rich in various species and have the highest number of
endemics of all vegetation belts (29 vs. 49 for the entire range). Pluviotropical
evergreen rain forest dominates the larger part of these altitudes, with tree species
like Schefflera mannii or Syzygium guineense frequently found in the canopy, and
Allophyllus bullatus forming a second layer. These forests merge gradually into
submontane evergreen forest from 800 m upwards. Species composition, however,
changes only slightly, being characterized by Syzygium staudtii, Schefflera
abyssinica, and still Allophyllus bullatus (Richards 1963; Hall 1973). From 2000
to 2600 m, orotropical cloud forest is the dominant forest type with species found on
all tropical African mountains like Nuxia congesta, Prunus africana, Xymalos
monospora, and Hypericum revolutum. These form the evergreen cloud forest,
together with Rapanea neurophylla. Bamboo is completely absent from
Mt. Cameroon but occurs other ranges in western central Africa. Hagenia abyssinica
does also not occur on Mt. Cameroon, although all other species show clear relation-
ships to the Hagenia-dominated upper montane forests common in other areas of
sub-Saharan Africa. At all altitudes, large parts of the forests have been replaced by
grasslands due to high volcanic activity and thus frequent fires. For the same reason,
scrub and thickets are frequently found all over the mountain, with Marantaceae
dominating below 2000 m and ericaceous species like Erica mannii, Agauria
salicifolia, and Myrica arborea forming a patchy ericaceous belt towards the
timberline ecotone. Dense altotropical tussock grasslands reach up to 3400 m, where

large Poaceae as Andropogon sp., Festuca abyssinica, Deschampsia mildbreadii,
and Loudetia camerunensis dominate, interspersed with small tree islands (Photo).
True giant rosette plants, Dendrosenecio or Lobelia species are absent from
Mt. Cameroon, though Crassocephalum mannii and Lobelia columnaris show
resemblances to this growth form. From 3400 m upwards to the summit region,
vegetation cover becomes more open and impoverished in species (Maitland 1932).
There are no clear vegetation differences with regard to slope exposition.
The Sudano-Ethiopian Mountains: Imatong, Simien, Bale
In striking contrast to the mountains in the strictly tropical part Africa, the Sudano-
Ethiopian mountains emerge from dry vegetation and semiarid surroundings. Thus,
the mountains of southern Sudan and Ethiopia are characterized by steep climatic
gradients and correspondingly, by clear altitudinal changes in the vegetation zona-
tion. All ranges are widely isolated from the eastern African mountains, with the
Imatong Mts. in southern Sudan being the least distant.
Their foot zone receives some 800 mm, while at 1900 m asl. some 2200 mm have
been recorded; the highest figure in Sudan. The lower slopes of the Imatong Mts.
(Fig. 22) are covered with lush evergreen submontane forest, where Ocotea
usambarensis, Olea welwitschii, and Chrysophyllum fulvum are common species (Jack-
son 1956). Above 2000 m, these forests merge into evergreen montane xeromorphic
forest with Podocarpus latifolius and Olea capensis. Above 2600 m, a mixture of
evergreen Bamboo and evergreen cloud forests with Hagenia abyssinica replaces these.
The highest zones, above 3000 m, are covered with dense thickets formed by Erica
arborea and Myrica salicifolia up to the summit region.
Fig. 22 Vegetation zonation in the Imatong Mts. (Bussmann 2006)

Fig. 23 Vegetation zonation of the Bale Mts. (Bussmann 2006)
The by far largest area of afromontane and afroalpine environments is found in

Ethiopia. This huge country displays notable differences in climate, and conse-
quently relatively moist ranges as the Bale Mts. in the southern part are found, as
well as extremely dry areas like the Simen Mts. in the north. The Bale Mountains
(Fig. 23) form the single largest afroalpine area, with a flora and vegetation being
transient between East Africa and the drier northern part of Ethiopia (Gashaw and
Fetene 1996; Weinert 1983). The southern foot zone emerges from dry Combretum-
Commiphora savannah at 390 mm annual precipitation, but the southern Harenna
escarpment receives around 850 mm at 3000 m asl. The central part of the range is a
huge plateau in some 4000 m asl., where conditions become progressively moister
towards the northern slope, which receives around 1000 mm (Miehe and Miehe
1994). This induces a clear zonation of very well pronounced vegetation belts. The
lower southern slopes are covered with dense evergreen submontane deciduous
forest from 1450 to 1900 m, where Afrocarpus gracilior, Filicium decipiens, and
Celtis africana as well as C. gomphophylla form an open canopy (Negatu and
Tadesse 1989; Bussmann 1997, Fig. 24). The undergrowth is often dominated by
very dense thickets of wild coffee (Coffea arabica). Mid-altitudes from 1900 to
2300 m are covered by evergreen submontane forest (with Ocotea kenyensis,
Aningeria adolfi-friederici, and Olea welwitschii as most characteristic tree species),
Fig. 24 Humid Afrocarpus gracilior, Filicium decipiens, and Celtis africana forest in Bale,
Ethiopia. (Photo R.W. Bussmann)
and thus resemble the Imatong Mountains closely. Above 2300 m, Cassipourea
malosana, Schefflera abyssinca, and Croton macrostachyus form the evergreen
montane forest, which slowly gives way to cloud forest communities with Hagenia
abyssinica and Hypericum revolutum from 2400 m onwards. Large areas of Bamboo
are found between 2800 and 3100 m altitude. Cultivation has been more intense on
the drier northern slopes, so only remnants of the natural vegetation survived. Dry
orotropical Cedar and Podocarpus forests have been described up to 3000 m
(Weinert 1983) (Fig. 25), but cloud forest with Kosso and Giant St. John’s Worth
is restricted to a band from 3000 to 3400 m.
Ericaceous vegetation with Erica trimera sl. starts to dominate the vegetation
from this zone upwards to 4000 m. As elsewhere, the vegetation of the ericaceous
belt has been severely altered by human influence, so mosaics of forests, scrub, and
afroalpine communities result. On the central Sanetti plateau, woody vegetation is
restricted to sheltered sites as outcrops or boulder streams up to 4000 m. Instead,
Helichrysum scrub and to a lesser extend tussock grasslands prevail up to the highest
peaks Fig. 26. Many of the grass and shrub species demonstrate clear affinities to the
eastern African Mts. (e.g., Pentaschistis pictigluma, Helichrysum citrispinum), but
the Giant Lobelias are present with only one endemic species (Lobelia
rhynchopetalum, Fig. 27), and the widespread forest species Lobelia gibberoa.
Giant Groundsels are not found in Ethiopia. Strangely, an endemic thistle (Echinops
longisetus) has developed a stem here resembling giant groundsels.
Much of the Ethiopian highlands would bear montane forests if untouched, hence
remnants of these forests still occur in the central part of the country (Tamrat 1993).
Fig. 25 Dry orotropical Cedar and Podocarpus forests in the Bale region, Ethiopia. (Photo
R.W. Bussmann)
Fig. 26 Afroalpine
vegetation on the Sanetti
Plateau. Note endangered
Ethiopian Wolf (Canis
simensis) in the middle.
Ethiopia. (Photo
R.W. Bussmann)
The plant communities are still strikingly similar to those described for Mt. Kenya,
but differences increase north-westwards, until the dry Simen Mts. (Fig. 28) form the
second largest afroalpine region in Ethiopia. Here, the precipitation regime finally
becomes unimodal; the foot zone is rather dry but near the timberline 1500 mm have
been measured (Hurni 1982). Savannahs below 2000 m are followed by evergreen
montane forest with Cedar, Olea europaea, Rapanea simensis, and finally Kosso
between 2000 and 3500 m (3300 m in the southwest). Between 2900 and 3700 m,
ericaceous vegetation takes over, but here Erica arborea is the only tree heather;
Fig. 27 Sanetti plateau with

dense population of Lobelia
rhynchopetalum
Fig. 28 Vegetation zonation in the Simien Mts. (Bussmann 2006)

Fig. 29 Simien mountains with dense Lobelia rhynchopetalum populations, Ethiopia. (Photo
R.W. Bussmann)
joined by Hypericum revolutum, as well as Nuxia congesta in the lower elevations

(Klötzli 1975; Verfaille 1978; Nievergelt et al. 1998). Above this belt, Festuca
macrophylla, Carex erythrorhiza, and Lobelia rhynchopetalum (Fig. 29) occur, the
latter with an inflorescence up to 5m tall. In the highest regions, stands are gradually
replaced by Festuca abyssinica and finally by Helichrysum citrispinum scrub up to
the summit region.
Maloti-Drakensberg: Mountain Vegetation in Southern Africa
Environmental Conditions
The Maloti-Drakensberg forms the highest part of the Great Escarpment which
bounds the interior plateau of southern Africa. The geological structure is relatively
homogenous and characterized by stratified Jurassic flood basalts in the upper
portion, which cap the underlying fine-grained Clarens Sandstone (c. 1800–
2200 m) and other formations of the Karroo Supergroup. For approximately
200 km the towering scarp of the High Drakensberg constitutes the boundary
between the eastern highlands of Lesotho and South Africa. An abrupt and rugged
barrier ranging from the foothills at about 1400 m to numerous peaks, buttresses,
pinnacles, and cutbacks of the escarpment edge lying roughly between 2800 and
3300 m marks the declivity towards South Africa. Some long spurs (the so-called
Little Berg) run out at right angles to the escarpment. The mountain divide consists
of two parts: the northern Berg, a north-east-facing portion between Mont aux
Sources (3282 m) and Giant’s Castle (3314 m), and the southern Berg, a south-
east-facing part of the escarpment between Giant’s Castle and Wilson’s Peak
(3310 m). Towards the west, the summit plateau gently slopes down to the foothills
and lowlands of Lesotho below 2000 m. Rounded slopes, small cliffs, and deeply
incised valleys characterize the treeless plateau. The highest peak Thabana
Ntlenyana (3482 m) lies about 4 km back from the edge of the escarpment (Hilliard
and Burtt 1987; Killick 1978a, 1990).
Mean annual precipitation in this high mountain area ranges from 1240 mm at the
foot of the escarpment (1370 m) to a maximum of about 2000 mm at an altitude of
2300 m. The escarpment edge receives about 1600 mm (rainfall data from the
Cathedral Peak area after Killick 1963). Letseng-la-draai (3050 m), located in the
rain shadow receives 713 mm, and Mokhotlong Town (2377 m) about 35 km to the
west of the escarpment receives only about 575 mm (Killick 1978b). Apart from
these steep gradients in rainfall distribution, the humidity conditions are generally
characterized by strong seasonal differences. Almost 80% of the annual precipitation
falls between October and March, resulting in generally humid conditions during the
growing season (Killick 1978b). Frequent thunderstorms and recurrent periods of
fog, which usually do not extent into Lesotho for more than 3 km contribute to
summer humidity. Snow falls frequently between April and September and is usually
restricted to the summit plateau and nearby slopes of the escarpment. Snow cover
seldom lasts for longer periods, in exceptional years it may last for two months.
Recurrent droughts force plants to sustain water stress. Generally, the southern
Drakensberg is colder and drier than the northern portion. The temperature regime
in the upper valley sections near the escarpment is characterized by a long frost
period, lasting for approximately 180 days (Grab 1997a). Mean annual temperature
is 5.7°C in Letseng-la-draai, and 11.5°C in Mokhotlong Town. Absolute minimum
temperature exceeds –20°C at the highest altitudes. Therefore, periglacial conditions
with Thufur, needle ice, patterned ground, and soil movements are common at higher
elevations (Grab 1994, 1997b, 1998). Prominent differences between the warmer
and drier north facing slopes and the colder and moister southerly slopes result from
different solar radiation income, especially during winter (Granger and Schulze
1977). Obviously, differences of snow cover duration and soil moisture are ecolog-
ically most important and mirrored by more lush vegetation of the southerly expo-
sures. Most plant species of the Drakensberg are adapted to intense solar radiation,
long periods of low temperature and drought, and solifluction due to frost heaving.
The frequency of strong winds, especially in spring and summer, is considered to be
“a powerful factor militating against tree growth” by Hilliard and Burtt (1987).
Phytogeographical Position and Altitudinal Zonation: Problems

of Nomenclature
The vertical sequence of the Maloti Drakensberg has been differentiated into a
montane (approximately 1280–1830 m), subalpine (1830–2870 m), and alpine belt
by Killick (1963, 1990). Other studies like those of Herbst and Roberts (1974) and
Morris et al. (1993) follow this altitudinal division. White (1983) includes the
“afroalpine” belt as an “archipelago-like region of extreme floristic impoverishment”
within the “afromontane region” On the other hand, Hilliard and Burtt (1987) reject
the possibility to include the slopes and summits of the Maloti-Drakensberg in an
“afromontane” and “afroalpine” region on the basis that floristic affinities with
adjacent lowlands of southern Africa are stronger than with the tropical mountains
of Tanzania and Kenya. They argue for the recognition of the area as a distinct
phytochorion, the “South-eastern Mountain Regional Mosaic” instead. In the most
recent overview of the vegetation of southern Africa, Cowling & Hilton Taylor
(1997) include the Drakensberg and Lesotho mountains together with the mountains
of the Cape Folded Belt in an “afromontane region.” In his latest paper, Killick
(1997) describes the treeless vegetation of the summit plateau as tundra. The
proposed nomenclature postulates a low-lying laural belt with supralaural forests
up to approximately 2000 m and largely tree-deficient grass-savannas on the upper
mountain slopes.
Altitudinal Zonation of Vegetation
Themeda triandra grasslands with Trachypogon spicatus, Hyparrhenia hirta,

Harpochloa falx predominantly covers the slopes between approximately 1600
and 2750 m. Large tussocks of Merxmuellera macowanii (1830–2550 m) and
Festuca costata (below 2900 m) occur along the streams. Depending on the kind
of fire regime some of these grass slopes up to about 2400 m contain open Protea
savannas, including 3–5 m high trees of Protea caffra and P. roupelliae with rounded
crowns, and the fire-resistant dwarf species Protea dracomontana. Locally, 2–4 m
high Aloe ferox (¼A. candelabrum), characteristic for drier conditions can be found
as a grassland constituent on steep rocky north-facing slopes in the southern Dra-
kensberg (Loteni valley) (Hilliard and Burtt 1987, Killick 1990). Small forest groves
with the 12–18 m high Yellowwood (Podocarpus latifolius, up to 1900 m) and
Olinia emarginata (up to 2150 m), often festooned with Usnea are confined to
narrow valleys and southern aspects along stream gullies. These forests are larger
and more numerous on the northern Berg (Hilliard and Burtt 1987). The forest
margins are characterized by the Rosaceae Leucosidea sericea, the commonest
woody plant on the Drakensberg (up to 2100 m), Buddleia salviifolia and Bowkeria
verticillata, or by up to 2.5 m high Miscanthus capensis-Cymbopogon validus
grasslands and the Berg bamboo Thamnocalamus tessellatus (1800–2000 m). The
altitudinal distribution of forest margins roughly extends from the valley floors to the
lowermost basalt cliffs at about 2000 m; some tree clumps persist up to 2400 m
(Hilliard and Burtt 1987). Evergreen thickets with the Thymelaeaceae Passerina
drakensbergensis (2100–2450 m on the northern Drakensberg), the Ericaceae
Phillipia evansii (1800–2300 m), the Compositae Euryops tysonii (1800–2500 m),
and the Cupressaceae Widdringtonia nodiflora (“Subalpine Fynbos” after Killick
1963) are limited in extent because of recurrent grass fires. Forest in Lesotho only
exists to a limited extent of 1 km2 (Leslie 1991, Low and Rebelo 1998) with the
largest stand in Tsehlanyane (2100 m), which accommodates a 6 m high Leucosidea
sericea forest.
The vegetation of the summit plateau of Lesotho between approximately 2850 m
and the highest peaks comprise grasslands, dwarf scrub communities, wetland
communities along the streams and moister slopes, and open scree and rock com-
munities, colonized by small and scattered turf patches along the basalt outcrops.
These vegetation types are best developed from December to the end of March. The
grasslands are usually dominated by the Gramineae Merxmuellera disticha, Festuca
caprina, Pentaschistis oreodoxa, Harpochloa falx, and the Cyperaceae Scirpus
falsus in moister places. Subtropical Themeda triandra grasslands are confined to
the lower slopes and warmer north-facing aspects below 3000 m. Generally, species
of temperate affinity like Festuca caprina are more abundant on the cooler aspects
and at higher altitudes. These mixed grasslands are distributed as mosaics, often
interspersed by dwarf scrubs and patches of up to 1 m high Merxmuellera
drakensbergensis stands at water surplus sites. The evergreen tussock grass is
adapted to low temperatures through accumulation of dead and decaying material
that acts as an insulator at the base of the plant. The most common dwarf scrubs are
Helichrysum trilineatum and Erica dominans, covering considerable areas with low
heathlands, especially in the vicinity of Mont Aux Sources. Apparently, this led
Killick (196, 1979) to consider Erica-Helichrysum heathland to be the climax
community of the uppermost altitudinal belt. The cushion-forming succulent
Euphorbia clavarioides is limited to steep rocky and mostly north-facing slopes
(Hilliard and Burtt 1987). Furthermore, the two karroid invader species Chrysocoma
ciliata and Pentzia cooperi (Morris et al. 1993) together with Artemisia afra
dominate the dwarf scrub communities between Sani Pass and Mokhotlong. Wet-
lands and flat marshy grasslands in the riverheads contrast to the surrounding
vegetation. The low-growing wetland vegetation contains a mixture of Gramineae
(Agrostis sp., Poa annua) and Cyperaceae (Scirpus falsus, S. ficinioides, Isolepis
fluitans, and Schoenoxiphium filiforme). Pools and silt patches, often waterlogged in
summer form a habitat for aquatic communities with Limosella inflata, Aponogeton
ranunculiflorus, and Crassula inanis surrounded by Merxmuellera
drakensbergensis tussocks and hygrophilous forbs like Ranunculus meyeri,
Kniphofia caulescens, Moraea alticola frequently accompanied by Trifolium
burchellianum (van Zinderen Bakker 1955, Killick 1978c, Backéus and Grab
1995, Schwabe 1995). Thufur are often colonized by Athrixia fontana.
According to own studies on Thabana Ntlenyana (3482 m), Giant´s Castle
(3314 m) and Hodgon´s Peak (3257 m) the following species reach the summits:
– Dwarf shrubs: dominated by a variety of everlastings such as Helichrysum

trilineatum, H. flanaganii, H. praecurrens, H. milfordiae (forms mats on rock
surfaces), H. subglomeratum, and H. witbergense together with Eumorphia
sericea, Clutia nana, Euryops decumbens, E. acraeus, Erica frigida, and
Lotononis galpinii. Hard cushions of Helichrysum pagophilum and Hirpicium
armerioides are conspicuous on cliffs and crags.
– Herbs: Cineraria geifolia, Glumicalyx flanaganii, G. montanus, Crassula

rubescens, Felicia uliginosa, F. rosulata, F. linearis, Ursinia montana,
Wahlenbergia lobulata, Cerastium arabidis, and Mossia intervallaris
– Grasses Festuca caprina, Koeleria capensis, Merxmuellera disticha,
M. drakensbergensis, Luzula africana, Poa binata, and Pentaschistis galpinii
References
Allaud C, Jeannel R. Le Mt. Kenya. Rev Gen Sci. 1914;25:639–44.
Backéus I, Grab S. Mires in Lesotho. Gunneria. 1995;70:243–50.
Beck E. Cold tolerance in tropical alpine plants. In: Rundel PW, Smith AP, Meinzer FC, editors.
Tropical alpine environments. Cambridge: Cambridge University Press; 1994. p. 77–110.
Beck E, Rehder H, Kokwaro JO. Classification and mapping of the vegetation of the alpine zone of
Mount Kenya (Kenya). Geographica Bernensia African Studies Series. 1990;A8:41–6.
Mt. Kenya Area
Berger P. Rainfall and Agroclimatology of the Laikipia Plateau, Kenya. Geographica Bernensiaca –
African Studies Series. 1989;A7:1–102., 3 maps.
Biedinger N. Ökologie und Diversität der Epiphyten-Vegetation in zentralafrikanischen Wäldern
(Ruanda, Zaire). – PhD thesis. Mathematisch-Naturwissenschaftliche Fakultät. Rheinische
Friedrich-Wilhelms-Universität. Bonn, 226. 1995.
Bussmann RW. The forests of Mount Kenya (Kenya) – vegetation, ecology, destruction and
management of a tropical mountain forest ecosystem. Dissertation Universität Bayreuth, Vol.
1–3. 1994.
Bussmann RW. The forest vegetation of the Harenna Escarpment (Bale Province, Ethiopia) –
syntaxonomy and phytogeographical affinities. Phytocoenologia. 1997;27:1–23.
Bussmann RW. Succession and regeneration patterns of East African mountain forests. Syst Geogr
Plants. 2001;71:959–74.
Bussmann RW. Vegetation zonation and nomenclature of African Mountains – an overview. Lyonia.
2006;111:41–66.
Bussmann RW, Beck E. The forests of Mount Kenya (Kenya), a phytosociological synopsis.
Phytocoenologia. 1995a;25(5):467–560.
Bussmann RW, Beck E. Regeneration- and cyclic processes in the Ocotea-Forests (Ocotea
usambarensis Engl.) of Mt. Kenya. Verh Ges Ökologie. 1995b;24:35–8.
Bussmann RW, Beck E. Regeneration- and succession processes in the Cedar-Forests (Juniperion
procerae) of Mt. Kenya. Ecotropica. 1995c;1:79–84.
Bussmann RW, Beck E. Vegetation units of Mount Kenya Forest Reserve – Comment. In: BITÖK
(editors) Vegetationsökologische, ethnobotanische und -faunistische Beiträge über Äquatorial-
Afrika. Bayreuther Forum Ökologie. 1999;64:17–28.
Bussmann RW, Lange S. Succession and regeneration patterns of East African mountain forests. In:
BITÖK, (editors), Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie. 1999;64:1–16.
Bytebier B, Bussmann RW. Vegetation of Mount Nyiru (Samburu District, Kenya), A checklist and
syntaxonomical survey. J E Afr Nat Hist. 2000/printed 2002;89:45–71.
Cable S, Cheek M. The plants of Mt. Cameroon. Kew: Royal Botanical Gardens; 1998. 198 pp
Cowling RM, Hilton-Taylor C. Phytogeography, flora and endemism. In: Cowling RM, Richardson
DM, Pierce SM, editors. Vegetation of Southern Africa. Cambridge: University Press; 1997.
p. 43–61.
Decurtins S. Hydrogeographical investigations on Mt. Kenya and its w and nw slopes. Laikipia
Reports. 1985;5:1–14.
Decurtins S. Hydrographical investigations in the Mount Kenya subcatchment of the Ewaso Ngiro
River. Geographica Bernensiaca – African Studies Series. 1992;A10:1–151.
Engler A. Die Pflanzenwelt Ostafrikas und der Nachbargebiete. Berlin: D. Reimer; 1895.
Fischer E. Die Vegetation des Parc National de Kahuzi – Biega, Sud Kivu, Zaire. Stuttgart: Franz
Steiner Verlag; 1996. 240pp
Gashaw M, Fetene M. Plant communities of the afroalpine vegetation of Sanetti Plateau, Bale
Mountains, Ethiopia. Sinet. 1996;19:65–86.
Grab S. Thufur in the Mohlesi valley, Lesotho, Southern Africa. Permafrost and Periglacial
Processes. 1994;5:111–8.
Grab S. Analysis and characteristics of high altitude air temperature data from northern Lesotho.
Implications for cryogeomorphic occurrences. Geooekoplus. 1997a;4:49–60.
Grab S. Thermal regime for a Thufa Apex and its adjoining depression, Mashai Valley, Lesotho.
Permafr Periglac Process. 1997b;8:437–45.
Grab S. Non-sorted patterned ground in the High Drakensberg, Southern Africa, Some New Data.
Geogr J. 1998;164:19–31.
Granger JE, Schulze RE. Incoming solar radiation patterns and vegetation response, Examples from
the Natal Drakensberg. Vegetatio. 1977;35:47–54.
Hall JB. Vegetational zones of the southern slopes of Mount Cameroon. Vegetatio. 1973;27:19–69.
Hauman L. La “region afroapline” en phytogeographie centro africaine. Webbia. 1955;XI:467–89.
Hedberg O. Vegetation belts of the East -African mountains. Sven Bot Tidskr. 1951;45:141–96.
Hedberg O. Altitudinal zonation of the vegetation on the east African Mountains. Proc Linnean Soc
Lond (Botany). 1955;165:134–6.
Hedberg O. Features of afro-alpine plant ecology. Acta Phytogeographica Suecica. 1964;49:1–144.
Heinrichs T. Geologie – Ostafrika. Berlin/Stuttgart: Gebrüder Borntraeger; 2001. 231 pp
Hemp A. Ecology of the pteridophytes on the southern slopes of Mt. Kilimanjaro. Part II, Habitat
selection. Plant Biol. 2001;3:493–523.
Hemp A. Ecology of the pteridophytes on the southern slopes of Mt. Kilimanjaro. I. Altitudinal
distribution. Plant Ecol. 2002;159:211–39.
Hemp A, Beck E. Erica excelsa as a fire-tolerating component of Mt. Kilimanjaro’s forests.
Phytocoenologia. 2001;31:449–75.
Herbst SN, Roberts BR. The Alpine vegetation of the Lesotho Drakensberg, a study in quantitative
floristics at Oxbow. J S Afr Bot. 1974;40:257–67.
Hilliard OM, Burtt BL. The Botany of the Southern Drakensberg. Cape Town. Ann Kirstenbosch
Botanic Gardens 1987;15:1–253.
Hurni H. Klima und Dynamik der Höhenstufung von der letzten Kaltzeit bis zur Gegenwart.
Geographica Bernensia G 1982;13.
Hutchins DE. Report on the forests of British East Africa. London: Darling & Son; 1909.
Jackson JK. The vegetation of the Imatong Mts., Sudan. J Ecol. 1956;44:341–74.
Karlowski U. Sekundäre Sukzession im afromontanen Nebelwald, Dynamik, Mechanismen und
Schutz der Biodiversität in zwei Habitaten des Berggorillas (Mgahinga Gorilla- und Bwindi
Impenetrable-Nationalpark) und im Echuya Forest, Uganda. – PhD. Mathematisch Naturwis-
senschaftliche Fakultät. Rheinische Friedrich Wilhelms Universität. Bonn, 178. 1995.
Killick DJB. An account of the plant ecology of the Cathedral Peak Area of the Natal Drakensberg.
Pretoria. Mem Bot Survey South Africa 1963;34:1–178.
Killick DJB. The Afro-Alpine Region. In: Werger MJA, editor. Biogeography and ecology of
Southern Africa. Monographiae Biologicae vol. 31. The Hague, Junk; 1978a. p. 515–560.
Killick DJB. Further data on the climate of the Alpine Vegetation Belt of Eastern Lesotho. Bothalia.
1978b;12:567–72.
Killick DJB. Notes on the Vegetation of the Sani Pass Area of the Southern Drakensberg. Bothalia.
1978c;12:537–42.
Killick DJB. African Mountain Heathlands. In: Specht RL, editor. Heathlands and related
Shrublands. Ecosystems of the world 9A. Amsterdam: Elsevier; 1979. p. 97–116.
Killick DJB. A field guide to the flora of the Natal Drakensberg. Johannesburg: Jonathan Ball and
Ad Donker; 1990.
Killick DJB. Alpine Tundra of Southern Africa. In: Wielgolaski FE, editor. Polar and Alpine
Tundra. Ecosystems of the world 3. Amsterdam: Elsevier; 1997. p. 199–209.
Klötzli F. Zur Waldfähigkeit der Gebirgssteppen Hoch-Semiens. Beitr naturk Forsch Südw Dtl
1975;Band 34:131–147.
Knapp R. Die Vegetation von Afrika unter Berücksichtigung von Umwelt, Entwicklung,
Wirtschaft, Agrar- und Forstgeographie. Jena: G. Fischer; 1973. 626 pp
Knox EB. The conservation status of the giant senecios and giant lobelias in Eastern Africa. Opera
Botanica. 1993;121:195–216.
Leibundgut C, et al. Hydrogeographical Map of Mount Kenya Area. Geographica Bernensiaca –
African Studies Series. 1986;A3:1–70, map
Leslie AD. Indigenous forest and woodland in the Kingdom of Lesotho. Suid-Afrikaanse
Bosboutydskrif. 1991;158:97–103.
Low BA, Rebelo AG. Vegetation of South Africa, Lesotho and Swaziland. Pretoria: Department of
Environmental Affairs and Tourism; 1998.
Mabberley DJ. Notes on the vegetation of the Cherangani Hills, N.W. Kenya. J East Afr Nat Hist
Soc Natl Museum. 1975;15:1–11.
Maitland TD. The grassland vegetation of the Cameroon Mountain. Bull Misc Inf, Royal Botanical
Gardens, Kew. 1932;9:417–35.
Miehe S. Vegetation ecology of the Jebel Marra Massif in the Semiarid Sudan. Berlin/Stuttgart:
J. Cramer; 1988. 171pp
Miehe S, Miehe G. Ericaceous forests and heathlands in the Bale Mountains of South Ethiopia –
ecology and man’s impact. Hamburg: Stiftung Walderhaltung in Afrika; 1994. 206 pp
Morris CD, Tainton NM, Boleme S. Classification of the Eastern Alpine Vegetation of Lesotho. Afr
J Range Forage Sci. 1993;10:47–53.
Ndam N, Healey J, Cheek M, Fraser P. Plant recovery on the 1922 and 1959 lava flows on Mount
Cameroon, Cameroon. Syst Geogr Plants. 2001;71:1023–32.
Negatu L, Tadesse M. An ecological study of the vegetation of the Harenna forest, Bale, Ethiopia.
Sinet. 1989;12:63–93.
Nievergelt B, Good T, Güttinger R. A survey on the flora and fauna of the Simen Mountains
National Park. Walia. 1998;(special issue):1–109.
Oliver EGH. Studies in the Ericoideae (Ericaceae). VII. The placing of the genus Philippia into
synonomy under Erica, the southern Africa species. S Afr J Bot. 1987;53:455–8.
Osmaston H. Glaciations, landscape and ecology. In: Osmaston H, Basalirwa C, Nyakaany J,
editors. The Rwenzori Mountains National Park, Uganda. Kampala: Makarere University;
1996. p. 49–65.
Osmaston H, Kaser G. Rwenzori Mountains National Park, Uganda. In: Gärtner E, Georger C,
editors. Glaciers and glaciations. Glasgow: Bartholomew; 2001.
Petters SW. Regional geology of Africa. Berlin/Heidelberg: Springer; 1991. 722pp
Rauh W. Vegetationsstudien im Hohen Atlas und dessen Vorland. Sitzungberichte der Heidelberger
Akademie der Wissenschaften – Mathematisch naturwissenschaftliche Klasse. 1952. p. 1–118.
Rehder H, Beck E, Kokwaro JO, Scheibe R. Vegetation analysis of the upper Teleki Valley (Mount
Kenya) and adjactent Areas. J E Afr Nat Hist Soc Nat Mus. 1981;171:1–8.
Rehder H, Beck E, Kokwaro JO. The afroalpine plant communities of Mt. Kenya (Kenya).
Phytocoenologia. 1988;16:433–63.
Richards PW. Ecological notes on west African vegetation III. The upland forests of Cameroon
Mountain. J Ecol. 1963;51:529–54.
Schlüter T. Geology of East Africa. Berlin: Borntraeger; 1997. 484pp
Schmitt K. The vegetation of the Aberdare National Park Kenya. Innsbruck: Wagner; 1991. 259pp
Schmitt K. On the afroalpine vegetation of the Ruwenzori Mountains, Uganda. Phytocoenologia.
1992;21:313–22.
Schwabe CA. Alpine Mires of the Eastern Highlands of Lesotho. In: Cowan GI, editor. Wetlands of
South Africa. Pretoria: Department of Environmental Affairs and Tourism; 1995.
Snowden JD. A study of the altitudinal zonation in South Kigezi and on Mounts Muhavura and
Mgahinga, Uganda. J Ecol. 1933;21:7–27.
Speck H. Soils of the Mount Kenya Area, their formation, ecological, andagricultural significance.
In: Winiger M, editor, Mt. Kenya Area. Contributions to ecology and socio-economy. African
Studies Series A1, 1986. p. 47–72.
Tamrat B. Vegetation ecology of remnant Afromontane forests on the Central Plateau of Shewa,
Ethiopia. Uppsala: Opulus Press AB; 1993. p. 64pp.
van Heist M. Accompanying report with the Land unit map of Mount Elgon National Park.
Kampala: Mount Elgon Conservation and Development Project; 1994a. p. 83pp.
van Heist M. Land unit map of Mount Elgon. Kampala: Mt.Elgon Conservation and Development
Project; 1994b.
van Heist M. Land unit map the Ruwenzori Mts. National Park. Fort Portal: WWF-RMCDP; 1999.
van Zinderen Bakker EM. A preliminary survey of the Peat Bogs of the Alpine Belt of Northern
Basutoland. Acta Geogr. 1955;14:413–22.
Verfaille M. The Ericaceous Belt of the Semien Mountains National Park, Ethiopia. Biol
Jbdodonea. 1978;46:210–23.
Weinert E. Vegetation zones in Bale Mountains near Goba, S. Ethiopia. – Wissenschaftliche
Zeitschrift der Martin Luther Universität Halle Wittenberg 1983, 41–67.
Weischet, W. & W. Endlicher. 2000. Regionale Klimatologie. Teil 2. Die Alte Welt. Teubner,
Stuttgart, Leipzig. 625 pp.
Wesche K. The high-altitude environment of Mt. Elgon (Uganda/Kenya) – Climate, vegetation and
the impact of fire. Ecotropical Monogr. 2002;2:1–253.
White F. The forests of Mount Kenya. Forest Society J Oxford. 1950;3(5):32–8.
White F. The vegetation of Africa. Paris: UNESCO; 1983.
Ethnobotany of Mountain Regions: Eastern
Africa

Grace N. Njoroge
Geography and Geology
The eastern African plains are bordered by a chain of mountains consisting of old
cristallinic Precambrian basement rocks, mainly extremely durable gneisses and
granites (Nyiro-Ndoto and Mathews Ranges, Shackleton 1946). A series of
Quarternary volcanic peaks like Mt. Kulal (2285 m), Mt. Marsabit (1707 m), and
the Huri Hills (1479 m) tower over the inland plains. Most regions of Central Kenya
are covered with tertiary to recent lavas and tuffs (Baker 1960; Mason 1963). The
soils of the plains consist mainly of Vertisols, Regosols, Lithosols, and Cambisols.
The mountain slopes are covered with humic Nitisols and Acrisols in case of the
basement formations and the lower volcanic areas, and deep humic Andosols above
about 2700 m (Mäckel 1986; Mäckel and Schultka 1988; Mäckel and Walter 1983;
Schmitt 1991; Speck 1983).
R. W. Bussmann (*)
G. N. Njoroge

Climate
The climate of East Africa shows the characteristic pattern of inner tropical regions
with large diurnal temperature oscillations, known as “Tageszeitenklima” (Troll
1959), but a small amplitude of the monthly means in the course of the year. The
only seasonality originates from the distribution of the precipitation, as the region is
characterized by two rainy seasons following the position of the inner-tropical
convergence zone (ITC). The "long rains" last from March to June, while the
“short rains” occur from October to November. Due to the equatorial position, the
daylight period is nearly constant all year round, which is an important factor for
vegetation growth.
According to Jätzold (1977, 1981), the study area is part of the hot, arid tropical
climate, with two short, subhumid seasons, with mean monthly temperatures of
26–20°C in the plains, in contrast to 17–19°C in the mountains (Gatab, 1657 m). The
average annual rainfall can be as low as 100–150 mm in the Hedad plain and Chalbi
desert, rising to 500 mm in the valleys of the Nyiro-Ndoto Mountains. In the
mountain forest zone, a rainfall of about 1200–2000 mm can be reached (Edwards
et al. 1979). The main rainfall is concentrated on two wet seasons from March–May
and October–December, extreme rainfalls occur, e.g., 175 mm within 6 h in Gatab on
Mt. Kulal (Mäckel and Walter 1983; Kenya Meteorological Department 1974,
1984).
The Main Source of Useful Plants: The Flora of East African

Mountain Forests and Woodlands
While the Alpine zones of the East African Mountain systems are certainly the most
well-known floristic zone of the region (Hedberg 1964; Figs. 1 and 2), they were
originally surrounded by dense forests, greatly limiting access for the local popula-
tion. In addition, traditionally the mountains were seen as seats of the gods, and thus
access was restricted by taboos (Bussmann 1994). For these very reasons, plant use
in the area has greatly been restricted to mountain forest and savannah ecosystems
(Bussmann 2006b, Bussmann et al. 2006, 2011, Kiefer and Bussmann 2004, 2008;
Luizza et al. 2013), and much of the areas in the African highlands have been
converted into dense agricultural zones Fig. 3).
Ecological observations on growth and germination of East African forest species
also exist (Bussmann 1999c, Bussmann and Lange 2000). Descriptions of the
regeneration processes in various East African forest types exist (Bussmann
2002a, b, 2004a, b, 2006a; Bussmann and Beck 1995b, c, 1999; Bussmann and
Lange 1999; Lange et al. 1997), based on field data particularly from Mt. Kenya.
Unfortunately, the natural forests of East Africa have been subjected to human
impact. Large forest tracts have been converted into agricultural fields, often up to
the alpine bog zone (Figs. 4, 5, and 6). Much of the forest clearing involves fire, often
devastating large areas (Figs. 7 and 8). Moreover, the forests, due to selective
logging, they also have undergone significant changes of their species composition.
In addition, the forests serve as the main source of firewood (Fig. 9). Although
Ethnobotany of Mountain Regions: Eastern Africa 45
Fig. 1 Mt. Kenya summit, with dense alpine vegetation dominated by Dendrosenecio
keniodendron. Kenya. (Photo R.W. Bussmann)
Fig. 2 Mt. Kenya summit and forests seen from the savannah zone, Kenya. (Photo
R.W. Bussmann)
private tree planting of exotic species has reduced the fuelwood deficit considerably
(Holmgren et al. 1994), the destruction of the natural forests has increased drastically
(Bussmann 1996, Fig. 10). Currently, an annual deforestation rate of 1.5% is
Fig. 3 Mt. Kenya summit surrounded by highly used agricultural and tea zone. Kenya. (Photo
R.W. Bussmann)
Fig. 4 Mt. Kenya, fields in destroyed forest areas on the Northern slopes. Kenya. (Photo
R.W. Bussmann)
Fig. 5 Bale Mountains, Ethiopia – fields in Juniperus procera dominated forests. Ethiopia. (Photo
R.W. Bussmann)
Fig. 6 Mt. Kenya, fields in the Juniperus procera forest zone. (Photo R.W. Bussmann)
Fig. 7 Bamboo destroyed by forest clearing, Aberdare National Park. Kenya. (Photo
R.W. Bussmann)
Fig. 8 Large forest clearing fire in the forest zone of Mt. Kenya. Kenya. (Photo R.W. Bussmann)
Fig. 9 Firewood collection in Bale Mountains National Park zone. Ethiopia. (Photo
R.W. Bussmann)
reached, partly driven by a human population growth rate of up to 3.8%, If current

rates of forest clearing continue, no natural forest will remain by the year 2040
(Barnes 1990). Today, approximately 2% of East Africa is still covered with
indigenous forests (Doute et al. 1981). In addition to providing timber and firewood,
the forests represent the largest water catchments systems.
Eastern African Forest Vegetation
Evergreen Submontane Forests
The Evergreen submontane forests of African mountains (Phytosociologically

known as Ocotetea usambarensis) (Bussmann 1994, 1997, 1999a; Bussmann and
Beck 1995a, b, 1999; Bussmann and Lange 1999) with their two storied tree layer
are differentiated by Ocotea usambarensis, often dominant in the upper canopy,
Xymalos monospora, Lasianthus kilimandscharicus, Pauridiantha holstii, and
Psychotria orophila in the lower tree and the often dense shrub stratum, and
Asplenium sandersonii, Asplenium elliottii, as well as Panicum calvum among the
herbal species. Character species of this class are Strombosia scheffleri and Apodytes
dimidiata in the upper tree layer and Tabernaemontana stapfiana, Ochna insculpta,
Macaranga kilimandscharica, and Peddiea fischeri in the lower canopy and the
shrub layer. Piper capense, Cyphostemma kilimandscharicum, Oplismenus hirtellus,
Fig. 10 Forest destruction in Central Kenya since about 1880
Plectranthus luteus, Begonia meyeri-johannis, and the ferns Dryopteris kilemensis,

Blotiella stipitata, Elaphoglossum lastii, Trichomanes borbonica, Asplenium
theciferum, and Oleandra distenta are character species of the herbal layer. The
Ocotetea usambarensis forests are found mainly on the very wet south-eastern and
southern slopes of Mount Kenya on altitudes between 1550 and 2550 m, growing on
humic Niti- and Acrisols and receiving an annual rainfall of 1500–2500 mm.
The lower forest types of Harenna in Ethiopia, at altitudes between 1900 and
2300 m, with Acrisols as substrate, clearly also belong to this vegetation class,
although the naming species and many other differential and characteristic species
do not occur in Ethiopia. Asplenium sandersonii and Panicum monticule, however,

occurred as differential species, whereas among the character species the epiphytic
fern Elaphoglossum lastii, Apodytes dimidiata, and Oleandra distenta, as well as
Trichomanes melanotricum and the shrubby Rubiacee Psychotria orophilat were
found. Oplismenus compositus, a very near relative to Oplismenus hirtellus, which is
a character species of the class further south, is regarded as a vicariant of that species
in Ethiopia (Bussmann 1997).
Evergreen and Evergreen Xeromorphic Montane Forests
The Cedar-forests of Mount Kenya, growing on humic Acrisols (Speck 1986) with
rainfall between 700 and 1500 mm, represent the typical vegetation of the altitu-
dinal range between 2500 and 2950 m. Phytosociologically, they form the alliances
Cassipourion malosanae and Juniperion procerae in the order Juniperetalia pro-
cerae and the class Juniperetea procerae (Bussmann 1994, 1999a, b, 2002,
Bussmann and Beck 1995a, 1999; Bussmann and Lange 1999; Bytebier and
Bussmann 2000). The mainly two-storied forests of the Cassipourion are found
mainly on the lower altitudes of the Juniperetalia procerae range, namely, between
2150 and 2650 m, and occupy especially the western and southwestern slopes of
Mount Kenya, where the rainfall is significantly higher than on the northern parts
of the mountain. Juniperus procera itself is often found only scattered in these
forests or is completely absent due to heavy logging and regeneration problems.
Dominant tree species of the Cassipourion are the Rhizophoracee Cassipourea
malosana and Olea capensis ssp. hochstetteri, the East African Olive, both
forming the upper canopy. Lepidotrichilia volkensii (Meliaceae) in the lower tree
and shrub strata as well as Ilex mitis (Aquifoliaceae), the African Holly, are further
important woody species.
In the open single or double storied Cedar forests, Juniperus procera grows about
50 m tall under favorable conditions. Other important tree species, mainly of the
lower canopy, are the African Olive (Olea europaea ssp. africana) and Podocarpus
latifolius. A patchwise dense grass layer with Stipa dregeana and Brachypodium
flexum, interspersed with herbs as Sanicula elata, Isoglossa gregorii, and
Achyranthes aspera, and low shrubs like Berberis holstii is also characteristic of
these open Cedar forests.
Evergreen Montane Bamboo Forests
The dense forests dominated by the African Bamboo, Oldeania africana (in the
following in this chapter we use the much better used synonym Sinarundinaria
alpina), which are found on various East African mountains, form a clear-cut class
(Sinarundinarietea alpinae Bussmann 1994, 1997, 1999a, b, 2002; Bussmann, Beck
1995a, 1999; Bussmann and Lange 1999; Bytebier and Bussmann 2000) of their
own. Sinarundinaria alpina (Poaceae), Impatiens hoehnelii (Balsamiaceae), and
Pseudocarum eminii (Umbelliferae) as well as Selaginella kraussiana

(Selaginellaceae), Cyperus dereilema, Anthriscus sylvestris (Umbelliferae), and
Cardamine obliqua (Cruciferae) are the most characterizing species of this
forest type.
Bamboo forests are restricted to a relatively narrow ecological range, mainly
depending on a soil temperature of 10–15 °C and the presence of very deep volcanic
soils, namely, humic Andosols. On Mount Kenya, the Sinarundinarietea alpinae are
found in a distinct belt around the whole mountain, interrupted only on the dry
northern slopes, being most extensive on the very wet southern parts.
The cyclic changes of the bamboo vegetation lead to regular changes of the
floristic composition and thus to difficulties in the classification of the different parts
of the cycle. Due to the fact that one single growth cycle of Sinarundinaria alpina is
supposed to take 15 (Clayton 1970) to about 40 years (Wimbush 1945; Agnew 1985,
and own observations give indications for even longer periods), all vegetation units
of the cycle are found for several years and are clearly differentiated from each other
by a unique floral composition. For that reason, the “stages” of the bamboo cycle
mentioned by different authors (Agnew 1985; Schmitt 1991) are regarded as clearly
differentiated subassociations of the respective higher taxonomic units.
Evergreen Subalpine Forests
Open evergreen, subalpine Hagenia forests extend from 2900 to 3300 m, where low
clouds and mist are frequent. The dominant tree is the “Kosso”-tree, Hagenia
abyssinica (Bruce), J.G. Gmel., of the Rose family.
Phytosociologically, the Hagenio abyssinicae – Hypericion revoluti, charac-
teristic for the wetter south-eastern to western slopes of Mt. Kenya, and the
Hagenio abyssinicae – Juniperion procerae of the drier North, make up the
order Hagenietalia abyssinicae, which is the only one of the class Hagenietea
abyssinicae (Bussmann 1994, 1997, 1999a, 2002; Bussmann and Beck 1995a,
1999; Bytebier and Bussmann 2000; Lange et al. 1997). The Hagenio – Hyper-
icion includes the pure Hagenietum abyssinicae with only low undergrowth
beneath the dominant Kosso trees, and the Hypericetum revoluti, where Hyper-
icum forms a dense shrub layer in the understory. A third association, the
Gnidietum is dominated by Gnidia glauca, a tall Thymeleace, which together
with Hagenia is forming the tree layer.
In the drier North of Mt. Kenya, the Hagenia-dominated forests are replaced by
mixed forests of conifers and broad-leafed species in which Hagenia abyssinica is
codominant (Hagenio-Juniperion). The Hagenio abyssinicae – Juniperetum procerae
is the only association in this alliance, which, however, comprises three sub-
associations: the Hagenio abyssinicae – Juniperetum procerae caricetosum
peregrinae, the Hagenio abyssincae – Juniperetum procerae pennisetetosum
clandestini, and the Hagenio abyssinicae – Juniperetum procerae brachypodietosum
flexi, which Bussmann (1994) interpreted as an early stage of succession after fire or
disturbance by heavy logging.
Regeneration Patterns of Eastern African Forests
Regeneration Cycle and Successional Patterns in the Ocotetea

sambarensis
Ocotea usambarensis, the most characteristic tree of this vegetation class is wide-
spread on East African mountains (White 1978, 1983; Friis 1992), inhabiting an
altitudinal range between 1300 and 2600 m. Although the species is heavily logged
for its valuable hardwood, nearly no research has been done on its regeneration, and
literature on this topic is therefore very rare.
Only Willan (1965) described the difficulties to regenerate Ocotea on the south-
ern slopes of Kilimanjaro and in the West Usambara Mountains. According to his
reports, Macaranga ssp. is dominant in gaps where Ocotea is absent, i.e., has died
naturally or has been logged out. Saplings of Ocotea are successfully outcompeted
by Macaranga, a pioneer species with a 50% higher growth rate than that of the
Camphor (Bussmann 1994). Willan proposed removal of all Macaranga trees before
replanting Ocotea.
During the fieldwork reported here, a great variety of the Ocotetea usambarensis
was examined. Nearly everywhere only very old Ocotea trees of approximately 300–
600 years of age (stem diameter 1.2–2.5 m) were found. Very few scattered young
trees (less than 100 years old) and seedlings were encountered. In the Mount Kenya
area, the main flowering of the dioecious Ocotea usambarensis occurs between early
March and late April, at the beginning of the long rains. A second flowering can be
observed after the start of the short rains in November. In most cases, however, only
few trees can be found flowering during these periods, whereas every 5–7 years
mass-flowering can be found. Most of the produced fruits are attacked by gall-insects
when still unripe or are eaten by birds. The few remaining seeds are only viable for a
few days, even under the most favorable conditions like high humidity and radiation.
Therefore, regeneration of Ocotea by seedlings is very rare (Bussmann and Beck
1995b; Bussmann and Lange 1999).
Notwithstanding the uniform age structure, numerous Ocotea saplings originat-
ing from old roots were observed. Seedlings and young specimens of the other tree
species of the Ocotetea usambarensis were found everywhere, indicating the excep-
tive character of Ocotea itself, while the other species follow normal regeneration
patterns and therefore form a kind of Mosaic structure.
For Ocotetea usambarensis the successional cycle shown in Fig. 11 appears to be
typical:
Equally old populations of Ocotea usambarensis trees die over a largely extended
period of time (1), creating a mosaic of gaps in which the Euphorbiacee Macaranga
kilimandscharica starts growing as secondary species (2). This leads to a vegetation
similar to the Macarangion kilimandscharicae (Bussmann, Beck 1995a, b), but only
on small scale, and does not change the structure of the forest to a significant extent.
Regeneration of Ocotea from seedlings is negligible, but numerous saplings from the
roots of a fallen tree, no longer suppressed, by the mother tree, start growing (3).
Macaranga kilimandscharica hereby plays an important role as shade tree, as
Fig. 11 Regeneration cycle of Ocotea usambarensis dominated forests in East African Mountains
Ocotea saplings apparently do not tolerate full sun. After the break down of the
relatively short-living Macaranga trees, the meanwhile well-established young
Ocotea trees close the gaps and by shading prevent further germination or establish-
ment of Macaranga kilimandscharica (4). Thus, Ocotea stands of a more or less
uniform age structure result.
In Ethiopia, Ocotea usambarensis itself is not present in the Ocotetea
usambarensis, being replaced by Ocotea kenyensis. No data are available on the
regeneration of this species itself. However, all observations indicated, that the
Ethiopian submontane forests regenerate also as Mosaic-Climax.
Under conditions of heavy logging and over-exploitation, a different successional
cycle was encountered in wide parts of the Camphor forests, leading to a drastic
change of the vegetation structure.
The removal of Ocotea usambarensis on a large scale (5) leads at once to
numerous large gaps, with many additional trees are killed during felling, removal,
and transport of Ocotea. The often-used logging practice of uprooting of the Ocotea
trees prevents the regeneration by root saplings. The big gaps are fast closed by
Macaranga kilimandscharica and at higher altitudes by Neoboutonia macrocalyx.
As Camphor does not regenerate in these man-made “gaps” after breakdown of the
secondary trees, these forests regenerate in a cycle of their own (6). The possibilities
for a regeneration to primary forest are unknown, and would at least require very
long timespans, and a complete logging stop around the gaps, to allow colonization.
The high population density of big herbivores, particularly elephants, was
claimed to be responsible for the failure of Ocotea regeneration. The influence of
megaherbivores on vegetation has however mainly been studied in savanna ecosys-
tems (Buechner and Dawkins 1961; Kortland 1984), and only few data exist on
forest habitats (Holloway 1965; Schmitt 1991; Schmitt and Beck 1992). The argu-
ment that especially elephants are responsible for the failure of Camphor regenera-
tion could not be corroborated by the data of this study: In areas with an estimated
equally high elephant population (as counted by dung), regions without heavy
logging showed considerable regeneration, whereas in logging areas, especially
when the Ocotea trees had been felled by uprooting, nearly no regeneration was
observed. Therefore, the main reasons for the failing regeneration of Ocotea
usambarensis are over-exploitation and unsustainable logging practices.
The Juniperetea procerae: Mosaic Climax of the Cassipourion

malosanae and fire-cycles in the Juniperion procerae
Two vegetation alliances, the Cassipourion malosanae (montane broadleaved for-

ests) and the Juniperion procerae (montane xeromorphic forests) are comprised in
the Juniperetalia procerae. Both alliances regenerate differently and therefore have to
be treated separately. The regeneration processes observed were found to be the
same in the whole research area.
Regeneration of the Cassipourion malosanae

No indications of a regenerative cycle could be observed in the Cassipourion
malosanae. Therefore, its associations are not linked by successional processes. In
general, regeneration follows a dynamic pattern. Trees of all age classes were found
in the Cassipourion forests suggesting a mosaic climax. Due to the environmental
conditions, especially the high rainfall, fires are rare in the Cassipourion and,
whenever occurring, they were limited to small areas. Juniperus procera occurs
only sporadically in the Cassipourion, as its germination and regeneration are
suppressed by heavy shade and the dense vegetation covering the forest floor for
this reason. Regenerating Cedar was only found in gaps where the bright light,
requiring seedlings compete with those of the broad-leaved species.
The Fire Cycles in the Juniperion procerae

In contrast to the Cassipourion malosanae, fire apparently plays an important role in
the regeneration cycle of the Juniperion procerae. Juniperus procera, the African
Pencil Cedar, typically grows in the drier montane, submontane, and subalpine areas
of the East African mountains between 1000 and 3000 m (Kerfoot 1961). Temper-
ature as well as rainfall apparently limit its spreading (Winiger 1979).
In his study of the South Aberdare Range, Wimbush (1937) suggested that
without fire the Juniperus forest is only a serial stage in the succession to a broad-
leaved forest, proposing a fire induced regeneration cycle.
After fire, herbaceous pioneer species and Juniperus procera seedlings invade the
gap (1), leading to a regenerating forest (2), where broad-leaved species as Olea
capensis ssp. Hochstetteri, Olea europaea ssp. cuspidate, and Olinia rochetiana
become established too. After about 20 years, the Juniperus trees have grown about
9 m high. When a closed canopy is established, the Cedar trees start to outgrow the
other species, reaching about 30 m after 80–90 years. If the number of Juniperus
trees established after a fire was high, a pure Cedar forest (3) results by suppression
of other trees. After about 200 years, the maximum height of about 40–46 m is
reached. Although frequently infected by the fungus Fomes juniperinus, thinning the
crowns, the Juniperus trees may live for another 300–700 years. In case, a burnt area
is mainly invaded by broad-leaved species, these create the canopy and only
scattered Juniperus specimens emerge. As Cedar cannot regenerate without fire,
natural gaps are filled with broad-leaved species. Therefore, pure broad-leaved
forests built the climax community (4) after some time. These broad-leaved forests
represent a stable climax, if not destroyed by fire (Fig. 12) (Bussmann and Beck
1995c; Bussmann and Lange 1999; Bussmann 2002a, 2004b).
The present study supported these findings of Wimbush (1937) only partly. The
Cassipourion malosanae can be seen as part (4) of the mentioned cycle, although the
impact of the rare fires appears to be very limited. Scattered Juniperus trees
regenerate in natural gaps, without fire, especially so at the upper limit of the
Cassipourion.
In areas where Juniperus procera has been removed by logging, more or less pure
stands of broad-leaved trees were encountered with only few Cedar trees left.
Fig. 12 Regeneration cycle of Juniuperus procera dominated forests in East African Mountains
However, most Cedar forests were in stage 4 of the succession cycle. Absolutely
pure stands of Cedar (stage 3) were never found at lower and medium altitudes.
Forests clearly dominated by Juniperus procera were seen only at higher altitudes
forming the upper borderline of the Juniperion procerae. These forests were included
in the Myrsino africanae – Juniperetum procerae (Bussmann and Beck 1995a). From
several large burnt areas on Mount Kenya, the following regeneration process was
concluded (Fig. 13).
After fire, a pioneer stage with a very high initial cover of Juniperus and Myrsine
africana develops (1), where soon broad-leaved trees start growing, producing
various independent climax subassociations (2), due to different ecological condi-
tions and different seed intake. However, Juniperus procera remains always the
dominant canopy species. These climax stages are open forests with only scattered
shrubby undergrowth, allowing regeneration of all tree species. Thus, seedlings and
young trees of Juniperus were often encountered in these forests, even when fire had
been absent for a longer period (3). This indicates, that at higher altitudes broad-
leaved forests in sensu Wimbush (1937) cannot replace the Juniperus forests, even if
burning has not occurred for long time. If a fire occurs, a new regeneration cycle
takes place.
Cyclic Processes in the Bamboo Forests
The African Bamboo, Sinarundinaria alpina, inhabits the highland and mountain
areas of East and Central Africa at altitudes between 1800 and 3400 m (Hedberg
1951; Dale and Greenway 1961). Bamboo forests prefer deep volcanic soils, espe-
cially Andosols (Speck 1986 and own observations).
According to Wimbush (1945), bamboo shoots can reach a height of up to 18 m
and a stem diameter of up to 10 cm New vegetative shoots are sprouting from old
rhizomes every 2–3 years. The main growing period starts some weeks after the long
rains and lasts for about 1–2 months. New shoots are to be encountered after each
long rainy season. However, maximum rates of biomass production with most of the
4500–7000 culms produced per hectare require longer periods of favorable climate.
When old culms die and fall over, young sprouts from the dense network of old
rhizomes will replace them, thus giving rise to a homogenous bamboo forest. In
undisturbed areas, where the lifespan of a culm is estimated as about 14 years, the
numbers of dying culms roughly equal new ones. Flowering, after which the culms
die, occurs in areas of about one to several hundred acres in more or less regular
intervals of at least 40 years (Wimbush 1945). On Mount Elgon, Clayton (1970)
observed a flowering cycle of only 15 years.
Agnew (1985), referring to a long-term knowledge of the vegetation of the
Aberdare Range distinguished a bamboo cycle with four stages:
1. Pioneer: Regeneration from sections of old rhizomes and rhizome fragments and
occasionally from fallen culms, lasting about 3–5 years during which dense
thickets of 1–3 m tall Sinarundinaria stems intersperse the still present Sambucus
Fig. 13 Regeneration cycle of broad-leaved forests domitaed by Cassipourea, Olea and

Podocarpus in East African Mountains
africana bushes of the previous cycle. No seedlings or even viable seeds have
ever been reported from stands of Sinarundinaria.
2. Building: During this period, the growing bamboo exceeds the average plant
height in the stand, and the vegetation becomes floristically depleted as the bright
light requiring species of the pioneer stage disappear and more shade tolerant
species of the following stage which might be survivors of a previous cycle are
getting momentum. The pH of the soil starts decreasing.
3. Mature: No further increase in culm height takes place during this stage. The
culms show even distribution and the soil pH is decreasing further.
4. Flowering and 5. Sambucus Stage: Within 2–3 years after mass-flowering the
culms die and fall and light-demanding species invade, above all Sambucus
africana. The soil pH increases again during this stage. The occurrence of trees
such as Podocarpus latifolius, Nuxia congesta, and Dombeya goetzii in the
bamboo therefore could be indicating the sites of frequent flowering.
In general, the sequences described by Agnew (1985) were also found on Mount
Kenya (Bussmann 1994, Bussmann and, Beck 1995a, Bussmann and Lange 1999),
Mt. Nyiro in Northern Kenya (Bussmann 2002, Bytebier and Bussmann 2000), and
the Harenna Escarpment in the Ethiopian Bale Mountains (Bussmann 1997). How-
ever, the vegetation showed considerable floristic differences on wet and drier sites
of bamboo forests, even if the areas compared were in the same stage of the cycle.
In the North of Mt. Kenya, young bamboo was apparently invading parts of the
Juniperion procerae with open undergrowth as a new habitat, forming dense, more or
less extended thickets. To clarify whether this represented a new colonization, very
young sprouts were excavated in several places, especially when forming outposts
far from young bamboo thickets. All samples indicated that the new shoots arose
from old rhizome fragments. Therefore, the areas which seemed to be newly
colonized, proved to be regenerating old bamboo stands, where Sinarundinaria
alpina had died many years ago, and very small vegetative bamboo sprouts had
been part of the heavily grazed pasture, so that the local people did not remember
bamboo in these locations (Mathenge, personal communication, 1993). Since no old
culms were found, they must have been decomposed a long time ago, as dead culms
normally stay for many years especially in the drier areas. Therefore, the time gap
between two subsequent bamboo cycles appears to be very long in this area, i.e.,
regeneration seems to be very slow.
Two explanations for that might be feasible:
– The region is fairly dry, i.e., Sinarundinaria alpina is at the limit of its distribution
and is therefore growing much slower than in other areas.
– The Sirimon area in the Northwest has a very high population of herbivores,
particularly buffaloes and elephants which are known to browse especially young
shoots and culms of bamboo (Wimbush 1945). The bamboo thickets in the area
show sincere browsing damage which decreases the growth rate and the degree of
regeneration. During this study, numerous game trails were found even in very
dense mature bamboo stands and new culms were often found browsed and
destroyed. These observations suggest even more that the elephant family groups
migrate through the bamboo during the growing season searching for fresh culms.
This corresponds closely to the findings of Agnew (1985).
In Ethiopia, no distinct Sambucus-stage could be observed. Apparently Sambucus

africana does not occur naturally in Ethiopia (Friis, personal communication, 1994);
however, after dieback of the flowered bamboo, other light demanding species occur,
until the pioneer stage takes over again.
During the presented study, on Mount Kenya and the neighboring Aberdare
Range, bamboo stands were mainly found on the wetter western, southern, and
southeastern slopes and were absent in the Northern areas. From this distribution, it
was concluded that Sinarundinaria alpina requires precipitation of about 800–
1000 mm per year although arid periods are tolerated. In clear correspondence to
the observations of Wimbush (1945), new culms were mainly produced after
prolonged rainy areas. During the research period of 1992–1996, the main culm
production occurred in large areas in May/June 1995, after 9 months of unusually
heavy rainfall. This, and the fact, that the newly produced culms were mostly much
larger and taller than the old ones clearly depicts the dependency of bamboo growth
on rainfall duration and intensity.
Only small areas of flowering or postflowering bamboo could be found, and no
synchronization of flowering among different populations (Mt. Kenya, Aberdares,
Mt. Nyiro, Harenna Escarpment, Bale Mts., Ethiopia) could be observed. In clear
contrast to the occurrence of mass-flowering effects suggested by Wimbush (1945),
this phenomenon did not occur anywhere in the study area. Instead, bamboo was
found flowering and later dying back only in little patches, after periods of favorable
climatic conditions. Therefore, it has to be concluded that in contrast to many Asian
and Latin American bamboo species, Sinarundinaria alpina shows no synchronized
flowering at all. Despite the flowering and the subsequent die back, seeds or
seedlings of the species were never found, and exclusively vegetative regeneration
of bamboo was observed in the whole growing area.
Of special interest is the occurrence of large areas of Bamboo forests in Ethiopia.
Although covering wide areas, (Harenna, Bussmann 1997), Illubabor (Friis, personal
communication, 1994) these forests were not recognized as specific forest types or
not mentioned at all by earlier authors. The large bamboo populations of South and
Central Kenya (Mt. Kenya, Aberdares, Mau, about 1000 km South of Harenna) are
linked to Harenna only by a small population occurring on top of Mt. Nyiro at the
southern end of Lake Turkana, which is also at least 500 km away from the Bale
mountains. As mentioned above, viable seeds of Sinarundinaria alpina have never
been found, and the species is known to regenerate only vegetatively, thus a
propagation by animals seems very unlikely. Therefore, the large gaps between the
main population and even these outposts pose some interesting questions regarding
the ecology and former distribution of Sinarundinaria, as influenced by the
palaeoclimate of the region. This requires further fieldwork.
Regeneration in the Subalpine Hagenia abyssinica forests
Already Schmitt (1991) and Bussmann (1994) supposed a cyclic regeneration of

Hagenia abyssinica depending on fire.
At first sight, it seems fairly incomprehensible that fire should be the decisive
factor for the regeneration of the Hagenia forests, which are predominately located
in the misty regions of the East African mountains, as high precipitation, as well as
frequent cloud formations, might prevent the vegetation and dead plant material
from becoming sufficiently dry to burn. Martin (1982), however, mentioned that
fires tend to be most frequent in areas with a combination of comparatively high rates
of primary production, slow decomposition rates and occasional drying of the fuels.
Even Houston (1995) stated that the alternation of wet and dry seasons produces
ideal conditions for fires in wet regions. On Mt. Kenya there are rainfall anomalies in
form of severe droughts, as for example the one occurring 1890 (Bussmann, personal
communication, 1994). In recent times, the rainfall of the year 1990 was likewise
extremely low, leading to the assumption that extraordinary dry periods, favoring the
outbreak of fires, occur in approximately 100 years’ time intervall. Thus, in suffi-
ciently wet years, a relatively high plant productivity, together with the large litter
production of an adult Hagenia, which is sheding the older leaves continually
(Miehe and Miehe 1994) and the low decomposition rates in the cold climate of
the subalpine forests, result in the accumulation of fuel, which seem to encourage the
spreading of fires in extremely dry years. The existence of charcoal layers in the soil
profiles, dug within the Hagenia forests, indicate, that fires are really a characteristic
feature of these misty forests. Field observations yielded that regeneration stages of
Hagenia abyssinica are always growing together with either young seedlings of
Juniperus procera and/or fire indicating vegetation. Further, the equal ages of the
Hagenia forest stands suggest that the Kosso is not regenerating successively, but all
of a sudden after a disturbing event like fire. Germination tests yielded that the
Hagenia seeds were not germinating, unless the dense undergrowth was removed.
Fire is the only natural factor that is able to destroy the undergrowth of larger areas,
and thus to provide an environment with reduced competition, suitable for germi-
nation of the seeds. Germination therefore depends on a fire event, which is
removing the competing undergrowth and further stimulates the germination of the
Kosso with a sudden heat surplus.
The proposed regeneration cycle of the Hagenia forests of Mt. Kenya (Lange
et al. 1997; Bussmann and Lange 1999) and other areas is shown in Fig. 14. It refers
only to the associations of the mixed Hagenia – Juniperus stands (alliance Hagenio
abyssinicae – Juniperion procerae), which are distributed in the drier areas, as
regeneration of Hagenia abyssinica was predominately found in these more fre-
quently burning areas. In contrast, no clear succession stage after fire could be
differentiated for associations of the second alliance of the class Hagenietea, the
Hagenio abyssinicae – Hypericion revoluti. It is supposed that the lower syntaxa of
this alliance are also subject to cyclic regeneration after fire, but further research in
recently burned areas of these pure Hagenia stands would be necessary, to verify this
assumption. Stage 1 of the cycle describes the climax forests with adult Kosso trees
Fig. 14 Regeneration cycle of Hagenia abyssinica dominated timberline forests in East African
Mountains
of uniform size and age, and Juniperus procera as codominating tree species. The
canopy of these forests is often closed to nearly 100 %, and the undergrowth mainly
consists of a dense herb and grass layer, whereas higher bushes are rarely found. The
conditions of high competition and intensive shading of the ground are not suitable
for the germination of Hagenia abyssinica. A fire event disturbes the climax
community by destroying herbs and leaf litter in the undergrowth, as well as all
roots in the upper soil layers, whereas the adult Kosso trees remain more or less
severly damaged, but still alive in most cases. Miehe and Miehe (1994) supposed
adult Hagenia trees to be almost fire resistant, as their bark is peeling in large pieces,
preventing a fire from reaching the essential vascular strands of the tree. Supporting
this assumption, many Kosso trees in the forests show sometimes heavy signs of
former burning, but they are still green and even flowering. After a fire, the heated
ground, now bare of vegetation and only covered with black ash, provides ideal
conditions for the germination of the Kosso seeds (2). It depends on the frequency of
fires, whether stage 3 or 4 of the regeneration cycle is developing in a certain area. In
case the time intervals between two fire events are sufficiently high (approximately
100 years), the young Hagenia trees are found scattered between a dense pioneer
vegetation of about 5–6 m height. Clutia abyssinica (Euphorbiaceae), Dombeya
torrida (Sterculiaceae), Helichrysum schimperi (Asteraceae), and Leonotis
mollissima (Lamiaceae) are characteristic species in the impenetrable shrub thicket
of this succession stage after fire. In contrast, more frequent burning results in a shift
to a less high and dense vegetation dominated by Gnidia glauca (Thymeleaceae) (4).
References
Agnew ADQ. Cyclic sequences in plant communities of the Aberdare mountains, Kenya. J East
Africa Nat Hist Soc Natl Mus. 1985;75:1–12.
Baker BH. Geology of the Baragoi Area. Geological Survey of Kenya. Nairobi: Government
printer; 1960.
Barnes RFW. Deforestation trends in tropical Africa. Afr J Ecol. 1990;28:161–73.
Buechner HK, Dawkins HC. Vegetation changes induced by elephants and fire in Murchinson Falls
National Park, Uganda. Ecology. 1961;42:752–66.
management of a tropical mountain forest ecosystem. Dissertation Universität Bayreuth, Bay-
reuth. 1994.
Bussmann RW. Destruction and management of Mt. Kenya’s forests. Ambio. 1996;25(5):314–7.
Bussmann RW. The forest vegetation of the Harenna escarpment (Bale Province, Ethiopia) syn-
taxonomy and phytogeographical affinities. Phytocoenologia. 1997;27(1):1–23.
Bussmann RW. Notes on the vegetation of Menagesha forest (Shoa Province, Ethiopia). In: BITÖK
(Ed.) Vegetationsökologische, ethnobotanische und -faunistische Beiträge über Äquatorial-
Afrika. Bayreuther Forum Ökologie. 1999a;64:105–18.
Bussmann RW. The relation of soil- and vegetation types in the forest belt of Mount Kenya (Kenya).
In: BITÖK (Ed.) Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie. 1999b;64:29–44.
Bussmann RW. Growth rates of important East African montane forest trees, with particular
reference to those of Mount Kenya. J East Afr Nat Hist. 1999c;881–2:69–78.
Bussmann RW. Islands in the desert – forest vegetation of Kenya’s smaller mountains and highland
areas Nyiru, Ndoto, Kulal, Marsabit, Loroghi, Ndare, Mukogodo, Porror, Mathews, Gakoe,
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002;91(1/2):27–79,
appendices 1-7 @. www.naturekenya.org/JournalEANH.htm
Bussmann RW. Succession and regeneration patterns in East African Mountain forests – a review.
Syst Geogr Plants. 2002a;712:959–74.
Bussmann RW. Vegetation ecology and regeneration of Tropical mountain forests. In: Ambasht RS,
Ambasht NK, editors. Modern trends in applied terrestrial ecology: Kluwer; 2002b. p. 195–223.
Bussmann RW. Diversity and use potential of montane forest ecosystems in Kenya. Biodiversity in
conversion: the influence of fragmentation and disturbance on the biodiversity of East African
highland rain forests. BIOTA East Africa, Final Report Phase I 2001-2004, 151–162. 2004a.
Bussmann RW. Regeneration and succession patterns in African, Andean and Pacific Tropical
Mountain Forests: the role of natural and anthropogenic disturbance. Lyonia. 2004b;61:98–111.
2006a;111:41–66.
Bussmann RW. Ethnobotany of the Samburu of Mt. Nyiru, South Turkana, Kenya. J Ethnobiol
Ethnomed. 2006b;2:35.
Phytocoenologia. 1995a;25(5):467–560.
Bussmann RW, Beck E. Regeneration- and cyclic processes in the Ocotea-Forests (Ocotea
usambarensis Engl.) of Mt. Kenya. Verh Ges Ökologie. 1995b;24:35–8.
Bussmann RW, Beck E. Regeneration- and succession processes in the Cedar-Forests (Juniperion
procerae) of Mt. Kenya. Ecotropica. 1995c;1:79–84.
Bussmann RW, Beck E. Vegetation units of Mount Kenya Forest Reserve – comment. In: BITÖK
(Ed.) Vegetationsökologische, ethnobotanische und -faunistische Beiträge über Äquatorial-
Afrika. Bayreuther Forum Ökologie. 1999;64:17–28.
Bussmann RW, Lange S. Succession and regeneration patterns of East African mountain forests. In
BITÖK (ed.) Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie. 1999;64:1–16.
Bussmann RW, Lange SH. Germination of important East African mountain forest trees. J East Afr
Nat Hist. 2000;891–2:101–11.
Bussmann RW, Gilbreath GG, Lutura M, Lutuluo R, Kunguru K, Wood N, Mathenge S. Plant use of
the Maasai of Sekenani Valley, Maasai Mara, Kenya. J Ethnobiol Ethnomed. 2006;2:22.
Bussmann RW, Swartzinsky P, Worede A, Evangelista P. Plant use in Odo-Bulu and Demaro, Bale
region, Ethiopia. J Ethnobiol Ethnomed. 2011;7:28.
Bytebier B, Bussmann RW. Vegetation of Mount Nyiru Samburu District, Kenya: a checklist and
syntaxonomical survey. J East Afr Nat Hist. 2000;891–2:45–71.
Clayton W. Gramineae (part 1). In: Turill WB, Milne-Redhead E, Hubbard CE, Polhill E, editors.
Flora of Tropical East Africa. Rotterdam: Balkema; 1970.
Dale IR, Greenway PJ. Kenya trees and shrubs. Nairobi: Buchanan; 1961.
Doute R, Ochanda N, Epp H. Forest cover mapping in Kenya using remote sensing techniques.
Nairobi: KREMU; 1981.
Edwards KA, Field CR, Hogg IGG. A preliminary analysis of climatological data from the Marsabit
District of Northern Kenya. UNEP Integrated Project on Arid Lands (IPAL) Technical report
B-1. Nairobi: UNEP; 1979.
Friis I. Forests, forest trees of Northeast Tropical Africa. Kew Bulletin additional series XV. London:
HMSO; 1992.
Hedberg O. Features of afro-alpine plant ecology. Acta Phytogeographica Suecica. 1964;49:1–144.
Hedberg O. Vegetation belts of the East African mountains. Svensk Bot Tidskr. 1951;45:140–99.
Holloway CW. The impact of big game on the forest policy in the Mount Kenya and North-East
Aberdare forest reserves. E Afr Agric For J. 1965;30(4):370–89.
Holmgren HP, Masakha EJ, Sjöholm H. Not all African land is being degraded: a recent Survey of
Trees on Farms in Kenya reveals rapidly increasing forest resources. Ambio. 1994;23(7):390–5.
Houston MA. Biological diversity – the coexistance of species on changing landscapes. Cambridge:
Cambridge University Press; 1995.
Jätzold R. Klimageographie, Serie E, Blatt 5, 1:1000000. Berlin/Stuttgart: Bornträger; 1977.
Jätzold R. Klimageographie Ostafrika – Afrika Kartenwerk, Serie E, Beiheft zu Blatt 5. Berlin/
Stuttgart: Bornträger; 1981.
Kenya Meteorological Department. Climatological statistics for Kenya. Nairobi: Kenya Meteoro-
logical Department; 1974. p. 1984.
Kerfoot O. Juniperus procera Endl. (The African Pencil Cedar) in Africa and Arabia – I. Taxonomic
affinities and geographical distribution. E Afr Agric For J. 1961;26:170–7.
Kiefer S, Bussmann RW. The meaning of regeneration strategies and anthropogenic influence for
the forest expansion in East African montane forest ecosystems – a modeling approach. Lyonia.
2004;62:161–9.
Kiefer S, Bussmann RW. Household energy demand and its challenges for forest management in the
Kakamega area, W-Kenya. Ethnobot Res Appl. 2008;6:353–71.
Kortland A. Vegetation research and the “bulldozer” herbivores of tropical Africa. In Chadwick AC,
Sutton SL, editors. Tropical rain-forest. The Leeds symposium. Leeds: Philosophical Society; 1984.
Lange SH, Bussmann RW, Beck E. Ecology and regeneration of the subalpine Hagenia abyssinica
forests of Mt. Kenya. Bot Acta. 1997;110(6):473–80.
Luizza MW, Young H, Kuroiwa C, Evangelista P, Worede A, Bussmann RW, Weimer A. Local
knowledge of plants and their uses among women in the Bale Mountains, Ethiopia. Ethnobot
Res Appl. 2013;11:315–39.
Mäckel R. Oberflächenformung in den Trockengebieten Nordkenyas. In: Büdel J, Busche D,
editors. Studien zur tropischen Reliefbildung. Berlin: Bornträger; 1986. p. 85–225.
Mäckel R, Schultka W. Vegetationsveränderungen und Morphodynamik im Ngare Ndare-Gebiet,
Kenia. In: Hagedorn J, Mensching HG, editors. Aktuelle Morphodynamik und Morphogenese
in den semiariden Randtropen und Subtropen. Abhandlungen der Akademie der Wissenschaften
Göttingen, Mathematisch-Physikalische Klasse 3/41. p. 253–76. 1988.
Mäckel R, Walter D. Die landschaftsökologische Bedeutung der Bergwälder für die Trockengebiete
Nordkenyas. Die Erde. 1983;114:211–35.
Martin RE. Fire history and its role in succession. In: Means JE, editor. Forest succession and stand
development in the Northwest. Corvalis: Forest Research Laboratory, Oregon State University;
1982.
Mason P. Geology of the Meru-Isiolo Area. Nairobi: Government Printer; 1963.
Miehe S, Miehe G. Ericaceous Forests and Heathlands in the Bale Mountains of South Ethiopia. In:
Ecology and man`s impact. Hamburg: Stiftung Walderhaltung in Afrika und Bundesforschung-
sanstalt für Forst- und Holzwirtschaft; 1994.
Schmitt K. The vegetation of the Aberdare National Park, Kenya Hochgebirgsforschung
1991;7:1–250. Innsbruck: Verlag Wagner.
Schmitt K, Beck E. Die Zerstörung eines Bergwalds in Kenia: Beispiele des Zusammenwirkens von
Mensch und Großwild. Jahrb Verein z Schutze der Bergwelt. 1992;57:207–26.
Shackleton RM. Geology of the country between Nanyuki and Maralal. Geological Survey of
Kenya, publication 11. Nairobi: Survey of Kenya; 1946.
Speck H. Mount Kenya Area. Bern: Lang; 1983.
Speck H. Soils of the Mount Kenya Area, their formation, ecological, and agricultural significance.
In: Winiger M, editor. Mt. Kenya Area. Contributions to ecology and ocio-economy. African
studies series A1; 1986. p. 47–72.
Troll C. Die tropischen Gebirge. Ihre dreidimensionale klimatische und pflanzengeographische
Zonierung. Bonner Geographische Abhandlungen. 1959;25:1–93.
White F. The Afromontane Region. In: Werger MJA, editor. Biogeography and ecology of Southern
Africa. The Hague: Junk; 1978. p. 463–513.
White F. The vegetation of Africa. Paris: UNESCO; 1983.
Willan RL. Natural regeneration of high forest in Tanganyika. E Afr Agric For J. 1965;31:43–53.
Wimbush SH. Natural succession in the Pencil Cedar forests of Kenya Colony. Empire For
J. 1937;16:49–53.
Wimbush SH. The African alpine Bamboo. Empire For J. 1945;24:33–9.
Winiger M. Bodentemperaturen und Niederschlag als Indikatoren einer klimatisch-ökologischen
Gliederung tropischer Gebirgsräume. Geomethodica. 1979;4:121–50.
Part II
Plants
Abutilon sp.
Abutilon angulatum (Guill. & Perr.) Mast.
Abutilon grandiflorum (Will.) Sweet.
Abutilon longicuspe Hochst. ex A. Rich.
Abutilon mauritianum (Jacq.) Medik.
Hibiscus sp.
Hibiscus calyphyllus Cav.
Hibiscus vitifolius L.
MALVACEAE

Grace N. Njoroge
Synonyms
Abutilon angulatum (Guill. & Perr.) Mast.: Abutilon intermedium Hochst ex Garcke;
Abutilon etveldeanum De Wild. & T. Durand; Bastardia angulata Guill. & Perr.
Abutilon grandiflorum (Will.) Sweet.: Abutilon tortuosum Guill. & Perr.; Sida
grandifolia Willd.
Abutilon mauritianum (Jacq.) Medik.: Abutilon longipes Mattei; Abutilon
zazibaricum Biojer ex Mast; Pavonia patens (Andrews) Chiov.; Sida
mauritiana Jacq.
Hibiscus calyphyllus Cav.: Hibiscus calycinus Willd.
Hibiscus vitifolius L.: Hibiscus jatrophifolius A. Rich.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Abutilon angulatum: Shambaa: Fiefie; English: Elephant’s ear, Fluted Abutilon

(Kokwaro 2009).
Abutilon hirtum: Gabbra: Matawi; Maa: Olorerirepi; Pokot: Kapiyan; Samburu:
Sulubei, Sulube; Turkana: Etoo, Asrilipong, Ekwangat (Beentje 1994; Gachathi
1989; Kokwaro 2009).
Abutilon grandiflorum: Luganda: Kifura; English: Hairy abutilon, Hairy Indian
mallow (Kokwaro 2009)
Abutilon mauritianum: English: Bush mallow, Country mallow, French: Mauve
des champs; Shambaa: Fiefie; Bende: Ikolelitambo; Samburu: Sulubei; Kikuyu:
Mukeu, Mwondwe, Mwondwe-ndogo; Marakwet: Jeptula; Pokomo: Malasa;
Digo: Chibangula mavi, Mbagula mavi; Swahili: Maumanda, Mjamanda, Mbiha,
Mopoahabari, Mpamba pori (Beentje 1994; Gachathi 1989; Kokwaro 2009).
Abutilon longicuspe: Kikuyu: Mondwe, Mwondwe; Ndorobo/Kipsigis:
Lolodongoiet; Kisii: Omorovianda; Maa: Osupikioi-oibor; Pokot: Cheptonge;
Taita: Muklundkunda, Turkana: Ekwanga; Samburu: Sulubei; Sulube; Swahili:
Mbiha (Beentje 1994; Gachathi 1989; Kokwaro 2009).
Hibiscus sp.: Samburu: Likimanshoi (Kokwaro 2009).
Hibiscus aponeurus: Luo: Omen; Luhya: Llundu, Lukhambi; Maa: Enkarani
(Kokwaro 2009).
Hibiscus calyphyllus: Maa: Osukupai naibor, Esubukioi narok; Digo: Kungala;
Kipsigis: Leltangoyet, Leledonget; Pokot: Kapenyan; Samburu: Sulube; Tugut:
O’tanya; Turkana: Nauru-kasiko; English: Lemon-eyed rose mallow (Beentje
1994; Gachathi 1989; Kokwaro 2009).
Hibiscus fuscus: Chagga: Kirundu; Meru: Mukluma; Kikuyu: Mugere;
Samburu: Lokimekii, Lokimek ol idonyo (Beentje 1994; Gachathi 1989; Kokwaro
2009).
Hibiscus ludwigii: Samburu: Lkimanshoi.
Hibiscus vitifolius: Kikuyu: Mugere; Samburu: Likimanshoi, nakodonyyo; Maa:
Osukupai naibor; English: Lemon-eyed rose mallow (Beentje 1994; Gachathi 1989;
Kokwaro 2009).
Botany and Ecology
Abutilon sp.: Biennial to perennial (rarely annual) erect or occasionally spreading

herbs or shrubs, variously pubescent, usually with stellate hairs. Leaves petiolate,
usually more or less ovate in outline with cordate base. Flowers generally yellow to
orange, rarely white, mauve or purple, small to medium-sized, axillary, solitary or
fascicled, rarely 2–4-nate on a common peduncle, sometimes on short axillary leafy
side-shoots, sometimes aggregated in terminal and lateral leafy pseudo-panicles;
pedicels usually articulated in the upper half often near the apex. Epicalyx absent.
Calyx with a cupular to campanulate tube; lobes 5, distinct, semi-orbicular to
lanceolate, usually acute to acuminate. Petals 5, connate at the base and adnate to
Abutilon sp. . . . 71
the base of the staminal tube, usually conspicuously longer than the calyx and in
open flowers usually spreading to rotate, generally obovate with a narrow sub-
unguiculate often ciliate basal portion. Staminal tube divided at the apex into
many filaments, dilated below, glabrous or stellate-pubescent; free parts of filaments
terete; anthers reniform. Carpels 5–40, 3–9-ovulate, in a circle around a distinct torus
and joined to form a subglobose gynoecium; style-branches as many as the carpels,
terete, filiform or clavate; stigmas simple to somewhat capitate. Fruit subglobose or
turbinate to hemispherical or almost disk-shaped, often truncate, depressed or
umbilicate at the apex; mericarps 5 to many, laterally compressed, follicular,
(1) 2–3 (9)-seeded, separating from the ultimately conical or subcylindric and
usually more or less produced or dilated to capitate torus and usually dehiscing by
the ventral suture, ultimately grey or brown to black, oblong to subrectangular,
reniform or more or less semi-orbicular, rounded to truncate at the base and rounded,
truncate or acute at the apex, muticous to mucronate, apiculate or awned at the upper
dorsal (outer) angle or at the apex, the ventral side with a usually distinct retrorse
tooth which originally fitted over and against the apex of the torus. Seeds reniform,
often unequally so and more or less comma-shaped, glabrous, puberulous or stellate-
tomentose, smooth, finely pitted or minutely papillose to verruculose; embryo
curved; cotyledons folded; endosperm scanty. Abutilon comprises 100–150 species
and is distributed in the tropics and subtropics. There is a need for further taxonom-
ical study as the circumscription of several species is obscure. Many species are a
component of East African Mountain forests (Bussmann 2002; Verdcourt and
Mwachala 2009).
Abutilon angulatum (Guill. & Perr.) Mast.: Perennial herb or shrub up to 5 m tall;
stem and branches angular; all parts greyish hairy, with very short stellate hairs and
without long simple hairs. Leaves alternate, simple; stipules subfalcate, 4 mm long,
reflexed, caducous; petiole 1–12(–24) cm long; blade roundish to broadly ovate, up
to 30 cm 26 cm but usually much smaller, cordate at the base, acute to acuminate
at the apex, margin toothed, upper surface dark grey-green, lower surface much
paler, 5–9-veined from the base. Inflorescence a large terminal and lateral panicle,
loose, much branched, ultimately leafless; peduncle 1–5 cm long, accrescent.
Flowers bisexual, regular; pedicel 1–5 cm long; epicalyx absent; calyx
cup-shaped, 6–15 mm long, 5-lobed, lobes ovate, acute, submucronate; petals
5, united at the base and adnate to the base of the staminal column, 9–22 mm
long, yellow to orange; staminal column 5–8 mm long, filaments 2–5 mm long;
ovary superior. Fruit a subglobose schizocarp of follicle-like mericarps, 10–
12 mm 8–9 mm, depressed, umbilicate; mericarps 20–40, 7–9 mm 5–6 mm,
rounded or with an obtuse dorsal angle, 1-seeded. Seeds c. 2.5 mm long, dark brown,
smooth to finely papillose, glabrous. In southern Africa flowering is in December–
May. Flowers open in the afternoon. A component of East African Mountain forests
(Bussmann 2002; Verdcourt and Mwachala 2009).
Abutilon grandiflorum (Will.) Sweet: Perennial herb or shrub up to 3 m tall;
branches covered with long and slender hairs. Leaves alternate, simple; stipules
subulate, caducous; petiole 5–20 cm long; blade ovate, up to 20 cm 15 cm, base
cordate, apex acute or subacuminate, margin toothed, 6–7-nerved, both surfaces
stellate hairy. Inflorescence axillary, 1–2-flowered; peduncle 4–5 cm long, up to

12 cm in mature fruit. Flowers bisexual, regular; epicalyx absent; calyx 5-lobed, 1–
1.5 cm long, accrescent, with broadly oval segments; petals 5, united at the base and
adnate to the base of the staminal column, yellow; stamens many, staminal column
very short; ovary superior. Fruit a subglobose schizocarp of follicle-like mericarps;
mericarps 8–10, oblong, 12 mm 7 mm, with an ascendant awn 1.5–2 mm long,
hairy, tardily separating, 2–5-seeded. Abutilon grandifolium is native to South
America. It is cultivated in the tropics and has widely become naturalized (Verdcourt
and Mwachala 2009).
Abutilon longicuspe Hochst. ex A. Rich.: Much-branched shrub up to 6 m tall; all
parts tomentellous, with or without long simple hairs. Leaves alternate, simple;
stipules linear, 6 mm long; petiole 2–19 cm long; blade broadly ovate up to
22 cm 18 cm, cordate at the base, acuminate to cuspidate at the apex, margin
toothed, upper surface dark green, lower surface much paler, both surfaces stellate-
velvety, palmately 5–7-veined. Inflorescence a terminal or lateral panicle up to
23 13 cm, many-flowered. Flowers bisexual, regular; pedicel 0.5–3(–4) cm
long; epicalyx absent; calyx 4–12 mm long, persistent, broadly cup-shaped,
5-lobed, segments acute; petals 5, free, 8–20 mm long, white to pink with purplish
center; stamens many, staminal column cylindrical, 2–7 mm long, purple, filaments
4–12 mm long; ovary superior. Fruit a subglobose schizocarp of follicle-like
mericarps, c. 9 mm 13–15 mm, umbilicate, downy; mericarps c. 20, reniform,
6–10 mm 4–7 mm, obtuse, 1-seeded. Seeds c. 3 mm 2 mm, smooth, glabrous.
Abutilon longicuspe is distributed from Sudan, Eritrea, and Ethiopia through DR
Congo and East Africa southward to Angola, Zimbabwe and Mozambique. It also
occurs in Egypt and Yemen. A component of East African Mountain forests
(Bussmann 2002; Verdcourt and Mwachala 2009).
Abutilon mauritianum (Jacq.) Medik.: Perennial herb or shrub up to 2.5(–4) m tall;
all parts pubescent or tomentose and sometimes also with long simple hairs; stems
glandular. Leaves alternate, simple; stipules linear-subulate, 4–9 mm 0.5 mm;
petiole up to 20 cm long; blade ovate to suborbicular, up to 18(–25) cm 16(–18)
cm, cordate at the base, acuminate at the apex, margin toothed; upper surface grey-
green, lower surface paler. Flowers solitary in leaf axils or on short axillary shoots,
bisexual, regular; pedicel 1.5–8(–12) cm long; epicalyx absent; calyx bell-shaped to
cup-shaped, 5-lobed, 6–22 mm long; petals 5, united at the base and adnate to the
base of the staminal column, 12–33(–38) mm 10–15(–25) mm, yellow, orange or
reddish; staminal column 5–7 mm long, filaments 3–5 mm long; ovary superior.
Fruit a subglobose schizocarp of follicle-like mericarps; mericarps 20–40, 10–
17 mm 4–6.5 mm, with a 3–5 mm long awn, ultimately stellately spreading,
black, 2–3 seeded. Seeds c. 2.5 mm long, dark brown, papillose, spiny-papillose
towards hilum. Abutilon mauritianum is widely distributed in the drier parts of
tropical Africa from Senegal eastward to Eritrea, Ethiopia, and Somalia, and south-
ward to Angola, Zambia, Zimbabwe Mozambique, and South Africa. The species
also occurs in the Comoros and the Seychelles and is recorded for Mauritius. A
Mwachala 2009).
Hibiscus sp.: Annual or perennial herbs (sometimes with annual shoots arising from
woody rootstocks or underground stems), shrublets, shrubs, or (rarely) small trees.
Fruit a loculicidally dehiscent capsule. Capsule winged or with very prominent
angles. Leaves petiolate, simple, lobed or digitately compound. Flowers usually
solitary, axillary, and often forming terminal racemose or corymbose inflorescences
by reduction of the upper leaves, medium to large, often yellow with a dark center or
red, pink, purplish or white; peduncle usually articulated. Epicalyx (occasionally
absent) of 5–20 bracts very variable in shape and length, free or adnate to the base of
the calyx. Calyx 5-merous, usually with 5 lobes (or rarely 5 or 10 teeth) joined at the
base or more rarely almost to the apex, persistent. Staminal tube truncate at the apex,
free parts of filaments very variable in length, sometimes whorled. Ovary 4–5-
locular; loculi 3 ovulate; style usually 5-branched. Many species are found as a
Mwachala 2009) (Figs. 1 and 2).
Hibiscus calyphyllus Cav.: Staminal tube up to 15 mm long; free parts of filaments
1.5–3 mm long. Petals up to 6 4.5 cm, obliquely obovate, pubescent outside,
Fig. 1 Hibiscus aponeurus

(Malvaceae), Maasai Mara,
Kenya. (Photo
R.W. Bussmann)
Fig. 2 Hibiscus aponeurus

Kenya. (Photo
R.W. Bussmann)
glabrous within. Calyx up to 16 mm long, stellate-tomentellous; lobes ovate to

ovate-lanceolate, usually 3-nerved, joined to nearly half-way. Epicalyx of 5 bracts,
stellate-pubescent; bracts up to 18 mm long, varying greatly in shape but nearly
always broadest near the middle then narrowed suddenly to a caudate tip which is a
prolongation of the midrib, joined for 3–4 mm at the base. Seeds 3 2.5 mm,
subreniform, tomentellous. Capsule 25 15 mm, ellipsoid; valves aristate. Style-
branches 4–5 mm long. Flowers up to 12 cm in diam., yellow usually with a
brownish or dark red centre, solitary, axillary; peduncle c. 7 (10) mm long,
stellate-pubescent, usually rather inconspicuously articulated near the base. Leaf-
lamina up to 12 12 cm, suborbicular in outline, obscurely or distinctly 3–5-lobed,
stellate-pubescent or stellate-pilosulose above, stellate-pubescent or stellate-pilose
or stellate-tomentose beneath, apex acute, margin serrate, base cordate, or sub-
cordate; petiole usually up to 9 (18) cm long, stellate-pilose; stipules up to 15 mm
long, filiform or subsetaceous, somewhat expanded at the base. Shrub or perennial
herb up to 3 m tall; stems tomentose or pubescent when young, later glabrescent
(Verdcourt and Mwachala 2009) (Fig. 3).
Fig. 3 Hibiscus calophyllos

Kenya. (Photo
R.W. Bussmann)
Hibiscus vitifolius L.: Shrub or perennial herb up to 1.5 m. tall, sometimes scandent
or scrambling; stems terete, nearly glabrous or with a very variable indumentum,
sometimes tomentose or tomentellous, often glandular, sometimes hispid or stellate-
hispid or aculeate and with these different types of hairs occurring in various
combinations. Leaf-lamina 3–15 3–15 cm, ovate to suborbicular; not lobed to
deeply 3–5 (7)-palmatilobed, with a variable indumentum (as on the stems) but
stellatehispid or bifurcate hairs, rather rare on the stems, are more frequent especially
on the under surface, margin serrate, base truncate to cordate 5–9-nerved; lobes acute
or bluntish; petiole 2–7 (18) cm long, hairy like the stems; stipules 3–5 mm long,
filiform. Flowers 5–9 cm in diam., yellow with a dark reddish or purple center,
solitary or forming terminal racemes by reduction of the upper leaves; peduncle 1–
1.5 (5) cm long, articulated above the middle. Epicalyx of c. 10 bracts; bracts 8–
10 mm long, filiform. Calyx up to 15 mm long in flower, accrescent, somewhat
scarious; lobes up to 15 10 mm, broadly elliptic, acuminate, joined to about half-
way. Petals up to c. 5 4.5 cm, glabrous or nearly so. Staminal tube up to 15 mm
long; free parts of filaments up to 5 mm long. Style-branches 1.5–3 mm long,
minutely glandular. Seeds 2.5 1.5 mm, irregularly reniform, glabrous. Capsule
10–12 12–15 mm winged, scarious, usually hispid; valves aristate (Verdcourt
and Mwachala 2009) (Figs. 4 and 5).
Local Medicinal Uses
Abutilon angulatum: In southern African traditional medicine the powdered root is

applied on burns. Root extracts are drunk for the treatment of cough and pneumonia,
and for easing labor pains. An infusion of the root forms part of preparations taken in
case of epilepsy and heartache. Cooked or raw leaves are eaten as a remedy for
hiccups. In Madagascar the flowers are made into a lotion applied in case of skin
problems. The root decoction is drunk as cough syrup, and to ease labor pain
(Kokwaro 2009). In Ethiopia, the root is chewed for chest pain (Teklehaymanot
and Giday 2010).
Fig. 4 Hibiscus
sp. (Malvaceae), Maasai
Mara, Kenya. (Photo
R.W. Bussmann)
Abutilon grandifolium: In Burkina Faso, decoctions of leafy or fruiting stems are

applied as enema for the treatment of measles, and the leaves and stems are used
internally and externally for the treatment of insect bites. In Uganda used as
abortificant. Root pieces used for conjunctivitis and worn to fend off magic
(Kokwaro 2009).
Abutilon longicuspe: In traditional medicine in Kenya, the plant is used to ease
childbirth and to expel the placenta. The root is used in Tanzania for the treatment of
stomach problems.
Abutilon mauritianum: Widely used in traditional African medicine. In Nigeria an
infusion of the root is taken as cooling drink in case of fever. In East Africa the root is
chewed as an expectorant, a root decoction is drunk against bronchitis and cramp in
the stomach, and a decoction of the root and bark for the treatment of diarrhea,
stomachache, coughs, and colds. In Ethiopia, the crushed fresh root is boiled and
taken orally in case of snake bites. In Nigeria, a bark extract is taken as a diuretic. In
Benin a decoction of leafy twigs is taken for the treatment of dysentery. In Nigeria,
leaves are used as rectal suppository to relieve soreness of the rectum in babies and
young children. In Tanzania, leaves are pounded to a pulp and applied on scabies. An
infusion of the macerated leaves is drunk in Nigeria for the treatment of venereal
diseases such as gonorrhea. A decoction of the leaves is taken as a gargle for sore
Fig. 5 Hibiscus sabdariffa

(Malvaceae), dried flowers,
Batumi Bazar, Georgia.
(Photo R.W. Bussmann &
N.Y. Paniagua-Zambrana)
throat and as a fomentation to relieve pain in dysentery and hemorrhoids. The leaf
sap is drunk in Tanzania against coughs and colds; it is also taken against malaria. In
Gabon, mucilage from the leaves is used in poultices applied on wounds to facilitate
the extraction of foreign bodies such as splinters. A mucilaginous extract from the
leaves is used as demulcent. In Uganda the head is washed with a leaf maceration
against headache, and the leaves are used against ascariasis. Leaf decoctions are
taken against diarrhea, gonorrhea, inflammation, bronchitis, and catarrh. The pow-
dered fruit is used in preparations used for the treatment of eczema in Tanzania.
Seeds are used against cough in Nigeria. In Kenya the species is said to be used to
ease childbirth and to expel the placenta. In Benin the fresh plant is ground with
kaolin in water and drunk against dysentery. In the DR Congo an infusion of the
whole plant is orally or rectally administered for the treatment of asthma and as a
tranquilizer. The leaves are used as bandage on open wounds and as toilet paper. The
leaf infusion is used for diarrhea. The root infusion to treat stomach cramp, bron-
chitis, diarrhea, colds, and cough (Kokwaro 2009). Abutilon mauritianum is also
used in Kenya for wound treatment (Njoroge et al. 2004). In Ethiopia the fresh root is
crushed and boiled and taken orally for snakebite (Teklehaymanot 2009).
Abutilon greveanum and Abutilon pseudocleistogamum: The sap of their flowers

is Abutilon hirtum, A. longicuspe, and A. mauritianum are used by the Samburu to
expel the placenta after birth, and is put on wounds to promote cicatrization
(Bussmann 2006).
Abutilon pseudocleistogamum: The leaves of are chewed as a hemostatic. The
decoction of the leaves is credited with oxytocic properties.
Abutilon indicum serves as abortifacient, aphrodisiac, to treat boils, bone fractures,
bronchitis, childbirth complications, colic, convulsions, as cooling agent, for cough,
diarrhea, dysentery, gonorrhea, headache, leprosy, paralysis, piles, as purgative, for
stomach complaints, ulcers, arthritis, and wounds (Verma et al. 2007).
Hibiscus fuscus serves for wound treatments and burns (Bussmann 2006; Njoroge
et al. 2004). Roots are used to treat chest problems. The juice from the stems is used
to induce childbirth (Gachathi 1989). Burned leaves and root are mixed with fat and
applied to wounds and burns (Kokwaro 2009).
Hibiscus rosa-sinensis is used in India for menorrhagia, dysentery, fever, headache,
burn, boil, skin disease, cough and cold, fatigue, hair fall, abortion, burning sensa-
tion in the stomach, twitching, and gastric problems (Raj et al. 2018). The leaf juice
is also used to treat infertility in women and irregular menstruation, while the root
extract as mouthwash (Debbarma et al. 2017).
Hibiscus calyphyllus and Hibiscus vitifolius: In Kenya and Tanzania, the leaves are
applied to wounds as a dressing. The vapor of boiled roots is inhaled, and the
decoction drunk to treat pneumonia. The leaves of Hibiscus calyphyllus are applied
to sores. The roots are boiled, and the steam inhaled and the decoction drunk to treat
pneumonia (Kokwaro 2009).
Local Food Uses
Abutilon angulatum: The leaves and flowers are cooked and eaten as a vegetable.
Hibiscus calyphyllus and Hibiscus vitifolius: The leaves are eaten as a vegetable,
especially in East Africa. They are collected from the wild during the rainy season,
wilted, chopped, and boiled mixed with other coarse vegetables. This vegetable is
eaten frequently locally, but in small amounts.
Local Handicraft and Other Uses
Abutilon species used as fiber plants in Madagascar include Abutilon greveanum

and Abutilon pseudocleistogamum, both endemic to Madagascar. Their fibers are
made into cordage. Abutilon angulatum: The stem bark yields a fiber locally used
for string and cordage. In Zambia, the fiber is used for stitching mats together. In
Madagascar the bark fiber is woven into cloth. The large and soft leaves are
sometimes used as toilet paper. The leaves and the inner tissue of the stems are
mixed with tobacco and smoked. Leaves are used as fodder. The stems are used as
tinder. Bees feed on the flowers. The plant has ornamental value.
Abutilon grandiflorum: cultivated in the tropics as a fiber plant and as an ornamen-

tal. Abutilon longicuspe: In Kenya, the stem is locally used to make string and
withies, e.g., for making baskets. In Tanzania, it yields string for building houses. In
Ethiopia the stem bark is used as rope. The cooked flowers are eaten as a vegetable.
The plant is also used as fodder, as ornamental and as a source of bee forage.
Abutilon mauritianum: The stem bark yields a strong fiber which is locally made
into string or rope, used for instance in fishing and for making hammocks. The bark
is used for making baskets in Kenya. Pregnant women in Gabon carry around the
waist a cord made of the fiber in the belief that it eases delivery. Wood from the stem
is used for cleaning teeth and the whole twigs are made into brooms. The leaves and
flowers are cooked and eaten as a vegetable. The seeds are eaten by nomadic people.
The plant is also used as fodder, as an ornamental and as a source of bee forage. In
Gabon, a mucilaginous extract from the bark is used as mordant in dyeing.
Abutilon hirtum, Abutilon longicuspe, and Abutilon mauritianum are browsed by
lifestock (Bussmann 2006). Used to extract fibers (Gachathi 1989).
Hibiscus calyphyllus and Hibiscus vitifolius: In Uganda, poles made from the stems
are used for building. The best fiber is made into rope in Uganda and Tanzania.
Hibiscus aponeurus is used in ceremonies by the Maasai, e.g., to cast curses, while
Hibiscus calyphyllus is used for construction (Bussmann et al. 2006, 2018).
Hibiscus sp., Hibiscus fuscus, Hibiscus ludwigii, and Hibiscus vitifolius are eaten
by wild animals but rarely by livestock. The bark can be used to make strings to join
sticks, and in ceremonial settings to make protective strings or burnt to mark the
body with ash (Bussmann 2006). The stems are used to weave winnowing trays
(Gachathi 1989).
In Ethiopia, various species of Hibiscus are used to make ropes used to tie posts in
construction. The species are also browsed by cattle and other livestock (Bussmann
et al. 2011). Often planted as ornamental (Mekonen et al. 2015).
References
Beentje H. Kenya trees and shrubs. Nairobi: National Museums of Kenya; 1994.
areas (Nyiru, Ndoto, Kulal, Marsabit, Loroghi, Ndare, Mukogodo, Porror, Mathews, Gakoe,
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi). J East Afr Nat Hist. 2002;91(1/2):27–79,
appendices 1–7. www.naturekenya.org/JournalEANH.htm
Ethnomed. 2006;2:35.
Bussmann RW, Paniagua-Zambrana NY, Wood N, Ole Njapit S, Ole Njapit JN, Ene Osoi GE, Kasoe
SP. Knowledge loss and change between 2002 and 2017 – a revisit of plant-use of the Maasai of
Sekenani Valley, Maasai Mara, Kenya. Econ Bot. 2018;72(2):207–16.
Debbarma M, Pala NA, Kumar M, Bussmann RW. Traditional knowledge of medicinal plants in
tribes of Tripura, Northjeast India. Afr J Tradit Complement Altern Med. 2017;14(4):156–68.
Gachathi FN. Kikuyu botanical dictionary. Nairobi: AMREF; 1989.

Kokwaro JO. Medicinal plants of East Africa. Nairobi: University of Nairobi Press; 2009.
Mekonen T, Giday M, Kelbessa K. Ethnobotanical study of homegarden plants in Sebeta-Awas
District of the Oromia Region of Ethiopia to assess use, species diversity and management
practices. J Ethnobiol Ethnomed. 2015;11:64.
Njoroge GN, Barbara G, Bussmann RW, Newton LE, Ngumi VW. Utilization of weed species as
source of traditional medicines in Central Kenya: optimizing resource efficiency (R.U.E.) in
agro-ecosystems. Lyonia. 2004;7(2):71–87.
Raj AJ, Biswakarma B, Pala NA, Shukla G, Vineeta V, Kumar M, Chakravarty S, Bussmann
RW. Indigenous uses of ethno-medicinal plants among forest dependent communities of
Northern Bengal, India. J Ethnobiol Ethnomed. 2018;14(1):8. https://doi.org/10.1186/s13002-
018-0208-9.
Teklehaymanot T. Ethnobotanical study of knowledge and medicinal plants use by the people in
Dek Island in Ethiopia. J Ethnopharmacol. 2009;124:69–78.
Teklehaymanot T, Giday MM. Quantitative ethnobotany of medicinal plants used by Kara and
Kwego semi-pastoralist people in lower Omo River Valley, Debub Omo Zone, Southern
Nations, Nationalities and Peoples Regional State, Ethiopia. J Ethnopharmacol.
2010;130:76–84.
Verdcourt B, Mwachala G. Flora of tropical East Africa: Malvaceae. Kew: Royal Botanic Gardens;
2009. 170 pp. ISBN: 978-1-84246-189-1.
Verma A, Kumar M, Bussmann RW. Medicinal plants in an urban environment: the medicinal flora
of Banares Hindu University, Varanasi, Uttar Pradesh. J Ethnobiol Ethnomed. 2007;3:35.
Acacia drepanolobium Harmes ex Sjostedt
Acacia nilotica (L.) Willd. ex Delile
Acacia senegal (L.) Willd.
Acacia seyal Delille
Acacia tortilis (Forssk.) Hayne
FABACEAE

Grace N. Njoroge
Synonyms
Acacia drepanolobium Harmes ex Sjostedt: Acacia formicarum Harms, Acacia

lathouwersii Staner, Vachellia drepanolobium (Harms ex Y. Sjöstedt) P.J.H. Hurter
Acacia nilotica (L.) Willd. ex Delile: Acacia adansonii Gioll. & Perr., Acacia
arabica (Lam.) Willd.) Acacia nilotica subsp. adansonii (Guill. & Perr.) Brenan,
Acacia scorpioides (L.) W. Wight., Acacia vera Willd., Mimosa arabica Lam.,
Mimosa nilotica L., Mimosa nilotica Thunb., Mimosa scorpioides L., Vachellia
nilotica (L.) P.J.H. Hurter & Mabb.
Acacia senegal (L.) Willd.: Mimosa senegal L., Mimosa senegalensis Houtt.,
Senegalia senegal (L.) Britton
Acacia seyal Delille: Acacia giraffa Sieber ex Steud., Acacia giraffae Hochst. ex
A. Rich., Acacia stenocarpa Hochst. ex A. Rich, Vachellia seyal (Delille)
P.J.H. Hurter
R. W. Bussmann (*)
G. N. Njoroge

Acacia tortilis (Forssk.) Hayne: Acacia perottettii Steud., Mimosa tortilis Forssk.,
Vachellia tortilis (Forssk.) Galasso & Banfi
Local Names
Acacia drepanolobium: Kamba: Kiunga, Muuga, Kipsigis: Mugurit, Luo: Adugo,

Gugna, Maa: Eluai, Redile: Fulai, Samburu: Luai, Rangau, Tugen: Ngowo,
Turkana: Eiyellel, Kamba: Kiunga (Beentje 1994; Kokwaro 2009).
Acacia nilotica: Bajun: Ntetwe, Borana: Burguge, Digo: Chigundigundi, Duruma:
Mgundi, Gabbra: Burkuke, Giriama: Munga, Kamba: Musemei, Kisemei, Kipsigis:
Chepitet, Maa: Ol-Kiloriti, Ol-Erb at, Ol-Kiroriti, Marakwet: Ngobgwa, Oromo:
Chalado, Pokot: Opokwo, Rendile: Gilorit, Mirgi, Samburu: Lokoloriti, Eldekeki,
Somali: Tugerr, Tugen: Chebiwo, Turkana: Ekapelimen, Swahili: Mjungu, Msemehi,
Mgunga, Mtetewe, Gogo: Chigundigundi, Mfuku, Shambaa: Mopira, English: Babul
acacia, Scented thorn, Scented-pod acacia (Beentje 1994; Kokwaro 2009).
Acacia senegal: Bajun: Mtengewa, Borana: Dimitu, Idado, Digo: Chikwata,
Gabbra: Iddaado, Kamba: Mung’ole, Musemei, Luo: Kiluor, Otiep, Maa:
Olderekesi, Oromo: Bura-dima, Sebonah, Pokot: Chemagayan, Rendile: Mirgi,
Samburu: Eldekeki, Lolderekesi, Somali: Adageti, Ethad-geri, Turkana: Ekunoit,
Swahili: Aiti, Kikwata, Mgunga, Acholi: Achika, Teso: Epujait, Luhya:
Kimunyenya, Chagga: Mwera, Nyankore: Omugando, English: Gum tree, Gum
arabic tree, Three-thorned acacia (Beentje 1994; Kokwaro 2009).
Acacia seyal: Borana: Wachu-Adi, Wacho-Dima, Gabbra: Waac’c’u, Kamba:
Kinyua, Kisewa, Kipsigis: Mugurt, Chepkomon, Luo: Ali, Maa: Oleregat, Ole-Rai,
Ol-Munishui, Marakwet: Reno, Pokot: Rena, Rendile: Fulai, Samburu: Lerai,
Somali: Fullai, Jikh, Tugen: Lelnet, Turkana: Ekoromait, Swahili: Mgunga, English:
White galled acacia, Whistling thorn, Shittim wood (Beentje 1994; Kokwaro 2009).
Acacia tortilis: Borana / Ilelewa: Dadacha, Gabbra: Daddacha, Ilelewa: Dadwota,
Kamba: Kilaa, Maa: Ol-Gorete, Ol-tepesi, Ol-Entepesi, Oromo: Gudis, Dabaso,
Dadech, Pokot: Ses, Rendile: Dahar, Samburu: Ltepes, Somali: Abak, Kura,
Tugen: Sietsiet, Seseya, Turkana: Ewoi, Ettior, Teso / Turkana: Etir, Swahili:
Munga, Mgunga, English: Whistling thorn (Beentje 1994, Kokwaro 2009).
Botany and Ecology
Acacia drepanolobium Harmes ex Sjostedt: Bush or small tree 1–5(–7.5) m high,

with short radiating branches from main stem, sometimes spreading at top. Young
branchlets shortly pubescent to puberulous, rarely glabrous, grey then going brown,
no powdery inner bark on twigs. Old bark black or grey, usually rough, sometimes
smooth. Stipules spinescent, mostly 1.5–4.5 (–7.5) cm long (some shorter ones often
also present), straight, grey or whitish, some fused at base into round “ant-galls”
1–3.5 cm in diameter, grape-purple going blackish. Petiole 2–5 (very rarely to 10)
mm long, glandular at the lowest of the 3–13 pairs of pinnae, rachis 0.8–4.5(–9) cm
Acacia drepanolobium Harmes ex Sjostedt. . . 83
Fig. 1 Acacia abyssinica

(Fabaceae), Bale Mountain
National Park, Demaro,
Ethiopia. (Photo
R.W. Bussmann)
long, glandular between the top 1–6 pairs of pinnae, leaflets 11–22 pairs, glabrous or
minutely ciliolate, subacute or acute sometimes obtuse at apex, 1.5–5.5 mm long,
0.7–1.25 (–1.75) mm wide. Flowers white or sometimes cream, in heads, involucel at
or rarely a short way above base of the glabrous puberulous or shortly pubescent
peduncle. Calyx 0.75–1.5(–2.5) mm, long, glabrous or ciliolate. Corolla glabrous
outside, sometimes puberulous on lobes, 3–4 mm long. Pods falcate or annular, thinly
coriaceous, finely longitudinally venose, glabrous, or puberulous, mostly attenuate or
even acuminate at ends, 4–7 cm long, 0.5–1.0 cm wide. Seeds mottled whitish-grey
and dark brown, smooth, irregularly quadrate or elliptic, compressed, 10–12 mm long,
4.5–5.5 mm wide, areole 5–6 mm long, 3–3.5 mm wide. Shrub or dwarf-tree grass-
land, “gregarious, usually on alkaline hard-pan grey soils with Lannea humilis and
Commiphora schimperi, or in fringing ‘mbuga’, or on dark clay cracking lime-
accumulating soils” 600–2680 m (Brenan 1959) (Figs. 1, 2, 3, and 4).
Acacia nilotica (L.) Willd. ex Delile: Tree (1.2)2.5–14 m tall, bark on trunk rough,
fissured, blackish or grey or brown, neither powdery nor peeling, young branchlets
from almost glabrous to subtomentose, glands inconspicuous or absent, bark of
twigs not flaking off, grey to brown. Stipules spinescent, up to 8(11) cm long,
straight or almost so, often deflexed, “ant-galls” and other prickles absent. Leaves
often with 1 (2) petiolar glands and other glands between all or only the topmost of
the 2–11(17) pairs of pinnae, leaflets 7–25(30) pairs, 1.5–7 0.5–1.5 mm, glabrous
to pubescent, not spinulose-mucronate at the apex, lateral nerves invisible beneath.
Fig. 2 Acacia abyssinica

(Fabaceae), Bale Mountain
National Park, Demaro,
Ethiopia. (Photo
R.W. Bussmann)
Fig. 3 Acacia
drepanolobium (Fabaceae),
samburu, Kenya. (Photo
R.W. Bussmann)
Flowers bright yellow, in axillary pedunculate heads 6–15 mm in diam., involucel

from near the base to c. half-way up the peduncle, very rarely somewhat higher
up. Calyx 1–2 mm long, subglabrous to pubescent. Corolla 2.5–3.5 mm long,
glabrous to pubescent outside. Pods especially variable, indehiscent, (4)8–17
(24) 1.3–2.2 cm, straight or curved, glabrous to grey-velvety, turgid. Seeds
deep-blackish-brown, 7–9 6–7 mm, smooth, subcircular, compressed, areole 6–
7 4.5–5 mm Acacia nilotica is native to the drylands of tropical Africa and western
Asia, eastwards as far as India, Myanmar, and Sri Lanka. In Africa, it occurs from
Senegal to Egypt and southwards through eastern Africa to Mozambique and
South Africa (Natal) and the Indian Ocean islands. It has been distributed throughout
the tropics and became naturalized in many areas, including Cape Verde, Jamaica,
Nepal, Indonesia, Vietnam, and Australia. It is widely cultivated in the Indian
subcontinent. (Brenan 1959).
Fig. 4 Acacia hookii

(Fabaceae), Sekenani, Maasai
Mara, Kenya. (Photo
R.W. Bussmann)
Acacia senegal (L.) Willd.: Shrub or tree up to 13 m tall, the bark grey to brown or
blackish, scaly, rough, young branchlets densely to sparsely pubescent, soon
glabrescent. Stipules not spinescent. Prickles just below the nodes, either in threes,
up to 7 mm long, the central one hooked downwards, the laterals curved upwards, or
else solitary, the laterals being absent. Leaves: petiole glandular or not (glands 0.5–
0.75 mm in diameter), rachis pubescent, glandular between the top 1–5 pairs of
pinnae, prickly or not, pinnae (2)3–6(12) pairs, 0.5–1.5(2.4, very rarely to 4 or more)
cm long, leaflets 7–25 pairs, 1–4(9) 0.5–1.75( 3) mm, linear-to elliptic-oblong,
ciliate on the margins only or hairy on the surface, or wholly subglabrous, lateral
nerves not visible or sometimes somewhat prominent beneath, apex obtuse to
subacute. Flowers white or cream, fragrant, sessile, in spikes 1.5–10 cm long on
peduncles 0.7–2 cm long, normally produced with the leaves, axis pubescent to
glabrous. Calyx 2–2.75(3.5) mm long, glabrous to somewhat pubescent. Corolla
2.75–4 mm long, exceeding the calyx, 5-lobed, glabrous outside. Stamen-filaments
4.5–7 mm long, free, anthers 0.2–0.25 mm, across, with a caducous gland. Ovary
glabrous, very shortly stipitate. Pods usually grey-brown, sometimes pale- or dark-
brown, dehiscent, (1.8)4–19 (1.2)2–3.4 cm, densely to sparsely appressed-
pubescent to puberulous, oblong, straight, venose, rounded to acuminate at the
apex. Seeds subcircular-lenticular, 8–12 mm in diam., central areole small to
medium, 2.5–6 2.5–5 mm, markedly impressed. Acacia senegal is widely
distributed in the drier parts of tropical Africa, from Senegal and Mauritania in the
west to Eritrea and Ethiopia in the north-east and to South Africa in the south. Of the
four recognized varieties var. senegal is the most widespread and is found through-
out the area of distribution (Brenan 1959).
The use of gum arabic (or gum acacia), which is derived from an exudate from the
bark, dates from the first Egyptian Dynasty (3400 B.C.). Gum Arabic has been used
for at least 4,000 years by local people for the preparation of food, in human and
veterinary medicine, in crafts, and as a cosmetic. Acacia senegal produces the only
acacia gum evaluated toxicologically as a safe food additive. Nowadays, the gum is
present in a wide range of everyday products. 60–75% of the world production of
gum Arabic is used in the food industry and in human and animal medicine (Brenan
1959) (Figs. 5, 6, and 7).
Acacia seyal Delille: Tree (2)3–9(12) m tall, bark on trunk powdery, white to
greenish-yellow or orange-red, young branchlets with few sparse hairs to almost
glabrous and with numerous reddish sessile glands (rarely, and not in our area, rather
densely puberulous), epidermis of twigs becoming reddish and conspicuously flak-
ing off to expose a greyish or reddish powdery bark. Stipules spinescent, up to 8 cm
long, “ant-galls” present or not, other prickles absent. Leaves often with a rather
large gland on the petiole and between the top 1–2 pairs of pinnae, pinnae (2)3–7
(8) pairs, leaflets (7)11–20 pairs, 3–8(10) 0.75–1.5(3) mm, in our area sparingly
ciliolate to glabrous, lateral nerves invisible beneath. Flowers bright yellow, in
Fig. 5 Acacia polyacantha

(Fabaceae), Sekenani, Maasai
Mara, Kenya. (Photo
R.W. Bussmann)
Fig. 6 Acacia senegal (Fabaceae), Sekenani, Maasai Mara, Kenya. (Photo R.W. Bussmann)
Fig. 7 Acacia senegal (Fabaceae), Amboseli, Kenya. (Photo R.W. Bussmann)
axillary pedunculate heads 10–13 mm in diameter borne on terminal or short lateral

shoots of the current season, involucel in the lower half of the peduncle, 2–4 mm
long, apex of bracteoles rounded to elliptic, sometimes pointed. Calyx 2–2.5 mm
Fig. 8 Acacia xanthophloea

(Fabaceae), Nairobi National
Park, Kenya. (Photo
R.W. Bussmann)
long, inconspicuously puberulous in upper part. Corolla 3.5–4 mm long, glabrous

outside. Pods dehiscent, (5)7–20(22) 0.5–0.9 cm, linear, falcate, constricted
between the seeds, finely longitudinally veined, glabrous except for some sessile
glands. Seeds olive to olive-brown, 7–9 4.5–5 mm, faintly and minutely wrinkled,
elliptic, compressed, areole 5–6 2.5–3.5 mm (Brenan 1959).
Acacia tortilis (Forssk.) Hayne: Tree 4–21 m. tall, occasionally a bush 1 m. high,
crown flat or spreading,10 bark grey to black, fissured, young branchlets glabrous to
densely pubescent, going brown to purplish-black. Stipules spinescent, some short
hooked and up to c. 5 mm long, mixed with other long straight whitish ones to c. 8
(10) cm long, “ant-galls” and other prickles absent. Leaves, rachis short, 2 cm long
or less, pinnae 2–10 pairs, 2–17 mm long, leaflets 6–19 pairs per pinna, usually very
small, 0.5–2.5(6) mm long, ciliate to glabrous. Flowers cream or whitish, in axillary
heads 5–10 mm in diameter on peduncles 0.4–2.4 cm long, involucel in the lower
half of the peduncle. Pods contorted or spirally twisted, longitudinally veined,
tomentellous to glabrous. Seeds olive to red-brown, 7 4.5–6 mm, smooth, elliptic,
compressed, areole 5–6 3–4 mm (Brenan 1959) (Figs. 8 and 9).
Acacia species are an important component of dry Afromontane forests (Bussmann
2002).
Acacia drepanolobium: Bark chewed for sore throat, root decoction given to
mothers after birth (Maasai), and for gastrointestinal problems (Beentje 1994;
Kokwaro 2009; Muthee et al. 2011).
Acacia nilotica: The species a wealth of medicinal uses. It is used for stomach upset
and pain, the bark is chewed to protect against scurvy, an infusion is taken for
dysentery and diarrhea. In Nigeria, it is one of the standard drugs for treating
diarrhea. It has also been used to eliminate stomach worms, as an antiseptic for
Fig. 9 Acacia xanthophloea

(Fabaceae), Nairobi National
Park, Kenya. (Photo
R.W. Bussmann)
open wounds and as an expectorant for treating coughs. In Africa, twigs are used as a
toothbrush. A decoction of the fruit is considered a febrifuge, and the seeds have
antimalarial, antidiabetic, antihypertensive, and antispasmodic activities. The bark,
gum, leaves, and pods are used in many traditional medicines, possessing anti-
inflammatory properties. The gum has been used as an emulsifying agent and
emollient. It is edible and is used to relieve throat and chest complaints. Bark
infusion used for digestion, the fruit juice for eye problems, as well a chest-problems
(Beentje 1994). The bark is chewed for sore throat and cough. Leaves are boiled and
the extract drunk as tea for chest-problems and pneumonia. Boiled bark and roots are
used for indigestion and stomach trouble, and sometimes used as aphrodisiac. The
bark decoction is uses for digestion and as stimulant, the root decoction for impo-
tence (Kokwaro 2009). In Ethiopia used externally for swellings and skin problems
(Wondimu et al. 2007). The Maasai use a bark decoction as stimulant (Muthee et al.
2011). Also used as digestive, to increase appetite, for chest pain, colds, backache,
stomach problems, and to treat diarrhea (Nankaya et al. 2019, 2020).
Acacia senegal / Acacia seyal: Bark, leaves, and gum are used as an astringent to
treat colds, ophthalmia, diarrhea, and hemorrhages. The seed contains a fat which is
used in medicine. A soothing and softening agent, gum Arabic is extensively
employed in folk medicines. Among many other uses, it is used internally for
coughs, diarrhea, dysentery, hemorrhages, and externally to cover inflamed areas.

Bark decoction used for malaria and diarrhea, root decoction also for diarrhea
(Beentje 1994). The root decoction is purgative and used for stomach-pain, and
diarrhea. The juice from the seed pods is used for eyesores. The bark decoction
serves for cough. Bark, gum and leaf juice are used for dysentery and stomach-ache.
Bark powder is applied to wounds. Leaves are chewed for toothache. The root
decoction also serves for diabetes (Kokwaro 2009). The bark is boiled for stomach
problems and abortion by the Samburu (Bussmann 2006), and a decoction is applied
externally for skin rashes (Njoroge et al. 2004).
Acacia tortilis: Bark decoction is used for diarrhea and stomach-ache (Beentje 1994;
Kokwaro 2009). Roots are used to treat skin diseases (Kokwaro 2009).
The bark of Acacia hookii is eaten by the Maasai for stomachache (Bussmann
et al. 2006), and similar uses are reported by the Samburu, who also use Acacia
etbaica for the same purpose. This species is also used to ease childbirth (Bussmann
2006). In Ethiopia the species serves for tonsilitis and gonorrhoea (Wondimu et al.
2007).
The bark decoction of Acacia concinna is used for toothache and body pain in
India, the fruit is used for diabetes (Debbarma et al. 2017). A bark paste of Acacia
leucophloea is applied to wounds (Muthu et al. 2006). Acacia polyacantha leaves
are boiled for abortion, while the bark of Acacia mearnsii is used for general
weakness (Njoroge and Bussmann 2006a). The bark of Acacia mellifera is chewed
to treat whooping cough (Njoroge and Bussmann 2006b), and for spleen pain
(Teklehaymanot et al. 2010). Acacia albida bark is used for diarrhea (Wondimu
et al. 2007). Acacia xanthophloea serves for skin problems and as stimulant among
the Maasai (Muthee et al. 2011).
Local Food Uses
Acacia drepanolobium: Young fruits and galls edible (Beentje 1994).

Acacia nilotica: The Hausa use roasted seeds as a food flavoring. In Tanzania, the
inner bark and the thick fruit pulp are boiled in water and drunk as a tea. The tannins
contribute to its many medicinal uses, the plant acting as a powerful astringent.
Acacia senegal / Acacia seyal: Seed may be dried and conserved for human
consumption mainly as an emergency food. In the food industry, gum Arabic is
used as a flavor fixative and emulsifier, to prevent crystallization of sugar in
confectionery, as a stabilizer in frozen dairy products, for its viscosity and adhesive
properties in bakery products, and as a foam stabilizer and clouding agent in beer.
The dried and preserved seeds of Acacia senegal are used as vegetables. The seeds
are sometimes eaten as a vegetable. In pharmaceuticals, it is used as a stabilizer for
emulsions, a binder and coating for tablets, and as an ingredient in cough drops and
syrups. The gum is edible (Beentje 1994).
Acacia tortilis: The gum is edible (Beentje 1994).
The fruit of Acacia concinna is eaten in India (Debbarma et al. 2017).
Acacia drepanolobium: Stems and branches used for fencing (Beentje 1994).
Acacia nilotica: Use for poles (Beentje 1994). The species is used in veterinary
medicine, for example, as a molluscicide to reduce liver-flukes in cattle. The pods are
desirable as fodder for cattle, and the leaves, young shoots, and young pods are
thought to aid milk production. Acacia nilotica wood burns without too much smoke
and provides good charcoal. The flowers provide pollen and nectar for bees. This
species is suitable for live fencing, mine timber, railway sleepers, boat building,
wheels, and water wells as its wood is durable and resistant to borers and termites.
The pods of Acacia nilotica are used traditionally in Nigeria and other sub-Saharan
countries for tanning leather and as a source of khaki-to-brown dyes if used without
mordant, or grey and black dyes for cotton combined with a mordant of iron-rich
mud. The latter process gives the typical dark grey stripes in the “hile,” “sampak,”
and “siole” traditional cotton textiles of Cameroon and in the “langtang” traditional
cotton weaving of the Mumuye of eastern Nigeria. These textiles were locally and
regionally used as currency well into the twentieth century and still have a great
cultural importance. Now they are mostly used as presents from a man to his future
wife and in major religious festivals, clan gatherings, and funerals. Black dyes
combining tannins most probably obtained from “sant” (acacia pods) with iron
mordants have been identified in ancient Egyptian textiles dating from the 18th
Dynasty (1542–1305 BC) onwards. The dried mature pods used in local tanneries in
Sudan produce a pinkish white leather of good quality. The tan stuff extracted from
the pods is internationally called by its Hausa name: “bagaruwa’” In contrast in India
and Pakistan, it is the bark, a by-product from timber plantations, that is used for
tanning and dyeing leather. The tannin produces a heavy leather which is firm,
durable, and hard, but combined with myrobalans (from Terminalia species) it
produces excellent leather. Acacia nilotica is widely used as a timber, source of
fodder, tannin and gum, and as a fence, shade, and fuel tree. The dark brown
heartwood is nearly twice as hard as teak and very shock resistant, and is used
widely in constructions, for railway sleepers, mine props, tool handles, and carts.
The trees make effective live fencing, are a good host for the semiparasitic sandal-
wood (Santalum album L.) and are important trees for lac insects (shellac) in the
Indian subcontinent. Gum collected from the trunk and branches was formerly used
in paints and medicines. It has properties similar to true gum arabic from Acacia
senegal and is frequently used in calico printing and dyeing as a thickening agent. It
is also used as sizing material for silk and cotton, and in paper manufacture in India.
The leaves and pods are an excellent fodder, rich in protein. The flowers yield a
honey of good quality. The bark of slender branches yields a fiber which is used for
the manufacture of paper or is made into coarse ropes in India. Acacia nilotica is also
a popular ornamental avenue tree. The pods have molluscicidal and algicidal prop-
erties and are added to ponds in Sudan to kill snail species that carry schistosomiasis
without affecting the fish. Bark decoction as panacea for livestock (Kokwaro 2009).
Acacia senegal / Acacia seyal: The foliage and pods are an important fodder source for
camels and goats. The wood is used for small-scale construction purposes and
agricultural implements, it yields a fuelwood of good quality that can be made into good
charcoal. The thorny branches are often used to make “dead fences” to enclose livestock
or protect agricultural fields. Being a very drought-resistant tree, it is planted for sand
dune fixation, windbreaks, and shelter belts in arid regions. The flowers are a source of
honey. Cordage is made from the roots, either directly or after beating to extract the
fibers, its strength makes it suitable for well ropes and fishing nets. The seed contains a
fat which is used for soap making. The leaves and pods are browsed by sheep, goats,
camels, impala, and giraffe. Gum Arabic is used in cosmetics as an adhesive for face
masks and powders, and to give a smooth feel to lotions. Acacia senegal wood is locally
valued for fuel wood and charcoal. Industrially, gum Arabic is applied as an adhesive, as
a protective colloid and safeguarding agent for inks, sensitizer for lithographic plates,
coating for special papers, sizing agent for cloth to give body to certain fabrics, and
coating to prevent metal corrosion. Gum Arabic is also used in the manufacture of
matches and ceramic pottery. In construction, the wood is used locally for poles and
fenceposts, the light-colored wood for tool handles and dark heartwood for weaver’s
shuttles. Strong ropes are made from the bark fibers of the long surface roots. Where the
trees are large (for example near the River Niger) they are cut into planks at least 12 cm
thick for making canoes for hunting hippopotamuses. The wood is hard and heavy and
takes a beautiful polish, with the sapwood being yellowish white and the heartwood
nearly black and irregular. The wood is made into throwing-sticks which, in contrast to
the Australian boomerang, can be made to fly straight and used for hunting and
pageantry. The flowers are a good source of honey, an important source of nutrition
and income generation to support rural livelihoods in marginal lands. The bark yields a
red dye (Beentje 1994). The leaves are eaten by livestock, and the species serves for
construction and fencing (Bussmann 2006). Used to treat livestock (Lulekal et al. 2008).
Acacia tortilis: Bark used for fibers and roots used for basketry. Fruits eaten by
livestock (Beentje 1994). The thorns are used for witchcraft (Kokwaro 2009). Often
used as firewood and for fencing (Lulekal et al. 2008).
All species are used for fencing and firewood (Kiefer and Bussmann 2008;
Bussmann et al. 2018). The Maasai use Acacia hookii and Acacia polyacantha
for fencing because they have long thorns. The same holds true for Acacia etbaica.
Both are browsed by elephants and goats and give good honey (Bussmann et al.
2006; Bussmann 2006). Acacia abyssinica serves for charcoal, firewood, and is
eaten by cattle and camels (Bussmann et al. 2011), and together with Acacia albida,
Acacia mearnsii, and Acacia saligna as shade plant, and for making tools (Mekonen
et al. 2015). Acacia pennata leaves are used as fish poison (Neuwinger 2004).
Acacia brevispica is used to remove nasal bots, and other parasites in cattle (Njoroge
et al. 2006; Wanzala 2017).
References
Brenan JPM. Flora of tropical East Africa: Leguminosae, subfamily Mimosoideae. London: Crown
Agents for Overseas Governments & Administrations; 1959.
appendices 1–7, www.naturekenya.org/JournalEANH.htm
Sekenani Valley, Maasai Mara, Kenya. Econ Bot. 2018;722:207–16.
tribes of Tripura, Northeast India. Afr J Tradit Complement Altern Med. 2017;144:156–68.
Lulekal E, Kelbessa E, Bekele T, Yineger H. An ethnobotanical study of medicinal plants in Mana
Angetu District, southeastern Ethiopia. J Ethnobiol Ethnomed. 2008;4:10. https://doi.org/
10.1186/1746-4269-4-10.
District of the Oromia region of Ethiopia to assess use, species diversity and management
Muthee JK, Gakuya DW, Mbaria JM, Kareru PG, Mulei CM, Njonge FK. Ethnobotanical study of
anthelmintic and other medicinal plants traditionally used in Loitoktok district of Kenya.
J Ethnopharmacol. 2011;135:15–21.
Muthu C, Ayyanar M, Raja N, Ignacimuthu S. Medicinal plants used by traditional healers in
Kancheepuram District of Tamil Nadu, India. J Ethnobiol Ethnomed. 2006;2:43.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Sustainability of the Loita Maasai childrens’
ethnomedicinal knowledge. Sustainability. 2019;11:5530. https://doi.org/10.3390/su11195530.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Medicinal plants of the Maasai of Kenya: a review.
Plants. 2020;9:44. https://doi.org/10.3390/plants9010044.
Neuwinger HD. Plants used for poison fishing in tropical Africa. Toxicon. 2004;44:417–30.
Njoroge GN, Bussmann RW. Traditional management of ear, nose and throat ENT diseases in
Central Kenya. J Ethnobiol Ethnomed. 2006a;2:54.
Njoroge GN, Bussmann RW. Herbal usage and informant consensus in ethnoveterinary management
of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006b;108:332–9.
source of traditional medicines in Central Kenya: optimizing resource efficiency R.U.E. in agro-
ecosystems. Lyonia. 2004;72:71–87.
Njoroge GN et al. Herbal usage and informant consensus in ethnoveterinary management of cattle
diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006;108:332–9
Teklehaymanot T, Giday M, Mutse M. Quantitative ethnobotany of medicinal plants used by Kara
and Kwego semi-pastoralist people in lower Omo River valley, Debub Omo zone, southern
nations, nationalities and peoples regional state, Ethiopia. J Ethnopharmacol. 2010;130:76–84.
Wanzala W. Potential of traditional knowledge of plants in the management of Arthropods in
livestock industry with focus on Acari ticks. Evid Based Complement Alternat Med. 2017;
https://doi.org/10.1155/2017/8647919. 8647919, 33 p.
Wondimu T, Asfaw Z, Kelbessa E. Ethnobotanical study of medicinal plants around ‘Dheeraa’
town, Arsi zone, Ethiopia. J Ethnopharmacol. 2007;112:152–61.
Acanthus eminens C.B. Clarke
Acanthus sennii Chiov.
ACANTHACEAE

Grace N. Njoroge
Local Names
Acanthus eminens: Kikuyu: Kitamani, Maa: Ol-digeledi, Marakwet: Tegilde,

Nandi/Kipsigit: Tegeldet, Okiek: Tikeltit, Sebei: Tekandet, Luhya: LIrakalu
(Beentje 1994; Kokwaro 2009).
Botany and Ecology
Acanthus eminens C.B. Clarke: Shrubby herb or shrub to 3 m tall, young stems
glabrous. Leaves with petiole 0.5–2(2.5) cm long, lamina narrowly elliptic to elliptic
or obovate in outline, largest 13–37 5–15 cm, deeply lobed with large triangular
spine-tipped lobes, each lobe with a spine on antrorse side, apex acuminate to
cuspidate, spine-tipped, base attenuate to cuneate, glabrous, each node with one to
several persistent sharp terete straw-coloured interpetiolar downwardly directed
pungent pale yellow spines to 1.5 cm long, occasionally some with a small lamina.
R. W. Bussmann (*)
G. N. Njoroge

Cymes solitary or also 2 from upper leaf-pair, (5–)7–30 cm long, rachis upwards
pubescent with hairs and few to many short to long capitate glands, with several
pairs of sterile bracts at base, sometimes merging into the vegetative leaves, bracts
green to purplish, ovate to elliptic, finely puberulous to sparsely pubescent and
upwards with few to many stalked capitate glands, 1.4–2.8 cm long in middle of
cyme, spine-tipped and with 4–8 teeth per side (rarely entire), bracteoles subulate to
linear, 5–11 mm long, entire. Calyx green to purplish, puberulous and with few to
many stalked capitate glands, dorsal and ventral sepals longer than bract, oblong-
obovate, similar or dorsal slightly longer, 2.1–4 cm long, dorsal entire or 2–3-
toothed, ventral 2–3-toothed or irregularly toothed, lateral lanceolate, 1.3–2.5 cm
long, acuminate. Corolla dark blue to dark purple (rarely pale blue to almost white),
tube 4–5 mm long below thickened rim which is 3–4 mm long, limb 5-lobed, 3.5–
5 3.5–5 cm, below puberulous with hairs and capitate glands, above glabrous,
callus with narrow central groove. Filaments 1.7–2.2 cm long, glabrous or with
stalked glands near apex, anthers 6–9 mm long, densely glandular dorsally. Capsule
2.3–2.5 cm long. Seed ellipsoid to round, 9–10 mm long, glabrous. In primary and
disturbed wet evergeen montane forest, forest margins, secondary grassland, often
forming large stands, occasionally used as a hedge plant, (1200–)1500–2600 m
(Vollesen 2008) (Fig. 1).
Fig. 1 Acanthus eminens

(Acanthaceae), Mt. Kenya
Afromontane forest, Kenya.
Acanthus eminens C.B. Clarke . . . 97
Fig. 2 Acanthus sennii

(Acanthaceae), Bale mountain
National Park, Odo Bulu
forest, Ethiopia. (Photo
R.W. Bussmann)
Acanthus sennii Chiov: Shrub, arising from a subterranean rhizome, with erect
stems 0.4–2.5 m tall. Stems simple or branched, glabrous to densely pubescent,
internodes 416 cm long. Leaves pinnatifid to strongly toothed, 10–23 6–16 cm,
oblong-ovate, margin and apex sharply spiny, surfaces glabrous or pubescent, bright
mid-green above. Inflorescence terminal, 5–13 cm long, with numerous densely
packed flowers subtended by spiny bracts. Calyx with unequal outer (to 33 10 mm)
and inner (to 17 5 mm) lobes, spiny at their apices. Corolla red, 60–63 mm,
formed of a tube of 10–11 mm and a large wide-spreading lip 50–55 mm long;
stamens with filaments 27–29 mm, bearing single-celled, white-hairy anthers to
5 mm Fruit a glabrous capsule (Kelbessa 2006) (Figs. 2, 3, 4, and 5).
Acanthus eminens: Leaf decoction used to treat liver and spleen problams
(Kokwaro 2009). The leaves are applied for skin infections, wounds and eye
problems (Jeruto et al. 2008).
Acanthus sennii: In Ethiopìa the leaves are crushed and put on wounds (Bussmann
et al. 2011; Luizza et al. 2013).

R.W. Bussmann)

R.W. Bussmann)
Acanthus eminens C.B. Clarke . . . 99

R.W. Bussmann)
Acanthus eminens and Acanthus sennii are both good honey plants (Bussmann
et al. 2011).
References
Jeruto P, Lukhoba C, Ouma G, Otieno D, Mutai C. An ethnobotanical study of medicinal plants
used by the Nandi people in Kenya. J Ethnopharmacol. 2008;116:370–6.
Kelbessa E. Acanthaceae. In: Hedberg I, Edwards S, Kelbessa E, Demissew S, Persson E, editors.
Flora of Ethiopia and Eritrea, vol. 5. Addis Ababa/Uppsala: The National Herbarium/Depart-
ment of Systematic Botany; 2006. p. 345–495.
Res Appl. 2013;11:315–39.
Vollesen K. Flora of tropical East Africa: Acanthaceae (Part 1). Kew: Royal Botanic Gardens; 2008.
286 pp. ISBN: 978-1-84246-217-1
Achyranthes aspera L.
AMARANTHACEAE

Grace N. Njoroge
Synonyms
Achyranthes aspera L.: Achyranthes argentea Lam.; Achyranthes aspera var.

indica L.; Achyranthes aspera var. obtusifolia (Lam.) Sues.; Achyranthes indica
(L.) Mill.; Achyranthes obtusifolia Lam.; Achyranthes robucta C.H. Wright;
Achyranthes sicula Roth; Cadelaria indica (L.) Raf.; Cadelaria sicula Raf.;
Centrostachys aspera (L.) Standl.; Centrostachys indica (L.) Standl.; Cyathula
geniculata Lour.; Stachyarpagophora aspersa (L.) M. Gómez
Local Names
Achyranthes aspera: Luo: Auch-Auch; Turu: Inyalo; Hehe: Lugeni; Luhya:

Lusayi; Swahili: Pulule; Kamba: Uthekethe (Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Achyranthes aspera L.: Perennial herb (sometimes woody and somewhat

suffrutescent), occasionally flowering in the first year, 0.2–2 m, stiffly erect to
subscandent or straggling and prostrate, simple to much branched, stems stout
to very weak, distinctly to obscurely 4-angled, striate or sulcate, subglabrous to
densely tomentose, the nodes shrunken when dry. Bracteoles 1.5–4.5(6) mm, the
basal wings one-third to one-quarter the length of the spine and more or less adnate
to it (sometimes free above or tearing free), typically tapering off above but not
rarely rounded or truncate. Perianth whitish or pale green to red or purple, segments
5, 3–7(10) mm long, the outer pair longest, narrowly lanceolate to lanceolate, very
acute, with a distinct midrib and 2 obscure to distinct lateral nerves, narrowly or
moderately pale-margined. Inflorescences at first dense, finally elongating to (5)8–
34(40) cm, peduncles (0.6–1)1–6(7.5) cm long. Leaves elliptic, oblong or ovate,
acute or acuminate to almost round and very obtuse, gradually or abruptly narrowed
below, (2–)5–22(–28) 1.3–8(–10) cm, indumentum varying from subglabrous
through subglabrous above and densely appressed-canescent below to densely
tomentose on both surfaces, petioles of main stem leaves 3–25(–30) mm, shortening
above and below. Bracteoles 1.5–4.5(–6) mm, the basal wings 1/3–1/4 the length of
the spine and adnate to it (sometimes free above or tearing free), typically tapering
off above but not rarely rounded or truncate. Perianth whitish or pale green to red or
purple, segments 5, 3–7(–10) mm, the outer pair longest, narrowly lanceolate to
lanceolate, very acute, with a distinct midrib and 2 obscure to distinct lateral nerves,
narrowly or moderately pale-margined. Inflorescences at first dense, finally elongat-
ing to (5–)8–34(–40) cm, peduncles (0.6–)1–6(–7.5) cm glabrous. Seed filling the
capsule, oblong-ovoid, smooth. Stamens 5, the filaments 1.5–4.5(–6) mm, alternat-
ing with subquadrate pseudostaminodes, typically the apex of the latter curves
slightly inwards as a narrow, crenate or entire, often very delicate flap, while from
the dorsal surface arises afimbriate-ciliate scale extending across the width of the
pseudostaminode, not rarely; however, this is reduced to a “stag’s-horn” process at
the centre of the dorsal surface or a shallow, dentate rim, or even becomes small and
filiform – or else subapical or apical so that the pseudostaminode appears simple
(this usually in small forms of var. sicula). Style slender, 1–4(–6) mm (Townsend
1985). Very common in the ground layer of drier Afromontane forests (Bytebier and
Bussmann 2000; Bussmann 2002a, b, 2006) (Figs. 1 and 2).
Achyranthes aspera: Powder of fire dried leaves is applied in cuts to stop bleeding
and in ankles to cure sprains. Also used for headache. The roots are chewed and
applied to cuts to stop bleeding. A root decoction serves to treat constipation,
especially in children, and to cure stitch. Pounded roots extracted in water serve
for venereal diseases. Burned leaves are applied to boils, and leaf paste is applied to
wounds to stop bleeding. The leaf juice is drunk for gonorrhea, and the root
Achyranthes aspera L. 103
Fig. 1 Achyranthes aspera

(Amaranthaceae), Pusac,
Amazonas, Peru. (Photo
R.W. Bussmann and Narel
Y. Paniagua-Zambrana)
Fig. 2 Achyranthes aspera

(Amaranthaceae), flowers,
Pusac, Amazonas, Peru.
(Photo R.W. Bussmann and
Narel Y. Paniagua-Zambrana)
decoction is used to stop miscarriages (Kokwaro 2009). Roots are boiled and the
decoction is drunk to make patients vomit if they have malaria (Bussmann et al.
2006; Bussmann 2006). Also used by the Kikuyu for backpain, toothache, and to
treat anthrax (Njoroge et al. 2004). In Ethiopia, very commonly used for amoebic
diarrhea and externally for wounds and bleeding (Giday et al. 2003, 2007, 2010;
Mekonen et al. 2015; Teklehaymanot et al. 2007; Teklehaymanot and Giday 2010).
The same use is observed in India (Muthu et al. 2006). It can also be used for cough
(Jeruto et al. 2008).
In India, the root is tied to people suffering from epilepsy and shivering
(Debbarma et al. 2017), as well as in decoction for indigestion, constipation, and
goiter (Reang et al. 2016). It is also used as contraceptive in women, asthma,
bleeding in delivery, boil, bronchitis, child birth, cold, cough, disability, dropsy,
dog bite, dysentery, ear complaints, headache, rabies, insect bite, leukoderma, liver
complaints, pain in ribs, body, piles, pneumonia, renal complaints, phlegm, scabies,
scorpion bite, skin diseases, snakebite, sore, stomachache, syphilis, toothache,
whooping cough, wounds, and eye complaints (Verma et al. 2007). In Pakistan,
the plant is used to heal wounds (Ur-Rahman et al. 2018), as well as to treat kidney
stones, ulcers, pneumonia, chest pain, dysmenorrhea and aerodontalgia (Umair et al.
2019). It is used to ease labor, to relieve body pain and fever, and to treat constipation
(Kunwar and Bussmann 2009; Malik et al. 2015; Sher et al. 2016). In Nepal, the
whole plant is used in leprosy, bronchitis, expectorant, asthma, cough, skin diseases,
sinusitis, headache, colic, boils, and piles. Plant has emetic properties. In Chinese
traditional medicine, hot water extract of the plant is used to alleviate arthritic pain
(Kunwar et al. 2009). In the Garwhal Himalaya, the root infusion is used for malarial
fever; the leaf extract facilitate delivery; and the plant decoction is used to treat
dropsy and bronchitis (Kumar et al. 2011).
Achyranthes aspera: Eaten by cattle (Bussmann et al. 2011; Bussmann 2006). The
root is crushed, boiled, and the extract used against rabies and anthrax (Bussmann
et al. 2011; Wondimu et al. 2007; Yineger et al. 2007).
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002a;91(1/2):27–79,
appendices 1–7. www.naturekenya.org/JournalEANH.htm
Syst Geogr Plants. 2002b;71(2):959–74.
syntaxonomical survey. J East Afr Nat Hist. 2000;89(1–2):45–71.
tribes of Tripura, Northeast India. Afr J Tradit Complement Altern Med. 2017;14(4):156–68.
Achyranthes aspera L. 105
Giday M, Asfaw Z, Elmqvist T, Woldu Z. An ethnobotanical study of medicinal plants used by the
Zay people in Ethiopia. J Ethnopharmacol. 2003;85:43–52.
Giday M, Teklehaymanot T, Animut A, Mekonnen Y. Medicinal plants of the Shinasha, Agew-awi
and Amhara peoples in northwest Ethiopia. J Ethnopharmacol. 2007;110:516–25.
Giday M, Asfaw Z, Woldu Z. Ethnomedicinal study of plants used by Sheko ethnic group of
Ethiopia. J Ethnopharmacol. 2010;132(1):75–85. https://doi.org/10.1016/j.jep.2010.07.046.
Kumar M, Bussmann RW, Mukesh J, Kumar P. Ethnomedicinal uses of plants close to rural
habitation in Garhwal Himalayan, India. J Med Plant Res. 2011;5(11):2252–60.
Kunwar RM, Bussmann RW. Medicinal, aromatic and dye plants of Baitadi and Darchula districts,
Nepal Himalaya: status, uses and management. In: Hartmann M, Weipert J, editors.
Biodiversität, Naturausstattung im Himalaya, vol. III. Erfurt: Verein der Freunde & Förderer
des Naturkundemuseums; 2009. p. 475–89.
Kunwar RM, Upreti Y, Burlakoti C, Chowdhary CL, Bussmann RW. Indigenous use and ethnophar-
macology of medicinal plants in Far-West Nepal. Ethnobot Res Appl. 2009;7:5–28.
Malik ZA, Bhat J, Ballabha R, Bussmann RW, Bhat AB. Ethnomedicinal plants traditionally used in
health care practices by inhabitants of Western Himalaya. J Ethnopharmacol. 2015;172:133–44.
ecosystems. Lyonia. 2004;7(2):71–87.
Reang I, Goswami S, Pala NA, Kumar K, Bussmann RW. Ethnoveterinary applications of medic-
inal plants by traditional herbal healers in Reang tribe of South district Tripura, India. Med
Aromat Plants. 2016;5(2):234.
Sher H, Bussmann RW, Hart R, de Boer HJ. Traditional use of medicinal plants among Kalasha,
Ismaeli and Sunni groups in Chitral District, Khyber Pakhtunkhwa province, Pakistan.
J Ethnopharmacol. 2016;188:57–69. https://doi.org/10.1016/j.jep.2016.04.059.
Teklehaymanot T, Giday M. Quantitative ethnobotany of medicinal plants used by Kara and Kwego
semi-pastoralist people in lower Omo River Valley, Debub Omo Zone, Southern Nations,
Nationalities and Peoples Regional State, Ethiopia. J Ethnopharmacol. 2010;130:76–84.
Teklehaymanot T, Giday M, Medhin G, Mekonnen Y. Knowledge and use of medicinal plants by
peoplearound Debre Libanos monastery in Ethiopia. J Ethnopharmacol. 2007;111:271–83.
Townsend CC. Amaranthaceae. Flora of tropical East Africa. Rotterdam/Boston: A.A. Balkema;
1985.
Umair M, Altaf M, Bussmann RW, Abbasi AM. Ethnomedicinal uses of the local flora in Chenab
riverine area, Punjab province Pakistan. J Ethnobiol Ethnomed. 2019;15:7. https://doi.org/
10.1186/s13002-019-0285-4.
Ur-Rahman I, Afsal A, Iqbal Z, Ijas F, Ali N, Asif M, Alam J, Majid A, Bussmann RW. Traditional
and ethnomedicinal dermatology practices in Pakistan. Clin Dermatol. 2018;36(3):310–9.
https://doi.org/10.1016/j.clindermatol.2018.03.018.
Verma A, Kumar M, Bussmann RW. Medicinal plants in an urban environment: The medicinal flora
town, Arsi Zone, Ethiopia. J Ethnopharmacol. 2007;112:152–61.
Yineger H, Kelbessa E, Bekele T, Lulekal E. Ethnoveterinary medicinal plants at Bale Mountains
National Park, Ethiopia. J Ethnopharmacol. 2007;112:55–70.
Acokanthera schimperi (A. DC.) Benth. &
Hook. f. ex Schweinf.
EUPHOPRBIACEAE

Grace N. Njoroge
Local Names
Acokanthera schimperi: Kikuyu: Murichu; Kipsigis: Kelyot; Fipa: Namikouko;

Maa: Ol-Moringye, Ol-Morijoi; Swahili: Muchungu; Borana: Kararu; Gabbra:
K’arraru; Giriama: Chungua; Kamba: Kivai; Muvai; Meru: Mururu; Kipsigis/
Nandi/Lumbwa: Kalyo, Keliot; Pokot: Kalyan; Samburu: Lmorijoi; Somali:
Marid; Tugen: Gelwon, Kelwon (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Acokanthera schimperi (A. DC.) Benth. & Hook. f. ex Schweinf.: Shrub or tree
1–10 m high, branching from the base; bark rough grey-brown; slash cream with fine
orange streaks; young branches glabrous or pubescent, conspicuously angled and
ribbed. Leaves petiolate, petiole 3–6 mm long; blade ovate, obovate, elliptic to
suborbicular, 1–8( 10) cm long, 1.1–6.5 cm wide, rounded or acute at the apex,
sometimes emarginate, cuneate or rounded at the base, coriaceous, glabrous;
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Acokanthera
cf. schimperi
(Euphorbiaceae), Maasai
Mara, Kenya. (Photo
R.W. Bussmann)
secondary veins obscure, especially beneath. Fruit purple when ripe, ellipsoid,
10–25 mm long; seeds up to 13 mm long. Corolla tube pink or reddish, 8–11 mm
long, glabrous or pubescent on external surface, inner surface pilose in the upper half
and wrinkled below; lobes pink outside, white inside, ovate, 2.5–3.5 mm long, 1.5–
2.5 mm wide, acuminate at the apex, pubescent at the base inside, glabrous or
pubescent outside, ciliate; stamens slightly exserted, inserted at 7–10 mm from the
base; anthers ovate, 1–1.5 mm long, acumen 0.2 mm long; pistil 7.5–10 mm long.
Inflorescence contracted, of many-flowered axillary cymes; sepals ovate, 1–2 mm
long, 0.5–1 mm wide, pubescent or glabrous, ciliate. Dry forest and forest margins,
wooded grassland, rocky bushland; 250–2200 m (Bussmann 2002; Omino 2002)
(Fig. 1).
Acokanthera schimperi: The root infusion is used to treat syphilis (Kokwaro 2009).
Acokanthera schimperi: The root, twigs, leaves, and bark infusion are used as arrow
poison (Beentje 1994; Jones 2007; Kokwaro 2009).
References
Bussmann RW. Islands in the desert - forest vegetation of Kenya’s smaller mountains and highland
appendices 1–7 @ www.naturekenya.org/JournalEANH.htm
Acokanthera schimperi (A. DC.) Benth. & Hook. f. ex Schweinf. 109
Gachati FN. Kikuyu botanical dictionary. Nairobi: AMREF; 1989.

Jones DE. Poison arrows. Austin: University of Texas Press; 2007.
Omino EA. Flora of tropical East Africa: apocynaceae (part 1). Rotterdam: Balkema; 2002. 114 pp.
ISBN 90-5809-409-X.
Adansonia digitata L.
MALVACEAE

Grace N. Njoroge
Synonyms
Adansonia bahobab L.; Adansonia baobab Gaertn.; Adansonia integrifolia Raf.;

Adansonia klima Pettigrew, K.L. Bell.; Bhagw.; Grinan, Jillani; Jean May., Wab.;
Adansonia scutula Steud.; Adansonia situla (Lour.) Spreng.; Adansonia
sphaerocarpa A. Chev.; Adansonia sulcata A. Chev.; Baobabus digitata (L.)
Kuntze; Ophelus sitularious Lour.
Local Names
Adansonia digitata: Swahili: Mbuyu, Mkuu hapingwa, Mkuu hafungwa, Muuyu;

Boni: Jah; Kamba: Kiamba, Muamba; Maa: Ol-Imisera; Oromo/Somali: Yak;
Pare: Maramba; Digo/Giriama/Swahili: Mbuyu; Taita: Mlamba; Sukuma:
Moandu; English: Babobab, Monkey-bread tree, Dead-rat tree, Cream-of-tartar
tree (Beentje 1994; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Adansonia digitata: A massive, deciduous tree with a trunk that can grow to an
immense girth. The bark is smooth. Borne at the ends of branches, leaves are usually
divided into 5–7 leaflets attached to a central point. Large (up to 20 cm in diameter),
white and pendent on long stalks. Each flower has five free petals and many stamens
(male parts). More or less cylindrical and up to 35 cm long and 13 cm wide. Each
fruit is filled with mealy pulp containing many small, dark brown seeds, each about
1 cm long and wide. The seeds have a reddish-black seed coat. Baobab occurs
naturally in most of mainland tropical Africa. Originally, it was absent from many
Central African countries, but it has been introduced in most of them. In mainland
tropical Africa, it is still absent from Rwanda, Burundi, Djibouti, and Uganda. It has
been introduced in Madagascar and many other Indian Ocean islands. In West
Africa, it often occurs in baobab orchards around villages. Outside Africa it has
been widely introduced in tropical and subtropical regions. In India it is a fairly old
introduction most likely brought by Arab traders and it now has many local uses
similar to those in Africa. In Barbados, two old trees, grown from seed imported in
1738 from West Africa, are considered one of the seven wonders of the island
(Beentje 1989) (Figs. 1, 2, 3, 4, and 5).
Fig. 1 Adansonia digitata (Malvaceae), Meru, Kenya. (Photo R.W. Bussmann)

Adansonia digitata L. 113
Fig. 2 Adansonia digitata

(Malvaceae), Tsavo, Kenya.
Adansonia digitata L.: Roots, bark, leaves, fruits and seeds are used medicinally for
an enormous range of ailments, among the more common of which are iron
deficiency, digestive system disorders, infections, and skin disorders. All plant
parts are used for treating fever. The bark gum is used for cleaning sores. A decoction
of the bark is used in Congo to bathe rickety children and in Tanzania to treat
toothache. Stem bark and fibers lining the fruit husk are used to treat amenorrhea.
The bark, fruit pulp, and seeds are used as an antidote for Strophanthus poisoning. In
Malawi baobab juice called “dambedza” is served as a cure for hangovers and
against constipation. In Zambia, a root infusion is used to bathe babies to promote
a smooth skin. A root decoction is taken with food in Sierra Leone for strength.
Fig. 3 Adansonia digitata (Malvaceae), Meru, Kenya. (Photo R.W. Bussmann)
Fig. 4 Adansonia za (Malvaceae), Sakhara, Madagascar. (Photo R.W. Bussmann)

Fig. 5 Adansonia za (Malvaceae), Sakhara, Madagascar. (Photo R.W. Bussmann &

Dried powdered roots are included in the treatment of malaria. Leaves are used as a
diaphoretic, expectorant, astringent, and as a fever prophylactic. Leaves are found in
a long list of treatments for ailments including asthma, fatigue, kidney, and bladder
diseases, diarrhea, and inflammations. Fruit pulp is used as an antidysenteric and in
the treatment of smallpox and measles. The seed oil is used for inflamed gums and to
ease toothache. A root decoction is used to treat laziness, the bark decoction for body
pains, menstrual problems, diabetes, anemia, diarrhea, and to bath weak infants.
Steam baths with bark are to treat high fever, and as diaphoretic. The fruit pulp serves
to treat diarrhea and fever, the leaf decoction serves for fever, filariasis, and as
astringent (Kokwaro 2009).
Local Food Uses
Adansonia digitata: An important fruit tree, the fruit pulp (rich in vitamin C) is eaten
on its own or mixed in porridge and is also used for making soft drinks. Seeds are
used as a thickener for soups, and leaves are eaten as a vegetable or in soups. The
leaves of baobab are used either fresh as a cooked vegetable or dried and powdered
as an ingredient of soups and sauces (Beentje 1994). The shoots and roots of
seedlings are eaten as well. The roots are boiled and eaten in West Africa in times
of famine. The flowers are eaten raw. The fruit contains soft, white, edible, and
nutritious flesh (“monkey bread”). It is used to curdle milk, and it is eaten as a sweet
and is used in making gruel and refreshing drinks and ice-cream. In Sudan, it is made
into a milk-like drink called “gubdi.” The powdered fruit flesh is added to cold
liquid, thus preserving vitamins. An emulsion of the fruit pulp may be used to
adulterate milk. The dried pulp is used as a substitute for cream of tartar in baking.
The seeds are eaten raw or roasted and are used to thicken and flavor soup.
Fermentation of the seed kernels improves the nutritional value. In coastal Kenya
and Tanzania, the pulp-coated seeds are colored and sugar-coated and sold as sweets.
The seeds are used to adulterate groundnuts and may be used as a coffee substitute.
An oil can be extracted from the seed kernels by boiling and distillation. It is
semifluid, golden yellow, gently scented, nondrying, and has a long shelf life, and
it is used for cooking.
Adansonia digitata: The bark fiber is stripped from the lower part of the trunk and is
used to make rope, string, cords for musical instruments, snares, fishing-nets, loin
cloths, sacking, baskets, mats, and waterproof hats. The root bark yields a fiber as
well and is used for dye (Beentje 1994). When the wood disintegrates fibers remain
that may be used as packing material. The leaves and fallen flowers are eaten by
livestock, and fruit pulp and seeds are fed during the dry season. Residues of oil
extraction are fed to livestock as well. The wood is used locally for canoes, wooden
platters and floats for fishing nets. It makes poor firewood and charcoal. Use of the
wood and bark for the paper industry is possible but not commercially viable. The
fruit husks are used as fuel and are made into containers and fishing floats. Trees with
hollow trunks have been used for a long time and are still used for water storage,
having capacities of up to 7500 l. Hollow trees are also used as tombs, meeting
places, prisons, stables, bus shelters, storage rooms, watchtowers, bathrooms, cool
rooms, and dairies. Pulp is used for smoking fish and the smoke is helpful in
repelling insects that pester livestock. The pollen mixed with water gives a glue
that is used in carpentry. The bark is used for tanning and green bark yields a dye. A
red dye is obtained from the roots. The powdered peduncle and the husk of the fruit
are used as a substitute for tobacco. Introductions outside its native range were made
mainly for use as an ornamental or shade tree. The extraordinarily shaped baobab
tree is surrounded by a wealth of legends, superstitions, folktales, and anecdotal
references throughout Africa. Fibers from the inner bark are used to make rope and
string for basketry, as well as for making beehives. Trunks that have been hollowed
by lightning or by humans have been employed imaginatively as a pub, toilet, prison,
and bus stop. In western Sudan, the trunks were used as water containers. The roots
produce a dye. Baobab also has some perceived magical uses. For example, it is said
that a decoction of the seeds will protect you against crocodiles and that flowers are
inhabited by spirits.
The bark is boiled as cure for Brucellosis in cattle (Kioko et al. 2015).
References
Beentje H. Flora of tropical East Africa: Bombacaceae. Rotterdam: Balkema; 1989. 9 pp. ISBN
90-6191-342-X.
Kioko J, Baker J, Shannon A, Kiffner C. Ethnoecological knowledge of ticks and treatment
of tick-borne diseases among Maasai people in Northern Tanzania. Veterinary World.
2015;8(6):755–762.
Aerva lanata (L.) Juss. ex Schult.
AMARANTHACEAE

Grace N. Njoroge
Synonyms
Aerva lanata (L.) Juss. ex Schult.: Achyranthes lanata L.; Achyranthes villosa
Forssk.; Aerva arachnoidea Gand.; Aerva elegans Moq.; Aerva floribunda Wight;
Aerva mozambicensis Gand.; Aerva pubescens Mart.; Aerva sansibarica Suess.;
Aerva tandalo Buch.-Ham. ex Dill.; Alternanthera pubescens Moq.; Amaranthus
aeruoides Hochst. & Steud. ex A. Rich.; Amaranthus lanatus Dum. Cours.; Celosia
lanata L.; Illecebrum lanatum (L.) L.; Illecebrum pubescens Willd.; Ouret lanata
Hiern; Uretia lanata Kuntze.
Local Names
Aerva lanata: Kipsigis: Baiy, Digo: Chivwa Kuku, Sukuma: Luwecha, Nyanja:
Mlomo wa Mbuya, Shambaa: Paramoyo
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Aerva lanata (L.) Juss. ex Schult.: Perennial herb, sometimes flowering in the first
year, frequently woody and suffrutescent below, prostrate to decumbent or erect
(occasionally scandent), stiff or weak and straggling, (0.1–)0.3–2 m, branched from
the base and often also above (upper branches short to long and slender even on the
same plant), but stem and main basal branches often not further branched. Stem and
branches terete, striate, densely lanate with whitish shaggy hairs, more rarely tomen-
tose or canescent. Leaves alternate, round to lanceolate- or ovate-elliptic, shortly or
more longly cuneate at the base, rounded and apiculate to acute at the apex, commonly
densely lanate or canescent on the lower surface and more thinly so above, sometimes
subglabrous on the upper surface, rarely glabrous throughout or thickly lanuginose,
those of the main stem 10–50 5–35 mm, those of the branches and upper part of the
stem reducing and often becoming very small, petioles from 2 cm down to almost
none. Spikes sessile, solitary or usually in axillary clusters on the main stems or long to
very short axillary branches, 0.4–1.5( 2) 0.3–0.4 cm, divergent, cylindrical, silky,
white to creamy, forming a long inflorescence which is leafy to the ultimate spikes,
bracts deltoid-ovate to oblong-ovate, 0.75–1 mm, membranous with a short arista
formed by the excurrent midrib, pilose, persistent, bracteoles similar or slightly
smaller, also persistent. Tepals densely lanate dorsally. Flowers ♀ or hermaphrodite
(probably also sometimes functionally ♂). Hermaphrodite flowers: outer 2 tepals
hyaline, oval-oblong, abruptly contracted at the tip to a distinct mucro formed by the
excurrent nerve, 0.75–1.75 mm without the mucro, inner 3 slightly shorter and
narrower, acute with a broad central green vitta along the midrib, which extends for
about three-quarters of the length of each tepal and is often furnished with a thickened
border, style short, stigmas very short and patent or divergent, anthers perfect,
probable male flowers similar but stigmas reduced, subcapitate or very short, scarcely
papillose. Female flowers: tepals sometimes similar to those of the hermaphrodite
flowers but commonly longer and narrower, tapering above, the outer up to 2.25 mm,
style slightly longer, stigmas distinctly longer and often equaling the style, linear,
divergent, or suberect, anthers absent, filaments reduced. Capsule rotund, compressed
1 mm Seed reniform, 0.6–0.8 mm, black, shining, the testa almost smooth in the
center, faintly reticulate around the margin. Grows from cultivated and disturbed
ground to woodland, bushland and grassland, at swamp and forest edges, open lava
screes and lava boulder-strewn hillsides, coastal sands, 0–2030 m. Widespread in the
drier parts of the tropics and subtropics of the Old World – in Africa from Sierra Leone
across to Egypt, S. to South Africa (rare) and Madagascar, also in Seychelles, Chagos
Archipelago, Asia from Arabia to Malaysia, Malayan Islands, Philippines, and New
Guinea (Townsend 1985).
Aerva lanata (L.) Juss. ex Schult. 121
Aerva lanata: Leaf infusion used to treat sore eyes, also for washing babies with
malaria. The smoke is used as inhalant. The whole plant is used to prevent lactation.
Roots are used to treat snakebites (Bekalo et al. 2009; Kokwaro 2009). In the
Caucasus used for kidney problems (Bussmann et al. 2017). As some in social
settings (Mohagheghzadeh and Faridi 2006). In India, used to treat cough, sore
throat and wounds (Muthu et al. 2006), also as anthelminthic, to treat boils and cuts,
burns, cholera, diarrhea, dropsy, ear problems, fever, headache, malaria, skin prob-
lems, snakebites, swellings, and white discharge in the urine (Verma et al. 2007).
Aerva sanguinolenta is used in India as diuretic (Kumar et al. 2011). Aerva javanica
serves in Ethiopia to treat cancer (Teklehaymanot 2009), bloody diarrhea, as
anthelminthic, and for chest pain (Teklehaymanot and Giday 2010).
References
Bekalo TH, Demissew Woodmata S, Asfaw Woldemariam Z. An ethnobotanical study of medicinal
plants used by local people in the lowlands of Konta Special Woreda, southern nations,
nationalities and peoples regional state, Ethiopia. J Ethnobiol Ethnomed. 2009;5:26. https://
doi.org/10.1186/1746-4269-5-26.
Bussmann RW, Paniagua Zambrana NY, Sikharulidze S, Kikvidze Z, Kikodze D, Tchelidze D,
Batsatsashvili K, Hart RE. Plants in the spa – the medicinal plant market of Borjomi, Sakartvelo
Republic of Georgia, Caucasus. Indian J Tradit Knowl. 2017;161:25–34.
habitation in Garhwal Himalayan, India. J Med Plant Res. 2011;511:2252–60.
Mohagheghzadeh A, Faridi P. Medicinal smokes. J Ethnopharmacol. 2006;108:161–84.
Teklehaymanot T, Giday MM. Quantitative ethnobotany of medicinal plants used by Kara and Kwego
Townsend CC. Amaranthaceae. In: Polhill RM, editor. Flora of tropical East Africa. Rotterdam/
Boston: A.A.B.alkema; 1985.
Afrocanthium keniense (Bullock) Lantz
Afrocanthium lactescens (Hiern) Lantz
Canthium glaucum Hiern.
RUBIACEAE

Grace N. Njoroge
Synonyms
Afrocanthium keniense (Bullock) Lantz: Canthium keniense Bullock

Afrocanthium lactescens (Hiern) Lantz: Canthium lactescens Hiern., Canthium
randii S. Moore, Canthium umbrosum Hiern, Plectronia lactescens (Hiern)
K. Schum., Plectronia psychotrioides K. Schum. ex De Wild., Plectronia randii
(S. Moore) Eyles, Plectronia umbrosa (Hiern) K. Schum.
Canthium glaucum Hiern.: Plectronia glauca (Hiern.) K. Schum.
Local Names
Afrocanthium keniense: Kikuyu: Mubiru-Wathi, Luhya: Esiobari (Beentje 1994;

Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Afrocanthium lactescens: Kipsigis: Kamolwet, Luhya: Kunkomwo, Maa:

Olkumi, Olmaldei, Marakwet: Kaibetwa, Kerbetwa, Pokot: Kamolwo, Buterwo,
Putere, Samburu: Larugach, Tugen: Keipetwo, Turkana: Amalerh (Beentje 1994)
Canthium glaucum: Swahili: Mtengenji, Giriama: Mfuranje (Beentje 1994).
Botany and Ecology
Afrocanthium keniense (Bullock) Lantz: Shrub or tree 2.5–8.5 m tall, glabrous,

young stems with fawn to greyish bark, lenticels obscure. Leaves restricted to new
growth at apex of stems and lateral shoots, blades elliptic, 6–14 cm long, 2.8–6.5 cm
wide, acute to acuminate at apex, acute to obtuse at base, midrib and 5–6 main pairs
of lateral nerves frequently turning black when dry, tertiary nerves apparent, rather
coarsely reticulate, domatia present as conspicuous tufts of whitish hair, petiole
3–5 mm long, stipules triangular, acuminate or somewhat lobed at apex, 0.5–1 cm
long, resembling bark when mature, without conspicuous hairs inside. Flowers
5-merous, restricted to top 1–4 nodes of woody stem, borne in pedunculate
5–20-flowered cymes, arranged to one side on each of the two main branches,
or sub-umbellate in few-flowered inflorescences, peduncle 0.3–1.5 cm long,
usually with lines of pubescence, pedicels 0.5–1 mm long, bracteoles inconspicuous.
Calyx-tube 1.5 mm long, limb reduced to a truncate rim 0.5 mm long, sometimes
obscurely dentate. Corolla yellowish green, tube 1.75–2 mm long, finely pubescent
at throat, lobes ovate, 1.75–2 mm long, 1.5 mm wide at base, acute and thickened at
apex, spreading or reflexed. Style slightly longer than corolla-tube, stigmatic knob
hemispherical, ribbed below with stigmatic surface clearly apparent on upper face.
Fruit heart-shaped, 1.3–1.4 cm long and wide, pedicels thickening and lengthening
to 3–4 mm long, pyrene narrowly obovoid, 1.4 cm long, 6.5 mm wide, coarsely
rugulose. In forest clumps, 1675–1800 m.
Afrocanthium lactescens (Hiern) Lantz: Tree 2.5–9 m tall, stems thick, often with
short internodes, covered with dark grey to reddish bark, glabrous or occasionally
pubescent, lenticels usually apparent. Leaves restricted to new growth at apex of
main shoots and short lateral branches, usually only one pair on each branch, blades
7–18 5–13 cm, broadly elliptic to round, acute to sub-acuminate at apex, acute to
obtuse or truncate at base, glabrous or occasionally pubescent beneath and sparsely
pubescent above, lateral nerves in 8–10 main pairs, acutely angled, often turning
black when dry, tertiary nerves apparent but not conspicuous, domatia usually
present, but not markedly conspicuous, petioles 5–20 mm long, stipules 6–12 mm
long, broadly triangular to ovate, often long-acuminate when young, resembling
bark when mature, not conspicuously hairy within. Flowers (4)5-merous, 20–50 in
pedunculate cymes, restricted to leafless nodes of previous season’s growth, cymes
branched 3–5 times, but ultimate segments with flowers arranged on one side,
peduncle 1–35 mm long, pubescent on either side, pedicels 0.5–3 mm long, bracte-
oles inconspicuous. Calyx tube 1.5 mm long, glabrous or sometimes sparsely
Afrocanthium keniense (Bullock) Lantz . . . 125
pubescent, limb reduced to a rim. Corolla cream to yellowish, tube 2–2.5 mm long,
finely pubescent at throat, rather wider than calyx tube, lobes 1.5–3 mm long, 1–
2 mm wide at base, triangular-ovate, acute, thickened towards the apex and some-
times appendaged inside, spreading or reflexed. Style shortly exceeding corolla tube.
Pollen presenter hemispherical, strongly ribbed below, 1 mm wide. Fruit yellowish,
edible, 7–12 8–12 mm, square in lateral view, straight-sided or tapering at base,
strongly bilobed and indented at apex.
Canthium glaucum Hiern.: Shrub, sometimes scandent, 1–5 m tall, young branches
glabrous or rarely pubescent, often lenticellate, armed with spines up to 15(21) mm
long, situated above brachyblasts (very abbreviated lateral branches), apparently
not developing into lateral branches, bark light to dark grey or brown. Leaves
restricted to lateral spurs and apex of stem, blades 2.4–6 1.4–3 cm, narrowly to
broadly elliptic, acute to subacuminate or sometimes obtuse at apex, acute to
cuneate at base, strongly or moderately discolorous, glabrous, or glabrous above
and sparsely pubescent at least on nerves beneath, or rarely pubescent on both
faces, lateral nerves usually in 3 main pairs, tertiary nerves apparent, domatia
present as small tufts of hair, petiole 1–1.6 mm long, glabrous to sparsely pubes-
cent, or rarely pubescent, stipules 1–4 mm long, triangular, apiculate, densely
pubescent within. Flowers (4)5-merous, in 2–8-flowered cymes, cymes subsessile
to pedunculate, peduncles up to 10 mm long, pedicels 3–10 mm long, glabrous or
sparsely pubescent, bracteoles inconspicuous. Calyx tube 1 mm long, glabrous or
rarely pubescent, limb divided often almost to base into triangular or narrowly
triangular lobes, 0.5–1 mm long. Corolla glabrous or rarely pubescent outside,
tube 1–1.25 mm long, with a ring of deflexed hairs at the throat and pubescent
immediately above, lobes 2–2.5 1.5 mm, triangular-lanceolate, scarcely apicu-
late to apiculate. Style slightly exceeding the corolla tube. Pollen presenter spher-
ical, 0.75 mm across, disk not prominent. Fruit pale red, edible, 9–12 mm long and
wide, square in outline, scarcely indented at apex, hardly tapered at base. Pyrene
7–12 4–6 mm, ellipsoid with ventral face flattened, triangular at point of
attachment, crested at the apex, rugulose (Verdcourt 1976; Bridson and Verdcourt
1988, 1991).
Both Canthium and Afrocanthium are common undergrowth species in all types
of Afromontane forests (Bussmann 2002a, b) (Figs. 1, 2, 3, 4, 5, 6, and 7).
Afrocanthium keniense: Leaves are pounded, mixed with fat, and rubbed over
new-born babies, and applied for skin swellings (Kokwaro 2009).
Afrocanthium latescens: Root decoction is drunk as purgative (Kokwaro 2009).
Some species are being used in Madagascar to treat Epilepsy (Razafindraibe et al.
2013). Canthium oligocarpum serves to treat eczema (Yineger et al. 2008).
Fig. 1 Canthium
cf. oligocarpum (Rubiaceae).
Mt. Kenya National Park,
Kenya. (Photo
R.W. Bussmann)
Fig. 2 Canthium
cf. oligocarpum (Rubiaceae).
Kenya. (Photo
R.W. Bussmann)
Fig. 3 Canthium sp. cf.

(Rubiaceae) Bale Mountains
National Park, Ethiopia.

(Rubiaceae). Bale Mountains
Local Food Uses
Afrocanthium latescens: Fruit edible (Beentje 1994).

Cantium glaucum: Fruit edible (Beentje 1994).
The fruits of all species are edible (Bussmann et al. 2011).


Afrocanthium latescens: Wood termite resistant (Beentje 1994).

The leaves of all species are browsed by cattle and wildlife (Bussmann et al.
2011).

References
Bridson DM, Verdcourt B. Flora of tropical East Africa: Rubiaceae (part 2). Rotterdam: Balkema;
1988.
Bridson DM, Verdcourt B. Flora of tropical East Africa: Rubiaceae (part 3). Rotterdam: Balkema;
1991.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002a;91(1/2):27–79.
Appendices 1–7 @ www.naturekenya.org/JournalEANH.htm.
Bussmann RW. Succession and regeneration patterns in east African Mountain forests – a review.

Razafindraibe HM, Razafiarison ZL, Raharimalala F, Rakotoarivony F, Randrianarivony T,
Rakotoarivelo N, Randrianasolo A, Kuhlman A, Bussmann RW. Medicinal plants used by
women from Agnalazaha littoral forest southeastern Madagascar. J Ethnobiol Ethnomed.
2013;9:73.
Verdcourt B. Flora of tropical East Africa: Rubiaceae (part 1). London: Crown Agents for Overseas
Governments & Administrations; 1976.
Yineger H, Kelbessa E, Bekele T, Lulekal E. Plants used in traditional management of human
ailments at Bale Mountains National Park, southeastern Ethiopia. J Med Plants Res.
2008;26:132–53.
Ageratum conyzioides L.
ASTERACEAE

Grace N. Njoroge
Synonyms
Ageratum conyzioides L.: Ageratum album Willd. ex Steud., Ageratum arsenei

B.L. Rob., Ageratum ciliare L., Ageratum conyzioides var. inaequipaleaceum Hieron.,
Ageratum cordifolium Poir., Ageratum hirtum Lam., Ageratum humile Salisb., Agera-
tum latifolium Cav., Ageratum latifolium var. galapageium B.L. Rob., Ageratum micro-
carpum (Benth.) Hemsl., Ageratum pinetorum (L.O. Williams) R.M. King & H. Rob.,
Ageratum suffruticosum Regel, Alomia microarpa (Benth.) B.L. Rob., Alomia
pinetorum L.O. Williams, Calestina microcarpa Benth., Carelia conyzioides (L.)
Kuntze, Coelestina microcarpa Benth., Eupatorium conyzioides (L.) E.H.L. Krause.
Local Names
Ageratum conyzoides: Kikuyu: Gathenge, Luhya: Ilusa, Swahili: Kundambara,

Chagga: Matawana, Luo: Oluoro-Chieng’, Padhola: Otikidiel (Gachati 1989;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Ageratum conyzioides L.: Herbs, annual, 50–100 cm tall, sometimes less than
10 cm, with inconspicuous main root. Stems robust, ca. 4 cm in diam. at base,
simple or branched from middle, stems and branches reddish, or green toward apex,
white powdery puberulent or densely spreading long tomentose. Leaves often with
axillary abortive buds, petiole 1–3 cm, densely white spreading villous, median
leaves ovate, elliptic, or oblong, 3–8 2–5 cm, upper leaves gradually smaller,
oblong, sometimes all leaves small, ca. 1 0.6 cm, both surfaces sparsely white
puberulent and yellow gland-dotted, basally 3-veined or obscurely 5-veined, base
obtuse or broadly cuneate, margin crenate-serrate, apex acute. Capitula small, 4–14,
in dense terminal corymbs, peduncle 0.5–1.5 cm, powdery puberulent, involucre
campanulate or hemispheric, about 5 mm in diam., phyllaries 2-seriate, oblong or
lanceolate-oblong, 3–4 mm, glabrous, margin lacerate, corollas 1.5–2.5 mm, gla-
brous or apically powdery puberulent, limb purplish, 5-lobed. Achenes black,
5-angled, 1.2–1.7 mm, sparsely white setuliferous, pappus scales 5 or awned,
1.5–3 mm. Flowering and fruiting year-round (Wu et al. 1994–2013) (Fig. 1).
Fig. 1 Ageratum conyzioides

(Asteraceae), Pusac,
R.W. Bussmann and
N.Y. Paniagua_Zambrana)
Ageratum conyzioides L. 133
Ageratum conyzioides: Leaf juice is highly hemostatic and applied to bleeding

wounds. Also used to treat sore-eyes, and intestinal problems, cough, and
stomach-ache. The root decoction can be used for chest complaints. As steam bath
the plant serves for colds in Ethiopia, and the crushed leaves are used to stop
bleeding (Yineger et al. 2008). IOt is also applied externally for back pain (Giday
et al. 2010) and shows antibacterial properties (Geyid et al. 2005). In Madagascar
used for stomach pain (Razafindraibe et al. 2013).
The leaf paste is applied in India to cuts and wounds (Kumar et al. 2011; Raj et al.
2018; Debbarma et al. 2017), as well as antibacterial and anti-inflammatory (Singh
et al. 2017), and as anthelminthic (Raj et al. 2018). In addition the species serves for
leprosy, headache, muscle pain, piles, anus prolapse, ringworm, scabies, skin prob-
lems, and uterine disorders (Verma et al. 2007). In Nepal the plant is used as
antiviral, a rather new incorporation in the local pharmacopoeia (Kunwar and
Bussmann 2009; Kunwar et al. 2015). Interestingly, no uses are reported by the
Maasai (Bussmann et al. 2006), and Oromo (Bussmann et al. 2011).
Ageratum conyzioides: The leaves are used as fish poison (Neuwinger 2004). Used
to kill headlice (Verma et al. 2008). An extract of Ageratum houstonianum is used
for tick control (Wanzala 2017).
References
Geyid A, Abebe D, Debella A, Makonnen Z, Aberra F, Teka F, Kebede UK, Yersaw K, Biza T,
Haile Mariam B, Guta M. Screening of some medicinal plants of Ethiopia for their anti-
microbial properties and chemical profiles. J Ethnopharmacol. 2005;97:421–7.
Ethiopia. J Ethnopharmacol. 2010;1321:75–85. https://doi.org/10.1016/j.jep.2010.07.046.
Kunwar RM, Bussmann RW. Medicinal, aromatic and dye plants of Baitadi and Darchula Districts,
Biodiversität, Naturausstattung im Himalaya, vol. III; 2009. p. 475–89, Erfurt.
Kunwar RM, Acharya RP, Chowdhary CL, Bussmann RW. Medicinal plant dynamics in indigenous
medicines in Farwest Nepal. J Ethnopharnacol. 2015;163:210–9.
Raj AJ, Biswakarma B, Pala NA, Shukla G, Vineeta V, Kumar M, Chakravarty S, Bussmann RW.
Indigenous uses of ethno-medicinal plants among forest dependent communities of Northern
Bengal, India. J Ethnobiol Ethnomed. 2018;14(1):8. https://doi.org/10.1186/s13002-018-0208-9.
women from Agnalazaha littoral forest (Southeastern Madagascar). J Ethnobiol Ethnomed.
2013;9:73.
Singh A, Nautiyal MC, Kunwar RM, Bussmann RW. Ethnomedicinal plants used by local inhab-
itants of Jakholi Block, Rudraprayag district, Western Himalaya, India. J Etnobiol Ethnomed.
2017;13(49). https://doi.org/10.1186/s13002-017-0178-3.
Wanzala W. Potential of traditional knowledge of plants in the management of arthropods in
livestock industry with focus on acari ticks. Evid-Based Complement Altern Med. 2017. Article
ID 8647919, 33 pages. https://doi.org/10.1155/2017/8647919.
Wu Z, Raven PH, Hong D e. Flora of China. Beijing/St. Louis: Science Press/Missouri Botanical
Garden Press; 1994–2013.
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plants Res.
2008;26:132–53.
Albizia anthelminthica Brogn.
Albizia gummifera (J.F. Gmel.) C.A. Sm.
FABACEAE

Grace N. Njoroge
Synonyms
Albizia gummifera (J.F. Gmel.) C.A. Sm.: Albizia sassa (Willd.) Chiov.
Local Names
Albizia anthelminthica: Kikuyu: Muguta, Swahili: Mporojo, Bono: Hobocho,

Borana: Howacho, Kamba: Mwowa, Kyalundathi, Muowa, Kyoa, Muoa, Maa:
Ol-Emugutan, Marakwet: Kitangwa, Oromo: Habachu, Pokot: Kamakitan,
Mukotonwo, Samburu: Olmukutan, Somali: Habasho, Reidep, Reidup, Gogo:
Mkutoni, Swahili: Mporojo (Beentje 1994; Gachati 1989; Kokwaro 2009).
Albizia gummifera: Kikuyu: Gathenge, Swahili: Mkenge, Mchapia tumbili,
Mchani mbao, Mshai, Ilelewa: Motemwelo, Kamba: Methia, Kipsigis/Nandu:
Seet, Seyet, Kavirondo: Mughonzulu, Kikiyu/Meru: Mukurue, Luhya:
Mukhonzuli, Kisii: Omugonkoro, Maa: Ol-Osepakupes, Marakwet/Tugen: Se,
Set, Samburu: Sogore, Sagorogurri, Taveta: Kiririgwi, Taita: Masarawaxchi,
R. W. Bussmann (*)
G. N. Njoroge

Turkana: Ekokwait, Chagga: Mboromo (Beentje 1994; Gachati 1989; Kokwaro

2009), English: Peacock flower, Smooth-bark flat-crown
Botany and Ecology
Albizia anthelminthica Brogn.: Shrub or tree to 10 m high, crown often somewhat

rounded. Bark grey to reddish-brown, smooth, young branchlets shortly pubescent
or glabrous, lenticellate, often forming abbreviated divaricate spine-tipped lateral
shoots. Leaves glabrous to shortly pubescent: petiole 0.5–2 cm long, a small gland
usually present midway along the petiole or just below the junction of the lowest
pinna pair, rhachis 0–3.5 cm long, usually terminating in a short rigid persistent
deflexed hook, a small gland often present near the junction of the top pinna pair,
pinnae 1–3(4) pairs, rhachillae 1.5–5 cm long, usually terminating in a short rigid
persistent deflexed hook, often a single stipel similarly bent near the base, leaflets (1)
2–4(6) pairs, (5)8–25(30) 4–18(24) mm (in our area), obliquely obovate or elliptic
to sub-rotund, mucronate apically, venose, glabrous or sparingly pubescent beneath,
especially on the midrib and lateral nerves. Inflorescences globose, peduncles
1–3 cm long, glabrous or sparingly pubescent. Flowers usually borne on leafless
branchlets, whitish, on pedicels 0.5–2.5 mm long, bracteoles rapidly deciduous.
Calyx greenish, 3–5 mm long, usually deeply slit unilaterally, glabrous throughout
or with a tuft of hairs at the apex of each lobe, sometimes sparingly pubescent
throughout. Corolla greenish, 6–12 mm long, glabrous throughout or puberulous
near the apices and margins of the lobes, lobes up to 5 mm long. Stamens 1.5–2.8 cm
long, united basally for up to 4 mm, tube not exserted beyond the corolla. Ovary 3 mm
long, glabrous, shortly stipitate. Pods straw-colored to light brown when mature, (6)
8–18 1.5–2.8 cm, oblong, valves thin, glabrous or occasionally puberulous all over,
venose, umbonate over the seeds, longitudinally dehiscent. Seeds 9–13 mm in diam.,
rounded, flattened (Brenan 1959).
Albizia gummifera (J.F. Gmel.) C.A. Sm.: Medium-sized deciduous tree up to 30 m
tall, bole straight and cylindrical, up to 75( 100) cm in diameter, without buttresses
or with small, thick, buttresses, bark yellowish to grey, usually smooth, inner bark
with clear gum, crown flattened, young branches finely pubescent, but soon
glabrescent. Leaves alternate, bipinnately compound with (3–)5–7( 8) pairs of
pinnae, stipules lanceolate, up to 7 mm long, caducous, petiole 2.5–4.5 cm long,
near the base of upper side with a sessile gland, rachis 4.5–11 cm long, pubescent,
leaflets in 6–17 pairs per pinna, sessile, obliquely rhombic, up to 2 cm 1 cm, often
auricled at proximal side of base, obtuse to acute at apex, glabrous but with some
hairs on midrib and margins. Inflorescence an axillary head on a 2.5–5 cm long
peduncle. Flowers bisexual, regular, 5-merous, reddish white, almost sessile, sub-
tended by up to 6 mm long, caducous bracteoles, calyx obconical, 2–5 mm long,
minutely pubescent outside, corolla 7–12 mm long, with 4.5–6 mm long tube,
pubescent outside, stamens numerous, 2.5–3.5 cm long, united into a tube for
most of their length, white in lower part and reddish in upper part, ovary superior,
ellipsoid, 1.5–2.5 mm long, gradually tapering into a 2.5–3.5 cm long style. Fruit an
oblong, flat pod 10–21 cm 2–4 cm, with stipe c. 1 cm long, glabrous, transversely
Albizia anthelminthica Brogn. . . . 137
Fig. 1 Albizzia gummifera (Fabaceae), Loita forest, Mara, Kenya. (Photo R.W: Bussmann)
veined, pale brown to reddish brown when ripe, opening with 2 papery valves,
9–12–seeded. Seeds flattened globose to broadly oblong, 8–12 mm 7–10 mm.
Albizia gummifera is widespread, occurring from eastern Nigeria to western Ethiopia
and Kenya, and south to Zimbabwe and Mozambique, also in central Madagascar
(Brenan 1959) (Figs. 1 and 2).
Albizzia anthelminthica: Bark infusion used as emetic, antimalarial and anthelmin-

tic (Beentje 1994; Kokwaro 2009). Also used as purgative. Roots used to treat fever,
gonorrhea, as emetic and purgative, and as aphrodisiac (Muthee et al. 2011; Nankaya
et al. 2019, 2020; Njoroge and Bussmann 2006). However, the plant is toxic, and
overdose can lead to death (Kokwaro 2009). Used for bone and joint pain, stomach
problems, as anthelminthic, for female health, malaria, and gonorrhea (Delbanco
et al. 2017). Sometimes used for asthma and diarrhea (Gafna et al. 2017).
Albizzia gummifera: In Kenya a bark infusion is taken to treat malaria (Beentje
1994). The twigs are used as toothbrush. Bark decoctions are used for menorrhagia,
postpartum hemorrhage, and to prevent abortion, also for whooping cough tonsilitis,
pneumonia, anthrax, kidney problems, anemia, and heart problems. The bark ash can
also be used. Roots are used to treat venereal diseases, the steam of boiled roots is
used for sore eyes. Root decoction used for measles and gastrointestinal problems
(Kokwaro 2009). In Uganda it is used to hasten parturition. The pounded bark is
used in Kenya as a snuff to treat headache, and in Tanzania, it is applied externally to
treat scabies. In eastern DR Congo a bark maceration is used as a body wash and
drink to treat psoriasis. The roots and leaves are purgative and are used in Mada-
gascar to treat diarrhea and eye troubles. In Malawi roots are soaked in water for
10 min, and the liquid is drunk to relieve the pain caused by sprains. In Madagascar,
Fig. 2 Albizzia gummifera

(Fabaceae), sapling, Loita
forest, Mara, Kenya. (Photo
R.W: Bussmann)
a leaf decoction is reputed to have antitussive activity and is administered to treat

asthma; leaves are applied to sores and fractures. In Kenya pounded roots are added
to a bath to treat skin diseases, and an extract of crushed pods is drunk to treat
stomach-ache. In Uganda, roots are used to treat sleeping sickness. Sometimes used
to relieve fatigue and cough in Madagascar (Razafindraibe et al. 2013). In Ethiopia
used for the treatment of ascaris, gonorrhea, and syphilis (Geyid et al. 2005). Albizia
zygida bark is used for malaria (Tabuti 2008), Albizia malacophylla for cough
(Giday et al. 2007).
The bark of Albizia procera is commonly used in medicine in India (Reang et al.
2016), while Albizia lebbek serves for boils, carbuncle, diarrhea, dysentery, gonor-
rhea, eyesores, gum ulcer, night blindness, and piles (Verma et al. 2007).
Albizzia anthelminthica: Wood hard (Beentje 1994). Nark or root decoction as

anthelminthic for livestock (Bussmann 2006; Kokwaro 2009). Used for vetetinary
diseases (Delbanco et al. 2017).
Albizia anthelminthica Brogn. . . . 139
Albizzia gummifera: The is used for light construction, furniture, cabinet work, and
various implements. It is also suitable for mine props, light flooring, joinery, interior
trim, panelling, framing, toys and novelties, sporting goods, boxes, crates, carvings,
peeled and sliced veneer, plywood, hardboard and particle board (Beentje 1994).
Logs are traditionally used for the construction of canoes. The wood is sometimes
used as firewood and for making charcoal. The wood pulp is suitable for paper
production. Albizia gummifera is planted as an ornamental shade tree. It is valued as
a shade tree for crops, e.g., in coffee plantations in Ethiopia, tea plantations in
Malawi, and vanilla plantations in Madagascar and is also planted or retained for
soil conservation and improvement. The gum from the bark is sometimes used in
confectionery. The foliage is browsed by goats. The flowers produce nectar for bees,
which often build nests in the trunk. The leaves are said to quicken the ripening of
bananas. Root decoction used for measles, foot and mouth disease and gastrointes-
tinal problems in cattle (Kokwaro 2009). The hard wood and bark are used as
excellent firewood (Bussmann et al. 2006; Kiefer and Bussmann 2004, 2008), the
wood serves to make weapons, walking sticks and especially spears (Bussmann et al.
2006). Albizia coriacea bark smoke is used as mosquito repellent (Tabuti 2008).
Albizia aurispara serves for construction and cultural uses in Madagascar
(Randrianarivony et al. 2016a, b).
References
Brenan JPM. Flora of tropical East Africa: Leguminosae, subfamily Mimosoideae. London: Crown
Delbanco AS, Burgess ND, Cuni-Sanchez A. Medicinal plant trade in Northern Kenya: economic
importance, uses and origin. Econ Bot. 2017;71(1):12–31.
Gafna DJ, Dolos K, Mahiri IO, Mahiri JG, Obando JA. Diversity of medicinal plants and anthropo-
genic threats in the Samburu central sub-county of Kenya. Afr J Tradit Complement Altern Med.
2017;14(5):72–9. https://doi.org/10.21010/ajtcam.v14i5.1010.21010/ajtcam.v14i5.10.
Geyid A, Abebe D, Debella A, Makonnen Z, Aberra F, Teka F, Kebede, Urga K, Yersaw K, Biza T,
2004;6(2):161–9.
anthelmintic and other medicinal plants traditionally used in Loitoktok district of Kenya. J
Ethnopharmacol. 2011;135:15–21.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Sustainability of the Loita Maasai Childrens’

ethnomedicinal knowledge. Sustainability. 2019;11:5530; https://doi.org/10.3390/su11195530.
2020;9:44. https://doi.org/10.3390/plants9010044.
Njoroge GN, Bussmann RW. Herbal usage and informant consensus in ethnoveterinary manage-
ment of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006;108:332–9.
Randrianarivony, et al. Value of useful goods and ecosystem services from Agnalavelo sacred forest
and their relationships with forest conservation. 2016a.
Randrianarivony TN, Andriamihajarivo TH, Rakotoarivony F, Rabarimanarivo M,
Randrianasolo A, Bussmann RW. Guide des plantes utiles d’Analavelona et de ses environs,
vol. 1. St. Louis: William L. Brown Center, MBG; 2016b. ISBN 978-0-9960231-4-6.
women from Agnalazaha littoral forest Southeastern Madagascar. J Ethnobiol Ethnomed.
2013;9:73.
inal plants by traditional herbal healers in Reang tribe of south district Tripura, India. Med
Aromat Plants. 2016;5:2.
Tabuti JRS. Herbal medicines used in the treatment of malaria in Budiope county, Uganda.
Alchemilla argyrophylla Oliv.
Alchemilla johnstonii Oliv.
ROSACEAE

Grace N. Njoroge
Synonyms
Alchemilla argyrophylla Oliv.: Alchemilla keniensis Baker f. ex De Wild.;

Alchemilla rammellii T.C.E. Fr.; Alchemilla robertii T.C.E. Fr.
Alchemilla johnstonii Oliv.: Alchemilla aberdarensis De Wild.; Alchemilla
alluaudii De Wild., Alchemilla ducis-aprutii Cortesi; Alchemilla geranioides
Rolfe; Alchemilla jaegeri Engl.; Alchemilla pentagona T.C.E. Fr.; Alchemilla
pseudomildbraedii T.C.E. Fr.; Alchemilla ulugurensis Engl.
Local Names
Alchemilla argyrophylla: Kikuyu: Gitamuhera (Gachati 1989); English: Ladies

mantle
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Alchemilla argyrophylla Oliv.: Afroalpine shrub, up to 2.5 m tall. Leaves densely

covered in silky silvery hairs. Flowers in axillary groups, normally 5–7, less than
2 mm long. Fruit hidden in the calyx (Beentje 1994; Graham 1960) (Fig. 1).
Alchemilla johnstonii Oliv.: A low straggling shrub with prostrate stems and erect
or procumbent branches, often dominant in carpeting masses; sometimes with
small and densely congested leaves. Stems woody or sometimes herbaceous at
the extremities, rooting rather infrequently at the nodes, usually reddish-brown,
covered (often thickly) with long, ascending or spreading, whitish hairs, rarely
glabrous. Stipules 3–8 ( 20) mm long, the lower half (approx.) membranous, the
upper half a spreading, foliaceous, lobed limb. Leaves petiolate, often leathery,
circular to reniform in outline, 3–5 ( 7)-lobed, 5–40 mm long and usually broader
than long; lobes often up to 1 cm deep, obtuse or rounded, apically entire or
dentate, sometimes appearing emarginate owing to the reduced size of the central
tooth, often markedly folded along the midribs and with the basal sinus varying
from wide to narrow and almost closed. Inflorescences usually short, simple or
branched; pedicels up to 3.5 mm long, sparsely hirsute with long, ascending hairs.
Calyx-lobes triangular to ovate-triangular, 1.5–2 mm long, with a few hairs on the
outside and margins; epicalyx-lobes lanceolate, shorter than the calyx-lobes.
Upland moor, moist bamboo-thicket, moor-grassland, on damp ground and often
Fig. 1 Alchemilla
argyrophylla (Rosaceae),
Naro Moru Route, Kenya.
(Photo: R. W. Bussmann)
Alchemilla argyrophylla Oliv. . . . 143
Fig. 2 Alchemilla fischeri

(Rosaceae), Mt. Kenya
National Park, Naro Moru
Route, Kenya. (Photo: R. W.
Bussmann)
in bogs; 2400–4260 m (Graham 1960). Common in the upper parts of Afromo-

ntane forests, especially Hagenia abyssinica dominated timberline forests
(Bussmann 2002, 2006a) (Figs. 2, 3, 4, 5, and 6).
Alchemilla haumannii leaves are chewed for stomach ache and applied to wounds
(Yineger et al. 2008).
Alchemilla species grow especially in higher altitudes, and thus are a typical
example for the fact that Afro-alpine species are very little used by the local
population (Bussmann 2006b), although are browsed by cattle (Bussmann et al.
2011; Luizza et al. 2013). Alchemilla abyssinica is used to treat blackleg in livestock
Fig. 3 Alchemilla fischeri

Bussmann)
Fig. 4 Alchemilla johnstonii

Bussmann)
Alchemilla argyrophylla Oliv. . . . 145
Fig. 5 Alchemilla johnstonii

Bussmann)
Fig. 6 Alchemilla
sp. (Rosaceae), Mt. Kenya
Bussmann)
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002;91(1/2):27–79.
Appendices 1–7 @ www.naturekenya.org/JournalEANH.htm
2006a;111:41–66.
Graham RA. Flora of Tropical East Africa: Rosaceae. London: Crown Agents for Overseas
Res Appl. 2013;11:315–39.
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plant Res.
2008;26:132–53.
Aloe sp.
Aloe vera (L.) Burm. f.
ASPHODELACEAE

Grace N. Njoroge
Synonyms
Aloe vera (L.) Burm. f.: Aloe barbadensis Mill., Aloe barbadensis var. chinensis
Haw., Aloe chinensis (Haw.) Baker, Aloe elongata Murray, Aloe flava Pers., Aloe
indica Royle, Aloe lanzae Tod., Aloe perfoliata var. barbadensis (Mill.) Aiton, Aloe
perfoliata var. vera L., Aloe vera var. chinensis (Haw.) A. Berger, Aloe vera var.
littoralis J. Koenig ex Baker, Aloe vulgaris Lam. Aloe rubescens DC.
Local Names
Aloe sp: Kikuyu: Kiruma, Luo: Ogaka, English, Aloe
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Aloe vera (L.) Burm. f.: Up to 1.5 m tall succulent. Flowers pedicellate, erect in bud,
pendent at anthesis, or Perianth of three outer and the three inner free or partially
fused subequal segments, cylindrical, rarely campanulate, rounded at the base or
tapering into the pedicel, sometimes constricted above the ovary, often trigonous,
sometimes slightly curved, rarely two-lipped, segments free towards the apex,
imbricate, the outer usually slightly longer than the inner. Stamens with flattened
filaments, anthers oblong, attached towards the base, exserted from the perianth-
tube. Ovary ovate-oblong, trigonous, three-locular, placentation central, style slen-
der, longer than the stamens, stigma capitate, very small. Capsule held erect,
sometimes stipitate, very obtusely three-angled, sub-globose to oblong, papyraceous
or slightly woody, dehiscing from the apex. Seeds many, in two longitudinal rows in
each locule, three-angled, sometimes obscurely so, the angles extended into papery
wings with two wings distinctly longer than the third. Perennial, acaulescent or
caulescent herbs, shrubs or trees, solitary (not forming basal offshoots) or suckering
(forming basal offshoots) to produce dense clumps or loose groups of plants. Roots
thickened. Leaves crowded spirally into rosettes, occasionally distichous especially
in young plants, usually noticeably succulent, with colorless or often yellow, brown
or purple, sometimes pungent and then sometimes poisonous sap, lamina linear,
lanceolate or deltoid, base sheathing, apex usually spine-tipped, margin with spine-
tipped teeth, or sometimes cartilaginous or ciliate, rarely entire, surface glabrous,
smooth or roughened, sometimes the lower or both surfaces spiny, uniformly green,
or often white-spotted, sometimes striated. Inflorescence pedunculate with flowers
lax to compact in one-several terminal and axillary, simple or paniculately arranged
racemes, inflorescence stalk (peduncle) stout, flattened at the base, with branches,
when present, usually from the middle and above, naked or with sterile bracts
below the racemes, racemes capitate to long-cylindrical, bracts scarious, rarely leaf-
like. Aloe vera is cultivated around the world. It has escaped from cultivation and
become naturalized in the Mediterranean, north Africa, the Indian subcontinent,
South America and the Caribbean (Carter 1994) (Figs. 1, 2, and 3).
Aloe vera: Cultivated around the world as a crop for its colorless jelly-like leaf
parenchyma (aloe gel). Used for a variety of purposes such as psoriasis, burns,
frostbite, and sores caused by the Herpes simplex virus. Aloe can help to lower
cholesterol levels and can help to lower blood glucose levels. The green outer layer
of the leaves yields a bitter, yellow exudate traditionally used as a laxative. Very
widely used globally to treat inflammation (external), vaginal inflammation, vaginal
ulcers, vaginal cancer, hair growth, skin embellishment, cataracts, eyes, wounds,
burns, weight loss, gastritis, inflammation, diabetes, cough, bronchitis, kidneys,
ulcers, cholesterol, cancer, asthma, bile (Bussmann and Sharon 2006; Bussmann
2006; Bussmann et al. 2018). Aloe macrocarpa is used to treat dandruff, hemor-
rhoids, and eye diseases (Yineger et al. 2008). Aloe secundiflora is used for eyes and
Aloe sp. . . . 149
Fig. 1 Aloe
sp. (Asphodelaceae),
Huanchaco, Peru. (Photo:
R. W. Bussmann)
diarrhea (Gafna et al. 2017), as well as wound treatment and malaria (Muthee et al.
2011). Aloe volkensii can be used for stomachache and diarrhea (Gachati 1989;
Kokwaro 2009; Nankaya et al. 2019).
Aloe sp. are used for eye problems, tuberculosis, polio, and boiled in strengthen-
ing soups (Bussmann 2006). Also used to treat external injuries (Teklehaymanot
et al. 2010). Aloe pulcherrima and various other species are used for preventing
miscarriages and easing labor, and to treat malaria (Giday et al. 2007; Luizza et al.
2013). Aloe kedogensis is used to treat typhoid, skin problems, malaria, colds, ear
problems, wounds, and coccidiosis (Jeruto et al. 2008).
Aloe vera: Used for good luck, happiness, and to fend off evil spirits (Bussmann and
Sharon 2006). Aloe trichosandra is used to treat anthrax in livestock (Giday et al.
2003). Aloe pulcherrima serves to treat bloating and blackleg in livestock
(Teklehaymanot et al. 2007). Aloe pubescens also is used in veterinary medicine
(Wondimu et al. 2007). Aloe volkensii serves to remedy anaplasmosis, corridor
disease, heart water, sweating sickness, east coast fever, red water, and lumpy skin
disease in livestock (Kioko et al. 2015). Aloe buettneri is used as fish poison
(Neuwinger 2004). Various species are used to prevent ticks, lice, and fleas in
livestock (Wanzala 2017).
Fig. 2 Aloe
sp. (Asphodelaceae),
Huanchaco, Peru. (Photo:
R. W. Bussmann)
Fig. 3 Aloe vera

(Asphodelaceae), Trujillo,
Peru. (Photo: R. W.
Bussmann)
References
Bussmann RW, Sharon D. Traditional plant use in Northern Peru: tracking two thousand years of
healing culture. J Ethnobiol Ethnomed. 2006;2:47.
Aloe sp. . . . 151

Batsatsashvili K, Hart RE. Unequal brothers – plant and fungal use in Guria and Racha,
Sakartvelo Republic of Georgia, Caucasus. Indian J Tradit Knowl. 2018;171:7–33.
Carter S. Flora of Tropical East Africa: Aloaceae. Rotterdam: Balkema; 1994.
Gafna DJ, Dolos K, Mahiri IO, Mahiri JG, Obando JA. Diversity of medicinal plants and
anthropogenic threats in the Samburu central sub-county of Kenya. Afr J Tradit Complement
Altern Med. 2017;14(5):72–9. https://doi.org/10.21010/ajtcam.v14i5.10.
and Amhara peoples in Northwest Ethiopia. J Ethnopharmacol. 2007;110(3):516–25.
Kioko J, Baker J, Shannon A, Kiffner C. Ethnoecological knowledge of ticks and treatment of tick-
borne diseases among Maasai people in Northern Tanzania. Vet World. 2015;8:755–62.
Res Appl. 2013;11:315–39.
people around Debre Libanos monastery in Ethiopia. J Ethnopharmacol. 2007;111:271–83.
Teklehaymanot T, Giday M, Mutse M. Quantitative ethnobotany of medicinal plants used by Kara and
Kwego semi-pastoralist people in lower Omo River Valley, Debub Omo Zone, Southern Nations,
https://doi.org/10.1155/2017/8647919. Article ID 8647919, 33 pages.
Wondimu T, Asfaw Z, Kelbessa E. Ethnobotanical study of medicinal plants around “Dheeraa”
2008;26:132–53.
Amaranthus hybridus L.
Amaranthus spinosus L.
AMARANTHACEAE

Grace N. Njoroge
Synonyms
Amaranthus hybridus L.: Amaranthus chlorostachys var. hybridus S. Watson,

Amaranthus chlorostachys Willd., Amaranthus cruentus L., Amaranthus hecticus
Willd, Amaranthus hybridus var. chlorostachys Beck, Amaranthus hybridus var.
quitensis (Kunth) Covas, Amaranthus hypochondriacus L., Amaranthus laetus
Willd., Amaranthus paniculatus L., Amaranthus quitensis Kunth, Amaranthus
retroflexus subsp. quitensis (Kunth) Bolós & Vigo, Amaranthus retroflexus var.
chlorostachys A. Gray, Amaranthus retroflexus var. hybridus A. Gray, Amaranthus
strictus Willd., Galliaria hybrida (L.) Nieuwl.
Amaranthus spinosus L.: Amaranthus caracasanus Kunth, Amaranthus diacanthus
Raf., Amaranthus spinosus fo. inermis Lauterb. & K. Schum., Amaranthus spinosus
var. basiscissus Thell, Amaranthus spinosus var. circumscissus Thell., Amaranthus
spinosus var. indehiscens Thell., Amaranthus spinosis var. purpurascens Moq.,
Amaranthus spinosus var. pygmaeus Hassk., Amaranthus spinosus var. rubicaulis
Hassk., Amaranthus spinosus var. viridicaulis Hassk., Galliaria spinosa (L.) Nieuwl.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Amaranthus hybridus: English: Amaranth

Amaranthus spinosus: Swahili: Mchicha, Mbere: Mangani, English: Spiny ama-
ranth, Prickly amaranth, Spiny pigweed
Botany and Ecology
Amaranthus hybridus L.: Annual, mostly pale green, sometimes reddish-tinged,

20–80 cm high, stem erect, simple or branched, grayish by dense short hairs, slightly
angled, leaves rather large, ovate-rhombic, obtuse or slightly notched at apex,
smooth above, shortly hairy beneath, on the margin, and on the petiole, this as
long as or somewhat longer than the blade, flowers in glomerules crowded in a green
dense and very compact paniculate inflorescence, disposed mainly at the summit of
the stem, branched in lower part, oblong-cylindric or ovoid-pyramidal, bracts lan-
ceolate, slenderly long-pointed, nearly twice as long as the perianth, tepals 5, sur-
passing the fruit, in staminate flowers lanceolate, in pistillate oblong-linear,
spatulately enlarged toward apex, obtuse or retuse, mucronulate, whitish-
membranous, the slender pale gray midrib mostly failing to reach the apex, stamens
5, capsule 1-seeded, circumscissile, shorter than perianth, seed 1 mm long, black or
blackish-brown, very shining, sharp-margined. In kitchen gardens, fallow areas,
along roads and rivers, often in settlements in irrigated fields, from the lowland to
the mid-mountain belt, up to 2200 m (Figs. 1, 2, 3, 4, 5, and 6).
Amaranthus spinosus L.: Annual herb, erect or slightly decumbent, up to 1.5 m tall.
Stem stout, sometimes reddish, usually branched, angular, glabrous or increasingly
furnished above (especially in the inflorescence) with short or longer, multicellular,
flocculent hairs. Leaves glabrous, or thinly pilose on the lower surface of the primary
venation, long-petiolate, petioles up to 9 cm long, sometimes longer than the
lamina), lamina lanceolate-ovate to rhomboid-ovate, elliptic, lanceolate-oblong or
Fig. 1 Amaranthus hybridus

(Amaranthaceae) in a garden
in Khevsureti, Georgia.
(Photo: R. W. Bussmann and
N. Y. Paniagua-Zambrana)
Amaranthus hybridus L. . . . 155


lanceolate, 1.5–12 cm 0.8–6 cm, subacute or more commonly blunt or retuse at the
apex with a distinct, fine colorless mucro, cuneate or attenuate at the base, each leaf-
axil bearing a pair of fine and slender to stout and compressed spines up to 2.5 cm
long. Flowers green, in the lower parts of the plant in axillary clusters 6–15 mm
diameter, towards the ends of the stem and branches the clusters leafless and
approximated to form simple or the terminal at least, branched spikes up to 15 cm

Fig. 5 Amaranthus
sp. (Amaranthaceae) in a
garden in Kartli, Georgia.

var. paniculatus
in Kakheti, Georgia. (Photo:
R. W. Bussmann and N. Y.
Paniagua-Zambrana)
long and 1 cm wide. Lower flower clusters entirely female, as are the lower flowers
of the spikes, upper flowers of spikes male, mostly for the apical 1/4–2/3 of each
spike. Bracts and bracteoles deltoid-ovate, pale-membranous, with an erect, com-
monly reddish awn formed by the excurrent green midrib: bracteoles shorter than to
a little exceeding the perianth, commonly smaller than the bracts. Perianth segments
5, those of the female flowers 1.5–2.5 mm long, narrowly oblong or spathulate-
oblong, obtuse or acute, mucronulate, frequently with a greenish dorsal finn, those of
the male flowers broadly lanceolate or lanceolate-oblong, acute or acuminate, only
the midrib green. Stigmas (2)3, flexuose or reflexed, 1–1.5 mm long. Capsule ovoid-
urceolate with a short, inflated beak below the style-base, 1.5 mm long, regularly or
irregularly circumcissile or more rarely indehiscent, the lid rugulose below the beak.
Seed 0.75–1 mm across, black, compressed, shining, very faintly reticulate.
Amaranthus spinosus originates probably from lowland tropical South and Central
America and was introduced into other warmer parts of the world from about
1700 A.D. onwards. At present it occurs in all tropical and subtropical regions.
Amaranthus caudatus: Leaf juice applied for ear diseases. Pounded leaf paste
rubbed on skin problems (Kokwaro 2009). The seeds of Amaranthus caudatus
are ingested to treat amoebas (Yineger et al. 2008).
Amaranthus hybridus/Amaranthus spinosus: The species has numerous medicinal

uses. The root is known as an effective diuretic. In South-East Asia a decoction of the
root is used to treat gonorrhea and is also applied as an emmenagogue and antipy-
retic. In many countries, including those in Africa, the bruised leaves are considered
a good emollient and applied externally in cases of eczema, burns, wounds, boils,
earache and hemorroids. The plant ash in solution is used to wash sores. Plant sap is
used as an eye wash to treat ophthalmia and convulsions in children. The leaves are
used in the Ural as diuretic and for dysentery. An infusion is used to treat colitis,
intestinal colic, and as a laxative for constipation, as hemostatic to treat hemoptysis,
and menstrual and hemorrhoid hemorrhages, as anti-protist and anti-bacterial, to
treat guinea worm and jaundice, and to treat headaches (Fedorov 1984). The whole
plant, fresh is used to treat general pain and heat in the breast (Béjar et al. 2002;
Bussmann and Sharon 2006a, 2007a). Fresh leaves and stems are used to treat
inflammations (general) (Bussmann and Sharon 2006b; Bussmann and Sharon
2015), and have antibacterial properties (Bussmann et al. 2010, 2011a).
Amaranthus spinosus: The whole fresh plant is used to treat flu, headache, and fever
(Béjar et al. 2002; Bussmann and Sharon 2006a, 2007a), also to treat amoebic
diarrhea and vomiting (Teklehaymanot et al. 2010).
Amaranthus viridis is used as astringent and emollient (Kunwar et al. 2012,
2013; Ur-Rahman et al. 2018). Amaranthus dubius and Amaranthus graecizans are
used for bone fractures (Giday et al. 2010), and the species also serves to treat cancer
and boils (Jeruto et al. 2008).
Amaranthus spinosus serves as vermifuge, for dysepsis, as diuretic, for cataracts
and constipation (Umair et al. 2019).
Sometimes Amaranthus is used to treat bilharzia (Rabearivony et al. 2015), and
iontestinal parasites (Razafindraibe et al. 2013).
Local Food Uses
Amaranthus hybridus/Amaranthus spinosus: In tropical Africa and elsewhere

leaves and young plants are collected for home consumption as a cooked, steamed,
or fried vegetable, especially during periods of drought (Mekonen et al. 2015).
Leaves are occasionally found for sale on markets. It has a bitter taste and is usually
eaten in small quantities as a substitute when no other vegetables are available. The
young leaves are cooked as herb pie. Young stems are used as a source of vitamins.
The seeds are eaten and contain fat 8.9%, protein 19%, starch 41%. The leaves and
shoots are used sometimes eaten (Batsatsashvili et al. 2017; Bussmann et al. 2014,
2016a, b, 2017, 2018; Bussmann 2017; Kunwar et al. 2012).
Amaranthus hybridus/Amaranthus spinosus: Used as forage and said to increase

the yield of milk in cattle. However, the spines can cause injury to the mouths of
grazing animals and cases of poisoning in cattle have also been reported. In Uganda,
the ash of burnt Amaranthus plants is used as a tenderizer in cooking tough

vegetables such as cowpea leaves and pigeon peas. The ash is also used as a
vegetable salt and in southern Africa it is used as a snuff, alone or with tobacco.
As fodder for pigs. The leaves are boiled with salt in Ethiopia to fatten livestock
(Bussmann et al. 2011b; Luizza et al. 2013). Used to make brooms. The whole fresh
plant is used to cook a soup (Béjar et al. 2002; Bussmann and Sharon 2006a, 2007a).
References
Batsatsashvili K, Mehdiyeva N, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S,
Tchelidze D, Alizade V, Paniagua Zambrana NY, Bussmann RW. Amaranthus retroflexus
L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer International
Publishing; 2017.
Béjar E, Bussmann RW, Roa C, Sharon D. Medicinal herbs of Southern Ecuador – Hierbas
Medicinales del Sur Ecuatoriano. San Diego: Latino Herbal Press; 2002. p. 340.
Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer International Publishing;
2017. XXVII, 746 p. ISBN 978-3-319-49411-1.
Bussmann RW, Sharon D. Traditional plant use in Loja province, Southern Ecuador. J Ethnobiol
Ethnomed. 2006a;2:44.
healing culture. J Ethnobiol Ethnomed. 2006b;2:47.
Bussmann RW, Sharon D. Plants of longevity – the medicinal flora of Vilcabamba. Plantas de
longevidad – La flora medicinal de Vilcabamba. Honolulu: Arogya; 2007a. ISBN 978-0-
9789962-2-2.
Bussmann RW, Glenn A, Sharon D. Antibacterial activity of medicinal plants of Northern Peru –
can traditional applications provide leads for modern science? Indian J Tradit Med. 2010;9
(4):742–53.
Bussmann RW, Glenn A, Sharon D, Chait G, Díaz D, Pourmand K, Jonat B, Somogy S,
Guardado G, Aguirre C, Meyer K, Rothrock A, Townesmith A. Antibacterial activity of
Northern Peruvian medicinal plants. Ethnobot Res Appl. 2011a;9:67–96.
region, Ethiopia. J Ethnobiol Ethnomed. 2011b;7:28.
Bussmann RW, Paniagua-Zambrana NY, Sikharulidze S, Kikvidze Z, Kikodze D, Jinjikhadze T,
Shanshiashvili T, Chelidze D, Batsatsashvili K, Bakanidze N. Wine, beer, snuff, medicine and
loss of diversity – ethnobotanical travels in the Georgian Caucasus. Ethnobot Res Appl.
2014;12:237–313.
Bussmann RW, Sharon D. Medicinal plants of the Andes and the Amazon – The magic and
medicinal flora of Northern Peru. William L. Brown Center, MBG, St. Louis. 2015. ISBN
978-0-9960231-2-2.
Khutsishvili M, Batsatsashvili K, Hart RE. A comparative ethnobotany of Khevsureti,
Samtskhe-Javakheti, Tusheti, Svaneti, and Racha-Lechkhumi, Republic of Georgia Sakartvelo,
Caucasus. J Ethnobiol Ethnomed 2016a;12:43. https://doi.org/10.1186/s13002-016-0110-2.
Batsatsashvili K, Hart RE. Medicinal and food plants of Svaneti and Lechkhumi, Sakartvelo
(Republic of Georgia), Caucasus. Medicinal and Aromatic Plants 2016b;5:266. https://doi.org/
10.4172/2167-0412.1000266.
Batsatsashvili K, Hart RE. Ethnobotany of Samtskhe-Javakheti, Sakartvelo Republic of Geor-
gia, Caucasus. Indian J of Traditional Knowledge 2017;161:7–24.

Batsatsashvili K, Hart RE. Unequal brothers – Plant and fungal use in Guria and Racha,
Sakartvelo Republic of Georgia, Caucasus. Indian J of Traditional Knowledge 2018;171:7–33.
Kunwar RM, Mahat L, Sharma LN, Shrestha KP, Kominee H, Bussmann RW. Underutilized plant
species in far-west Nepal – a valuable resource being wasted. J Mt Sci. 2012;9:589–600.
Kunwar RM, Mahat L, Acharya RP, Bussmann RW. Medicinal plants, traditional medicine, markets
and management in far-west Nepal. J Ethnobiol Ethnomed. 2013;9:24.
Res Appl. 2013;11:315–39.
Rabearivony ADN, Razafiarison ZL, Raharimalala F, Rakotoarivony F, Randrianarivony T,
Rakotoarivelo N, Randrianasolo A, Kuhlman A, Bussmann RW. Ethnobotanical study of the
plants known by men at Ambalabe, Madagascar. Ethnobot Res Appl. 2015;14:123–38.
2013;9:73.
Teklehaymanot T, Giday M, Mutse M. Quantitative ethnobotany of medicinal plants used by Kara
and Kwego semi-pastoralist people in lower Omo River Valley, Debub Omo Zone, Southern
Nations, Nationalities and Peoples Regional State, Ethiopia. J Ethnopharmacol. 2010;130:76–
84.
10.1186/s13002-019-0285-4.
2008;26:132–53.
Anthocleista zambesiaca Baker
GENTIANACEAE

Grace N. Njoroge
Synonyms
Anthocleista zambesiaca Baker: Anthocleista grandiflora Gilg.
Local Names
Anthocleista zambesiaca: Kikuyu: Mutunguru, Kipsigit: Masomboet, Sagalituet,

Meru: Mutete (Beentje 1994; Gachati 1989)
Botany and Ecology
Anthocleista zambesiaca Baker: Tree 7–27 m high, bark transversely striate,

branchlets unarmed. Leaves sessile or rarely shortly petiolate with petiole up to
2 cm long, very variable in size, lamina usually membranous, sometimes more or
less leathery, oblanceolate to broadly obovate, 20–60( 120) cm long, 8–30( 50)
R. W. Bussmann (*)
G. N. Njoroge

cm wide, rounded or subacute, cuneate at the base, not or inconspicuously auriculate,

margin more or less enrolled and crinkled, lateral nerves 9–14, prominent below,
tertiary nerves conspicuous. Inflorescence 19–40 cm long, 10–25 cm broad, usually
erect and shorter than the subtending leaves, primary peduncles up to 25 cm long,
pedicels fairly stout, 4–10 mm long, bracts leathery, broadly ovate-deltoid, acute,
4–8 mm long. Flowers fragrant. Corolla-tube 2.5–3 cm long, corolla-lobes (10–) 12
( 16), oblong or very narrowly elliptic, 1.3–2.2 cm long, often recurved, buds
usually tapered and definitely broadest below the apex. Anthers 6–7 mm long. Ovary
ovoid to narrowly ovoid, 5–9 mm long, style 2.5–4 cm long, stigma sub-globose,
2–3 mm in diameter, just exserted in the open flower. Outer calyx-lobes orbicular,
rather fleshy or leathery, rugose, 6–10 mm long, rounded to subacute, often rather
spreading and not clasping the corolla-tube, inner lobes subequal to outer ones,
sometimes broader, less leathery and rugose, subacute, rounded or rarely emarginate,
not or scarcely overlapping each other. Fruit erect, more or less ovoid, 3–3.5 cm
long, 2–2.5 cm in diameter, usually beaked or pointed at the apex, wrinkled with the
stalk thickened, and the calyx woody, spreading, the lobes shriveling and becoming
subacute (Bruce and Lewis 1960). An important component of wet Afromontane
forests (Bussmann 2002a, b; Kindt et al. 2007) (Fig. 1).
Anthocleista madagascariensis leaves are used to treat stomach ulcers, diarrhea,

malaria, constipation, abdominal pain, and severe headache in Madagascar, where
the species is often sold in local markets (Randriamiharisoa et al. 2015).
Anthocleista zambesiaca Sometimes used as firewood but smoking a lot. Also as

construction timber. Anthocleista madagascariensis is used for the same purpose in
Madagascar (Rakotoarivelo et al. 2013).
Anthocleista zambesiaca Baker 163
Fig. 1 Anthocleista
zambesiaca (Gentianaceae),
large leaved plant on the right
of the image, lower Mt. Kenya
forests, Kenya. (Photo
R.W. Bussmann)
References
Bruce EA, Lewis J. Flora of tropical East Africa: Loganiaceae. London: Crown Agents for Overseas
Kindt R, Lillesø JPB, van Breugel P. Comparisons between original and current composition of
indigenous tree species around Mount Kenya. Afr J Ecol. 2007;45(4):633–44.
Rakotoarivelo NH, Razanatsima AA, Rakotoarivony F, Kuhlman A, Andriambololonera S,

Randrianasolo A, Bussmann RW. Guide de plantes d’Ambalabe – Vol. 1. St. Louis: William
L. Brown Center, MBG; 2013. ISBN 978-0-9848415-7-8.
Randriamiharisoa MN, Kuhlman AR, Jeannoda V, Rabarison H, Rakotoarivelo N,
Randrianarivony T, Raktoarivony F, Randrianasolo A, Bussmann RW. Medicinal plants sold
in the markets of Antananarivo, Madagascar. J Ethnobiol Ethnomed. 2015;11:60. https://doi.
org/10.1186/s13002-015-0046-y.
Artemisia absinthium L.
Artemisia afra Jacq. ex Willd.
ASTERACEAE

Grace N. Njoroge
Local Names
Artemisia absinthium: English: Common wormwood, Absinthe

Artemisia afra: Kikuyu: Mutathia, Mutasia, Muhato, Kipsigis: Sisimwet, Luo:
Nyumba, Maa: Ol-tikambu, Oldigambu, Marakwet: Sesimwa, Shambaa: Fifi,
Sukuma: Ushimeli (Beentje 1994; Gachati 1989; Kokwaro 2009)
Botany and Ecology
Artemisia absinthium L.: Perennial. Whole plant grayish from short approximate
hairs, sericeous-tomentose. Root vertical, thick, perennial. Stem 60–100 cm high,
herbaceous, erect, with short branches, leafy, sometimes basally with short nonflow-
ering branches bearing long-petiolate leaves. Leaves with 6–9 cm long and 3–7 cm
wide lamina broadly ovate, almost thrice pinnately dissected, terminal lobe lanceo-
late, short-acuminate; middle cauline leaves short petiolate, lacking lobes at base of
petiole, twice pinnately dissected; upper leaves subsessile, simple pinnate or twice
R. W. Bussmann (*)
G. N. Njoroge

ternate; bracteal leaves ternate or simple, undivided, narrowly lanceolate; lobes of all
leaves linear-oblong, scarcely acuminate, undivided or with few teeth, usually
3–20 mm long and 1–4 mm wide. Capitula globose, 2.5–3.5 mm in diameter,
drooping or spreading into narrowly paniculate inflorescence. Involucral bracts
oblong-ovate, outer almost as long as inner bracts, hairy on spine, with wide scarious
margin. Receptacle convex, hairy. Peripheral florets pistillate, usually 25, their
corollas filiform-tubular; stigma lobes exserted from tube, linear, arcuate; disk florets
numerous (usually 60), their corollas conical, glabrous, sometimes weakly hairy;
anthers linear, apical appendages of anthers obtusely angular, with round apex, basal
appendages short, obtuse; stigma lobes broadly linear, after flowering somewhat
curved. Achenes about 1 mm long, oblong-cuneate, rather flat, finely sulcate, with a
round, slightly convex at apex. Flowering July to August. Ural, Caucasus, Altai,
Middle Asia, in meadows, steppes, on river banks, forest margins, fields, rocky
areas, in crops and fields, in gardens, along irrigation ditches, around settlements as
weed, on recent fallow lands, near roads, dwellings, in kitchen gardens and gardens,
less often in crop fields, meadows, pastures and forest edge; in forest, forest-steppe
and steppe zones, as well as in mountains. (Shishkin and Boborov 1961) (Figs. 1, 2,
and 3).
Fig. 1 Artemisia absinthium

(Asteraceae), Capadokia,
Turkey. (Photo
R.W. Bussmann and
Artemisia absinthium L. . . . 167
Fig. 2 Artemisia absinthium (Asteraceae), Kartli, Georgia. (Photo R.W. Bussmann and
Fig. 3 Artemisia absinthium

(Asteraceae), Svaneti,
Georgia. (Photo
R.W. Bussmann and
Artemisia afra Jacq. ex Willd.: Shrub, up to 2 m high, very aromatic. Leaves

alternate, petiolate, blade narrowly ovate in outline, bipinnatipartite, ultimate seg-
ments linear, entire or toothed, upper surface glabrous, lower surface canescent.
Heads disciform, subglobose, about 3 mm in diameter, many in terminal, racemose
panicles. Involucral bracts in a few rows, imbricate, canescent. Receptacle epaleate.
Flowers: outer female florets tubular, fertile, disc florets bisexual or functionally male,
yellow, February–May Fruit with cypsela narrowly obovoid, smooth or ribbed.
Pappus absent. Forest margins and along streams. Dry Grassland, Mesic Grassland,
Dwarf Karroid Shrubland. Forest margins, grassland and often along streams, in
sandy or rocky soil. Stream-sides and damp slopes. Common in the mountain forest –
subalpine zone transition zone, and in disturbed habitats at forest edges (Bussmann
2006) (Figs. 4, 5, 6, and 7).
Fig. 4 Artemisia afra

(Asteraceae), Mt. Kenya
National Park, Kenya. (Photo
R.W. Bussmann)

(Asteraceae), Bale Mountains
National Park border zone,
Oromyia, Ethiopia. (Photo
R.W. Bussmann)
Phytochemistry
Artemisia absinthium: Cyclitols (quiberachite), organic acids, essential oils

(cadinene, guaiazulene, ketopelenolide a, ketopelenolide b, hydroxyketopelenolide,
caryophyllene, selenene, a-pinen, nerol, tujone, tujol, p-cymene, camphene, cineole,
phellandrene, geraniol, elemol, a-chymachalene, a-cadine, limonene, eucalptol,
y-terpinene, linalool, isoborneol, camphor, a-bizabolol, hamazulene, methylha-
mazulene, ethylhamazulene, tuyen, sabinene, a-terpinene, ocimene, terpinolene,
n-hexanol, nonanal, farnesene, lavandulol, a-curcumine, germacrene D, sabinol),
sesquiterpenoids (absintine, anabsintine, artamarine, artamaridine, artamaridinine,
artamarinin, artabsine, artbine, arabsine, anabsine, artemoline, absintolide, iso-
absintine, ketopelenolide, artbine, arabine, arlatine, arenolide, parsirIn B, parsherIn
C), lignans (lyrioresinol A, lyrioresinol C, sezartemine, episazeartemIn A,
episepartemIn B, diazezartemine, sesamine, epieudesmine, fargesine, yangambin,
epiangambine, diayangambine, askhantine, epiaschantine), flavonoids (artemetine,
quercetine, kaempferol, isoramnetine, apigenin), fatty acids (lauric, myristic,

National Park border zone,
Oromyia, Ethiopia. (Photo
R.W. Bussmann)
palmitic, stearic, oleic, linoleic, arachidic), carbohydrates (inuline), polyacetylene

compounds, phenolcarboxylic acids, vitamins (C), steriods, coumnarins
(scopoletine, umbelliferon), fatty acids (lauric, myristic, palmitic, stearic, oleic,
linoleic, arachidic), tannins (Sokolov 1993).
Artemisia afra: Heated plant material is fermented and given to children with sore
throat and for fever. Also for indigestion. Boiled roots are used as anthelminthic. The
leaf juice serves as emetic. Ash from the whole plant is used as expectorant and for
abdominal pain. Extract of boiled leaves drunk for colds, flu, headache, and fever. A
steam bath is used for flu and colds. Crushed leaves can be applied for congestion,
and leaf juice as ear drops (Kokwaro 2009). A very important medicinal species in
Ethiopia, and often sold in local markets and in the medicinal plant trade (Bekalo
et al. 2009). The species is also used to treat fever, and burned for colds, when the
smoke is inhaled (Yineger et al. 2008). The antibacterial activity of Artemisia afra
and Artemisia abyssinica has been shown (Geyid et al. 2005). The species is also
used to treat epilepsy (Yineger et al. 2007). Artemisia absinthium is often planted in
Ethiopia as medicine (Mekonnen et al. 2015), and, e.g., used for wounds (Yineger
et al. 2007).
Fig. 7 Artemisia
sp. (Asteraceae), Bale
Mountains National Park
border zone, Oromyia,
Ethiopia (Photo
R.W. Bussmann)
Artemisia abyssinica has been shown to be acaricidal (Wanzala 2017).
References
plants used by local people in the lowlands of Konta special Woreda, southern nations,
doi.org/10.1186/1746-4269-5-26.
2006;11(1):41–66.

Shishkin BK, Boborov EG. Flora of the USSR, Volume 26: Compositae Giseke (altern. Asteraceae
Dumort); Leningrad:Akademia Nauk; 1961 (English 1995). 1072 pages.
Sokolov PD, editor. Plant resources of the USSR: flowering plants, their chemical composition, use;
volume 7. Family Asteraceae (Compositae). Leningrad: Akademia Nauk; 1993.
352 p. (in Russian).
livestock industry with focus on acari ticks. Evid-Based Complement Alternat Med.
2017:8647919. 33 pages. https://doi.org/10.1155/2017/8647919.
2008;26:132–53.
Asparagus africanus Lam.
Asparagus falcatus L.
ASPARAGACEAE

Grace N. Njoroge
Synonyms
Asparagus africanus Lam.: Protasparagus africanus (Lam.) Oberm.

Asparagus falcatus L.: Protasparagus falcatus (L.) Oberm.
Local Names
Asparagus africanus: Gonja: Chichilazyuka, Maasai: Embare Baba, Hehe:

Kapalang’anga, Taita: Lwafumbo, Giriama: Murikano (Gachati 1989; Kokwaro
2009), English: Asparagus
Asparagus falcatus: Kikuyu: Rurura, Sukuma: Moungwi, Shambaa, Mungwe,
Digo: Mwinka Ngulu (Gachati 1989; Kokwaro 2009)
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Asparagus africanus Lam.: Erect or scandent shrub, 0.5–3(5) m high. Stems many,
pubescent or glabrous, smooth, green with spreading branches and branchlets.
Spines straight or reflexed, sharp, 5–10 mm long. Cladodes 3–12 per fascicle,
filiform, slightly curved, apiculate, unequally long, 5–15(20) mm long. Flowers 2–
8 in each cladode fascicle, pedicels 5–10 mm long, articulated in lower half. Tepals
narrowly obovate, 2.5–4 mm long, whitish. Stamens with small yellow anthers.
Ovary with 4 ovules per locule, obovoid, style and stigmas very short. Berry 5–6 mm
in diameter, orange to red. A common component in the undergrowth of
Afromontane forests (Bussmann 2002) (Fig. 1).
Asparagus falcatus L.: Climbing shrub to 4 m or more. Branches glabrous. Stems
smooth, terete to angled, usually pale. Spines 1–17 mm long, recurved, also on
terminal branches. Cladodes fasciculate, 3–6 together, straight or falcate, smooth,
with a distinct midvein, (15)25–120 2–5 mm. Racemes simple, 1–3, axillary, 15–
55 mm long, present on main stems and branches, bracts ovate, acuminate, mem-
branous, pedicels 3–7 mm long, articulated below middle or above. Tepals white,
broadly elliptic to obovate, 3–3.5 mm long. Stamens with yellow anthers. Ovary
with 6 ovules in each locule, style short, about 0.5 mm long including stigma.
Berry globose or solitary or 2-lobed, 7–10 mm in diameter, red, 1–2 seeded.
Fig. 1 Asparagus africanus

(Asparagaceae), Mt. Kenya
R:W. Bussmann)
Asparagus africanus Lam. . . . 175
Fig. 2 Asparagus falcatus

R:W. Bussmann)
Widespread in the warmer parts of southern Africa, tropical East Africa to India and
Sri Lanka. Coastal forest, sand dunes, miombo woodland, along forest margins,
rivers and streams, and among rocky outcrops, 0–1800 m. (Demissew 2006). A
common component in the undergrowth of Afromontane forests (Bussmann 2002)
(Figs. 2, 3, and 4).
Asparagus africanus: Roots chewed as remedy for sore throat and cough. The root
infusion is used to treat gonorrhea and other venereal diseases. Leaves are applied to
wounds, and a leaf decoction is used to cure bubonic plague (Taita). The root
infusion is used to ease childbirth. A decoction of the whole plant is drunk for
mental problems (Kokwaro 2009). Juice is put on pimples (Bussmann et al. 2011;
Lulekal et al. 2008). Used to treat diarrhea and involuntary weight-loss
(Teklehaymanot et al. 2007), as well as skin lesions (Teklehaymanot 2009).
Fig. 3 Asparagus falcatus (Asparagaceae), Bale Moutains National Park, Odo-Bulu, Ethiopia.
(Photo R:W. Bussmann)
Fig. 4 Asparagus setaceus

(Asparagaceae), Bale
Moutains National Park,
Odo-Bulu, Ethiopia. (Photo
R:W. Bussmann)
Asparagus falcatus: Root tubers used to treat gonorrhea and to cure hernias in
children. Also drunk as anthelminthic (Kokwaro 2009). Juice is put on pimples
(Bussmann et al. 2011).
Asparagus setaceus juice is used to treat herpes (Giday et al. 2009). The leaves are
chewed for cough (Njoroge and Bussmann 2006), and the juice is applied to boils
(Njoroge and Bussmann 2007). Used also for back pain in women, postpartum pain,
and to treat sexually transmitted diseases (Njoroge and Bussmann 2009), as well as
abscesses, diarrhea, toothache, wounds, and sores. Used to treat urinary problems in
Ethiopia (Wondimu et al. 2007), plant powder for dermal issues (Yineger et al. 2008).
Asparagus aethiopicus is used as anti-hypertensive (Desta 1995).
Asparagus racemosa has medicinal applications in Ethiopia (Bekalo et al. 2009).
Used as galactagogue (Kunwar et al. 2018). Asparagus root is tonic, aphrodisiac,
diuretic, carminative, appetizer, and antispasmodic. It is useful in mental disorders,
dyspepsia, and diarrhea. Tender shoots are tonic and useful in dysentery and rheuma-
tism. Also used in tuberculosis, measles, diarrhea, epilepsy, and liver problems and
considered as a galactagogue, aphrodisiac, diuretic, antispasmodic, and nerve tonic. It is
also considered to prevent ageing, impart immunity, improve mental functions, and add
vigor and vitality to the body. Root extract increases mammary gland in postpartum
period acting as a galactagogue. Dysmenorrhoea and premenstrual syndrome were
found to be symptom free after usage of root extract. The plant extract is
cardioprotective, antitumor, antifungal, anti-ulcer, immunostimulatory, and diuretic.
The crude saponins from the shoots are useful in dyspepsia, and extracts of plant
rhizomes reduce blood sugar levels (Kunwar and Bussmann 2009; Kunwar et al.
2009a, 2013; Raj et al. 2018).
The roots of all Asparagus species used as diuretic (Fayvush et al. 2017).
Asparagus adscendens also serves as galactagogue (Singh et al. 2017). Asparagus
filicinus is used for diabetes and diarrhea (Bhat et al. 2013). Asparagus simulans is
used in Madagascar to treat epilepsy and stomach pain (Razafindraibe et al. 2013).
Local Food Uses
Asparagus africanus/Asparagus falcatus: The large root stores lots of water that
can be collected (Bussmann et al. 2006).
Asparagus africanus/Asparagus falcatus: Eaten by goats and cattle and threaded

for calabashes (Bussmann 2006; Bussmann et al. 2011; Luizza et al. 2013). Aspar-
agus africanus: Used to wash away the spirit of a person who died (Bussmann et al.
2006). Used to make brooms (Bussmann et al. 2011). Also used to treat horses
(Giday et al. 2003). The root extract is used as insect repellent (Giday et al. 2007),
and to treat rabies (Yineger and Yewhalaw 2007). Asparagus racemosus roots used
for fermentation of local beer (Kunwar et al. 2009b), and ethnoveterinary purposes
(Reang et al. 2016).
References
doi.org/10.1186/1746-4269-5-26.
Bhat J, Kumar M, Bussmann RW. Ecological status and traditional knowledge of medicinal plants
in Kedarnath Wildlife Sanctuary of Garhwal Himalaya, India. J Ethnobiol Ethnomed. 2013;9:1.
http://www.ethnobiomed.com/content/9/1/1.
Bussmann RW, Batsatsashvili K, Kikvidze Z. Asparagus persicus Baker, Asparagus sp. In:
Batsatsashvili K, Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions Central
Asia and Altai. Cham: Springer International Publishing; 2020; https://doi.org/10.1007/978-3-
319-77087-1_24-1.
Demissew S. Flora of tropical East Africa: Asparagaceae. Kew: Royal Botanic Gardens; 2006.
Desta B. Ethiopian traditional herbal drugs. Part I: studies on the toxicity and therapeutic activity of
local taenicidal medications. J Ethnopharmacol. 1995;45:27–33.
Fayvush G, Aleksanyan A, Mehdiyeva N, Alizade V, Batsatsashvili K, Kikvidze Z, Khutsishvili M,
Maisaia I, Sikharulidze S, Tchelidze D, Paniagua Zambrana NY, Bussmann RW. Asparagus
officinalis L., Asparagus verticillatus L. In: Bussmann RW, editor. Ethnobotany of the Cauca-
sus. Cham: Springer International Publishing; 2017.
and Amhara proples in northwest Ethipopia. Joruna. J Ethnopharmacol. 2007;110(3):516–25.
https://doi.org/10.1016/j.jep.2006.10.011.
Giday M, Asfaw Z, Woldu Z, Teklehaymanot T. Medicinal plant knowledge of the Bench ethnic
group of Ethiopia: an ethnobotanical investigation. J Ethnobiol Ethnomed. 2009;5:34. https://
doi.org/10.1186/1746-4269-5-34.
Biodiversität, Naturausstattung im Himalaya, Bd. III. Universität Ehrfurt, Erfurt; 2009.
p. 475–89.
Kunwar RM, Evans A, Mainali J, Ansari AS, Rimal B, Bussmann RW. Change in forest and
vegetation cover influencing distribution and uses of plants in the Kailash Sacred Landscape,
Nepal. Environ Develop Sustain. 2018. https://doi.org/10.1007/s10668-018-0254-4.
macology of medicinal plants in Far-west Nepal. Ethnobot Res Appl. 2009a;7:5–28.
Kunwar RM, Chowdhary L, Bussmann RW. Diversity, utilization and management of medicinal
plants in Baitadi and Darchula districts, farwest Nepal. Initiation. 2009b;21:157–64.
Kunwar RM, Fadiman M, Cameron M, Bussmann RW, Thapa-Magar KB, Rimal B, Sapkota
P. Cross-cultural comparison of plant use knowledge in Baitadi and Darchula districts, Nepal
Himalaya. J Ethnobiol Ethnomed. 2018;14:40. https://doi.org/10.1186/s13002-018-0242-7.
Luizza MW, Young H, Kuroiwa M, Evangelista P, Worede A, Bussmann RW, Weimer A. Local
Res Appl. 2013;11:315–39.
10.1186/1746-4269-4-10.
Central Kenya. J Ethnobiol Ethnomed. 2006;2:54.
Njoroge GN, Bussmann RW. Ethnotherapeutic management of skin diseases among the Kikuyus of
Central Kenya. J Ethnopharmacol. 2007;111:303–7.
Njoroge GN, Bussmann RW. Ethnotherapeutic management of Sexually Transmitted Diseases
STDs and reproductive health conditions in Central Province, Kenya. Indian J Tradit Knowl.
2009;8(2):262–9.
018-0208-9.
2013;9:73.
inal plants by traditional herbal healers in Reang tribe of south district Tripura, India. Med
Aromat Plants. 2016:5–2.
Singh A, Nautiyal M, Kunwar RM, Bussmann RW. Ethnomedicinal plants used by local inhabitants
of Jakholi Block, Rudraprayag district, Western Himalaya, India. J Etnobiol Ethnomed.
2017;13:49. https://doi.org/10.1186/s13002-017-0178-3.
Yineger H, Yewhalaw D. Traditional medicinal plant knowledge and use by local healers in Sekoru
District, Jimma Zone, Southwestern Ethiopia. J Ethnobiol Ethnomed. 2007;3:24.
2008;26:132–53.
Asystasia gangetica (L.) T. Anderson
ACANTHACEAE

Grace N. Njoroge
Synonyms
Asystasia gangetica (L.) T. Anderson: Asystasia bojeriana Nees, Asystasia

comorensis Nees, Asystasia comorensis var. humilis Nees, Asystasia
coromandeliana Burkill & C.B. Clarke, Asystasia coromandeliana Nees, Justicia
gangetica L., Ruellia zeylanica Roxb.
Local Names
Asystasia gangetica: Digo: Mforfor (Kokwaro 2009), English: Tropical primrose,

Chinese violet, French: Herbe le rail, mange-tout, herbe pistache, pistache marron,
Portuguese: Asistasía branca, Swahili: Fuchwe, mtikini, kichwamangwo
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Asystasia gangetica (L.) T. Anderson: Perennial herb, with usually ascending,

branched, quadrangular stem up to 2 m long, often rooting at the lower nodes.
Leaves opposite, simple, stipules absent, petiole 0.5–6 cm long, blade ovate to
lanceolate, 3–8( 13) cm 1.5–4.5( 7) cm, base cuneate to cordate, apex acumi-
nate or acute, margin entire, glabrous to sparsely pubescent, with 4–6 lateral veins at
each side of the midrib, provided with cystoliths. Inflorescence a terminal raceme up
to 25 cm long, with flowers directed to one side. Flowers bisexual, slightly zygo-
morphic, 5-merous, pedicel up to 3 mm long, calyx with lanceolate lobes 4–10 mm
long, corolla funnel-shaped, up to about 2.5( 4) cm long, usually white with
purplish spots inside lower lobe, with rounded lobes c. 1 cm wide, lower lobe
slightly longer, stamens 4, 2 shorter and 2 longer, ovary superior, densely pubescent,
2-celled, style up to 1.5( 2) cm long, stigma with 2 short lobes. Fruit a club-shaped
capsule 2–3 cm long, pubescent and glandular, usually 4-seeded. Seeds ovoid,
flattened, 4–5 mm long, grey to brown, with crenate margins, tuberculate, supported
by retinacula. Native in tropical Africa, Arabia, and tropical Asia, but has been
introduced in many other tropical regions, where it often naturalized. It occurs
throughout tropical Africa (Vollesen 2008) and is often found in Croton and Vepris
dominated Afromontane forests (Bussmann 2002).
Asystasia gangetica: Africa an infusion of the plant is used to ease pain during
childbirth, and the sap is applied to sores, wounds, piles, and in embrocations to treat
stiff neck and enlarged spleen in children. Powdered roots are considered analgesic
and used in treating stomach-ache and snakebites. A leaf decoction is used as
analgesic and to treat epilepsy and urethral discharge. In Nigeria, the leaves are
used to treat asthma. In India, the sap is applied to swellings, it is also used as a
vermifuge and to treat rheumatism. In the Moluccas (Indonesia) the juice, together
with lime and onion juice, is recommended for dry coughs with an irritated throat
and discomfort in the chest. Leaves crushed and boiled in water as anthelminthic
(Kokwaro 2009). The leaves of Asystasia schimperi are used by the Nandi to treat
cough and skin problems (Jeruto et al. 2008). In Ethiopia Asystasia excellens leaves
are boiled and applied to wounds to stop bleeding (Yineger et al. 2008).
Local Food Uses
Asystasia gangetica: Locally used as a potherb and leafy vegetable, mainly in times
of scarcity. In Kenya and Uganda, it is locally a popular vegetable, mixed with beans
and groundnut or sesame paste. It is also often prepared in a mix with other leafy
vegetables.
Asystasia gangetica (L.) T. Anderson 183
Asystasia gangetica: In Tanzania, plants are pounded with water to make a wash
against fleas for young animals. Occasionally planted as an ornamental. Because of
its ability to grow under shade and its nutritive value, used as a forage for cattle,
goats and sheep in South-East Asia, it is either grazed or cut for stall feeding.
Excessive consumption by sheep can result in bloat.
References
Vollesen K. Flora of tropical East Africa: Acanthaceae (Part 1). Kew: Royal Botanic Gardens; 2008.
286 pp. ISBN 978-1-84246-217-1 [Fl. Trop. E. Africa, Acanth. (Part 1)].
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plant Res. 2008;2
(6):132–53.
Azadirachta indica A. Juss.
MELIACEAE

Grace N. Njoroge
Synonyms
Azadirachta indica A. Juss.: Melia azadirachta L., Melia indica (A. Juss.) Brandis
Local Names
Azadirachta indica: Swahili: Mwarumbaini, English: Neem (Kokwaro 2009).
Botany and Ecology
Azadirachta indica A. Juss.: Evergreen trees with simple trichomes, up to 15 m

high, short and robust trunk, brown, wrinkled and fissured bark, polygamous plants.
Leaves up to 40 cm long, with 4–9 pairs of opposite leaflets, leaflets ovate-lanceolate
to lanceolate, strongly falcate, up to 9 cm long and 3 cm wide, apex long
R. W. Bussmann (*)
G. N. Njoroge

accumulated, base highly asymmetrical, margin thickly serrated, glabrous. Inflores-

cence an axillary panicle, up to 35 cm long, calyx 5-lobed to below midway, petals
5, free, imbricated, white, cylindrical staminal tube, slightly expanded at the apex,
terminated by 10 appendages, these rounded or truncated, emarginated or bi-lobed,
and sometimes joined and forming a flight, annular disc, fused to the base of the
ovary, 3-locular ovary, each locule with 2 collateral ovules, sparsely expanded style,
ending in 3 acute and partially fused stigmatic wolves, cream color. Ellipsoid drupe,
1.5–1.8 cm long, yellow, seeds 1–2 with cartilaginous endocarp (Styles and White
1991). Commonly planted in agroforestry (Onyango et al. 2004).
Azarirachta indica: Oil used for skin diseases, leprosy, and eczema. The leaf
decoction is drunk for malaria. Branches used as toothbrush. Bark and leaves are
applied externally for wounds, rheumatism, and to stimulate liver function (Giday
et al. 2007; Kokwaro 2009). The leaves are also used to reduce fever (Kunwar et al.
2010). Part of incense mixtures is used for tuberculosis, smallpox, measles, skin
problems, rheumatism, cardiac problems, and as antifungal (Mohagheghzadeh and
Faridi 2006). Used as antimalarial (Muthee et al. 2011; Muthaura et al. 2007;
Njoroge and Bussmann 2006a; Tabuti 2008), and for ear, nose, and throat diseases
(Njoroge and Bussmann 2006b), as well as stomach problems (Muthee et al. 2011).
The species is also used as anthelminthic (Tadesse et al. 2009). In India used to
treat antipyretic, antiseptic, blood purifier, boils, cholera, diabetes, diarrhea, dys-
entery, dyspepsia, heart complaints, inflammation of gums, jaundice, leprosy, liver,
complaints, malaria, measles, piles, rheumatism, scabies, skin disease, small pox,
snake bite, spleen complaints, syphilis, tonic, tuberculosis, tumors, ulcer, urine
complaints, venereal diseases, and wounds (Verma et al. 2007; Raj et al. 2018).
Fresh leaves are consumed during stomach problem, cough, cold, pimples, fever,
smallpox, and chicken pox (Debbarma et al. 2017). In Madagascar, the leaf
decoction is used to treat colic in infants (Randrianarivony et al. 2016). The leaves
are sold in local markets as remedy for asthenia, diabetes, albumin, rheumatism,
pelvic pain, boils, hepatitis, kidney stones, burns, constipation, and high choles-
terol (Randriamiharisoa et al. 2015). Sometimes used for diabetes (Ullah et al.
2019), and in Pakistan also for malaria, fever, as blood purifier, and for headache
(Umair et al. 2019).
Azarirachta indica: Leaf decoction used in poultry for diarrhea, scaly legs, pox
lesions (Kokwaro 2009), and other veterinary purposes (Njoroge and Bussmann
2006c). As insecticide against tick infestations, and tsetse flies (Wanzala 2017).
Azadirachta indica A. Juss. 187
References
and Amhara proples in northwest Ethipopia. J Ethnopharmacol. 2007;110(3):516–25.
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in Far-west Nepal: a
pharmacological appraisal. J Ethnobiol Ethnomed. 2010;6:35.
Muthaura CN, Rukunga GM, Chhabra SC, Mungai GM, Njagi ENM. Traditional antimalarial
phytotherapy remedies used by the Kwale community of the Kenyan Coast. J Ethnopharmacol.
2007;14:377–86.
Njoroge GN, Bussmann RW. Phytotherapeutic management of diversity and utilization of antima-
larial ethnophytotherapeutic remedies among the Kikuyus Central Kenya. J Ethnobiol
Njoroge GN, Bussmann RW. Traditional management of Ear, Nose and Throat ENT diseases in
Central Kenya. J Ethnobiol Ethnomed. 2006b;2:54.
of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006c;108:332–9.
Onyango JC, Nyunja RAO, Bussmann RW. Conservation of biodiversity in the East African
Tropical Forest. Lyonia. 2004;72:151–7.
RW. Indigenous uses of ethno-medicinal plants among forest dependent communities of Northern
Bengal, India. J Ethnobiol Ethnomed. 2018;141:8. https://doi.org/10.1186/s13002-018-0208-9.
org/10.1186/s13002-015-0046-y.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016. ISBN 978-0-9960231-4-6.
Styles BT, White F. Flora of Tropical East Africa: Meliaceae. Rotterdam: Balkema; 1991. 66 pp.
ISBN 90-6191-356-X [Fl. Trop. E. Africa, Mel.].
Tadesse M, Eguale T, Giday M, Mussa A. Ovicidal and larvicidal activity of crude extracts of
Maesa lanceolata, and Pleactranthus punctatus against Haemonchus contortus.
Ullah M, Mehmood S, Ali M, Bussmann RW, Aldosari A, Ali Khan R, Ullah R, Hussain W,
Rahman Shah MA. An ethnopharmacological study of plants used for treatment of diabetes in
the Southern and Tribal regions of Khyber Pakhtunkhwa province, Pakistan. Ethnobot Res
Appl. 2019;188. https://doi.org/10.32859/era.18.8.1-20.
10.1186/s13002-019-0285-4.
livestock industry with focus on Acari Ticks. Evid Based Complement Altern Med. 2017;
Article ID 8647919, 33 p. https://doi.org/10.1155/2017/8647919.
Balanites aegyptiaca (L.) Delile
Balanites pedicellaris Mildbr. & Schltr.
Balanites rotundifolia (Tiegh.) Blatt.
ZYGOPHYLLACEAE

Grace N. Njoroge
Synonyms
Balanites aegyptiaca (L.) Delile: Agalid aegyptiaca (L.) Kuntze, Ximenia

aegyptiaca L.
Balanites pedicellaris Mildbr. & Schltr.: Balanites australis Bremek.
Balanites rotundifolia (Tiegh.) Blatt.: Synonyms: Agalid rotundifolia Tiegh.
Local Names
Balanites aegyptiaca: Swahili: Mkonga, Mnyira, Mchunju, Mguguni, Borana:

Baddan, Gabbra: Baddano, Kamba: Kilului, Mulului, Kipsigis: Ngoswet, Luo:
Othoo, Maa: Olngoswa, Marakwet: Ngoswa, Tunyunwa, Pokot: Tuyunwo,
Samburu: Lowvai, Tugen: Ngonswo, Turkana: Eroronyit, Luganda: Musongole
(Beentje 1994; Kokwaro 2009), English: Torchwood, Green thorn.
Balanites pedicellaris: Ilelewa: Mubadana, Oromo: Baddan, Pokot: Lomion,
Loma, Marakwet: Lomion, Taita: Kikowa, Turkana: Elamach (Beentje 1994;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Balanites rotundifolius: Borana/Oromo: Baddan, Ilelewa: Mubadada, Pokot/

Marakwet: Lomion, Pokomo: Daddan, Rendile: Kulum, Samburu: Sarei,
Somali: Kullan, Tugen: Tirikiwa, Turkana: Ebei (Beentje 1994; Kokwaro 2009).
Botany and Ecology
Balanites aegyptiaca (L.) Delile: Small evergreen tree (rarely a shrub) up to 15 m

tall, spiny, with a flat or rounded crown, stem with a reticulate dark brown or grey
(rarely green) bark, branches green or greyish, stiff and brittle, always armed with
stout simple green or yellowish spines. Leaves petiolate, leaflets shortly petiolulate,
lamina 2.5–6 1.5–4 cm, slightly asymmetric, elliptic to elliptic-obovate, apex
subacute to obtuse, sometimes slightly refuse, base cuneate or rarely rounded,
coriaceous, puberulous when young, later glabrescent or sometimes remaining
puberulous on the lower surface, secondary nerves 4–6 pairs, prominent beneath,
petiole (0.5)0.8–2 cm long, puberulous or glabrescent, canaliculate. Flowers in
usually few-flowered sessile or shortly pedunculate fascicles, pedicels up to
1.5 cm long, densely greyish-pubescent. Flowers 1.4 cm in diameter. Sepals
5, 2.7 2.7–3.25 mm, ovate or ovate-lanceolate, coriaceous, caducous, densely
pubescent outside, with long silky whitish hairs inside. Petals 7.2–9.5 2–2.4 (2.8)
mm, narrowly elliptic or elliptic-oblong, rarely lanceolate-oblong, glabrous on both
surfaces. Stamens with the anthers 1.7–2.2 mm long, ovate or ovate-oblong, gla-
brous, filaments 3.175 mm long. Ovary densely covered with long silky hairs. Drupe
yellowish or green, up to 5 2.5 cm, usually subcylindric, more rarely narrowly
ellipsoid or subobclavate, finely puberulous, sometimes glabrescent. Occurs from
south-eastern Kenya south to Swaziland and the province of KwaZulu-Natal in
South Africa, with its center of diversity in Mozambique (El Hadidi 1985) (Fig. 1).
Balanites pedicellaris Mildbr. & Schltr.: Much-branched shrub or small tree with
pendulous branches, up to 6 m or taller, variable in shape and size. Branches
yellowish or greyish-green or grey, rather stiff, usually spiny, younger parts densely
puberulous, glabrous when older, spines rather stout, simple. Leaves shortly petio-
late, leaflets sessile, 1–3 (4) 0.5–2.3 (3) cm, obovate, broadly obovate, obcuneate
or rarely obovate-elliptic, apex rounded, rarely subtruncate, entire or slightly retuse,
base always narrowly cuneate, sub-succulent, very shortly and densely puberulous to
sub-tomentose when young, later glabrescent, secondary nerves 3–5 pairs, not very
prominent beneath, petiole 1.5–4 (6) mm long, rather stout, densely pubescent at
first, later glabrescent. Flowers c. 1.4 cm in diameter, in few- to many-flowered
sessile or rarely shortly pedunculate fascicles, pedicels up to 2 cm long, fairly stout,
densely puberulous, very rarely glabrous. Sepals 6.5–7.5 3.2–5 mm, ovate to
ovate-lanceolate, coriaceous, densely tomentose outside, with long whitish silky
hairs inside, acute, caducous. Petals 6.7–9 (2.5) 2.8–3.4 mm, narrowly elliptic
with an acute apex, glabrous on both surfaces. Stamens with oblong anthers 2–
2.8 mm long, filaments 3.75 mm long. Ovary densely covered with whitish silky
hairs. Drupe orange, subglobose or broadly ellipsoid, usually somewhat flattened on
Balanites aegyptiaca (L.) Delile . . . 191
Fig. 1 Balanites aegyptiaca (Zygophyllaceae), Nairobi National Park, Kenya. (Photo

R.W. Bussmann)
both ends, 1.2–2.5 1.5–2 cm, finely puberulous when young, later usually
glabrescent (El Hadidi 1985).
Balanites rotundifolia (Tiegh.) Blatt.: A spiny evergreen shrub or small tree up to 6
(–8) m high (frequently much shorter), with a low bushy habit, densely branched, or
with a trunk up to 40 cm diameter, bark grey-brown, strongly fissured, sap gummy,
branchlets grey-green or yellowish-brown and, like the young spines, glabrous or
puberulous to pubescent at first, glabrescent. Primary spines borne on the parent axis
at a varying angle, (1–)2–5(–10) cm above the axil, 2–11 cm long, 2–5 mm diameter
at the base, subulate, smooth or sometimes shallowly grooved, green or yellowish-
green, spinuliferous with some branch-spines, spinules 0.5–2 cm long grading to
branch-spines. Leaves on the stems and spines, sessile or with a petiole up to 4 mm
long, stipules 1–2(–3) mm long, triangular, puberulous, often persistent, leaflets
sessile or subsessile, orbicular to broadly obovate or obovate-elliptic, 0.7–6.5 cm
long, 0.8–4.8 cm wide, coriaceous, frequently concave and undulate, apex rounded
or sometimes emarginate to truncate and abruptly apiculate, base rounded or broadly
cuneate, glabrous or puberulous to pubescent, eventually glabrescent, rarely
setulose, foliole linear, 0.7–4 mm long, sometimes caducous. Inflorescence on the
stems and spines, a few to 12-flowered fascicle or sometimes clustered on a short
peduncle up to 2.8 cm long, axillary or rarely terminal on a leafy shoot, pedicels 0.2–
1.6 cm long. Flowers 4-merous, sepals ovate, 3–5.5 mm long, 3 mm wide, acute to
acuminate, sparsely pubescent outside, the glabrous margin narrow, petals yellowish
green or olive-green, obovate-elliptic, 3–6.5 mm long, 1.8–2.5 mm wide, acute or
obtuse and sometimes irregular at the apex, narrowing to the base, glabrous within,
stamens 8, spreading-erect, anthers 1.5–2 mm long, 0.5–1 mm wide, ovary 1–
1.5 mm high, glabrous or densely pubescent to pilose, style 1–2 mm long. Fruit,
not elongating in early development, swelling at first proximally, with slower
expansion of the glabrous style-base, eventually ripening orange-yellow, ovoid to
broadly ellipsoid when mature, 2.5–3 cm long, 1.8–2.5 cm diameter, rounded at both
ends, thin, hard, brittle and smooth on the outside, fibrous within enclosing a dense
pale layer, the endocarp hard and oily, the seed becoming free inside the fruit
(El Hadidi 1985).
Balanites aegyptiaca: The roots and bark are widely used in purgative medicines.
The roots are used to treat malaria (Beentje 1994; Kokwaro 2009). Root decoction
also used for edema, abdominal pains, as purgative and anthelminthic. Molten gum
mixed in porridge is used for pneumonia and chest problems (Kokwaro 2009). The
gum is used to treat skin problems (Wondimu et al. 2007). Root and bark used to
treaty stomach problems and cowpox (Muthee et al. 2011).
Balanites pedicellaris: Root infusion used as emetic (Beentje 1994). Pounded roots
in water used to treat diarrhea and fever (Kokwaro 2009).
Balanites rotundifolia: Leaves used to treat wounds (Beentje 1994). Boiled roots in
water used to treat abdominal pain and fever. Leaf paste applied to wounds
(Kokwaro 2009). The roots are boiled, and the decoction drunk for diarrhea and to
stop weight loss and applied to body rushes in Ethiopia (Teklehaymanot et al. 2010).
Local Food Uses
Balanites aegyptiaca: The fruits and seed oil are edible, the oil is used in cooking,
and as a massage oil (Beentje 1994). The leaves and fruits serve as famine foods
(Balemie and Kebebew 2006).
Balanites pedicellaris: Fruit edible (Beentje 1994).
Balanites rotundifolia: Fruit edible (Beentje 1994).
Balanites aegyptiaca: Oil pressed from the seed kernel is used in Limpopo Province
of South Africa as a dressing for hides and skins. In some regions the oil or seeds are
burnt as torches, hence the common name “torchwood,” the wood produces a good
charcoal. The timber is useful for building poles, tool handles, grain mortars, stools
and for carving and turnery, in Swaziland it is used to make wagons. In southern
Malawi the fruits are used to make leg rattles. Although the fruit is edible to
mammals, the fruit exudate is used in fish poison and is lethal to the freshwater
Balanites aegyptiaca (L.) Delile . . . 193
snails and water fleas (Cyclops spp.) that are vectors of bilharzia and Guinea worm,
respectively. Occasionally been planted as an ornamental. Gum used as glue. Wood
used for utensils and tool handles (Beentje 1994). Dried leaves used to treat eye
problems in cows (Kokwaro 2009). Used as fuelwood, together with Balanites
glabra (Tian 2017). Used as fish poison (Neuwinger 2004).
Balanites pedicellaris: Used as fish poison (Neuwinger 2004).
Balanites rotundifolia: Wood used for utensils and spears, gum used as glue
(Beentje 1994). Used as fish poison (Neuwinger 2004).
References
Balemie K, Kebebew F. Ethnobotanical study of wild edible plants in Derashe and Kucha Districts,
South Ethiopia. J Ethnobiol Ethnomed. 2006;2:53. https://doi.org/10.1186/1746-4269-2-53.
El Hadidi MN. Flora of tropical East Africa: Zygophyllaceae. Rotterdam: Balkema; 1985.
Teklehaymanot T, Giday M, Mutse M. Quantitative ethnobotany of medicinal plants used by Kara and
Kwego semi-pastoralist people in lower Omo River Valley, Debub Omo Zone, Southern Nations,
Tian X. Ethnobotanical knowledge acquisition during daily chores: the firewood collection of
pastoral Maasai girls in Southern Kenya. J Ethnobiol Ethnomed. 2017;13:2. https://doi.org/
10.1186/s13002-016-0131-x.
Basella alba L.
BASELLACEAE

Grace N. Njoroge
Synonyms
Basella alba L.: Basella alba var. cordifolia (Lam.) M.R. Almeida, Basella
cordifolia Lam., Basella crassifolia Salisb., Basella japonica Burm. f., Basella
lucida L., Basella nigra Lour., Basella rubra L., Basella volubilis Salisb., Gandola
alba Rumph. ex. L., Gandola nigra (Lour.) Raf., Gandola rubra Rumph. ex L.
Local Names
Basella alba: Kikuyu: Murerema, Luhya: Indelema, Pokot: Rachan (Beentje 1994;
Gachathi 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Basella alba L.: Glabrous annual or shortly lived perennial, succulent tangled
twiner, stems much branched, 2–10 m long, sometimes almost leafless, greenish or
reddish. Leaf-lamina ovate to suborbicular, (2–)5–15 cm long, (1.25–)5–13.5 cm
broad, acute or acuminate (less commonly obtuse), usually widely cordate at the
base, lateral nerves 4–5 on either side, petiole (1–)2.5–6.5 cm long. Flowers white,
rose or purplish, (3–)4–5 mm long, in long-peduncled spikes, 2.5–15(–25) cm long,
usually unbranched (in African specimens at least) but branched in some cultivated
forms. Perianth fleshy, urceolate, somewhat saccate at the base, lobes short, ovate,
about one-third the length of the tube, not opening. Fruits 0.5 cm in diameter, 4–
7 5–10 mm, red, white or black, surface crinkly in the dry state. In thickets, forest
edges, margins of cultivated land and swampy ground, frequently by rivers or
streams, 0 (cultivated)–2450 m. Often in drier Afromontane forests (Bussmann
2002a, b) (Figs. 1, 2, 3, and 4).
Fig. 1 Basella alba

(Basellaceae), flowering
spike, Demaro, Bale
Mountains National Park,
Ethiopia. (Photo
R.W. Bussmann)
Basella alba L. 197
Fig. 2 Basella alba

(Basellaceae), flowers,
Demaro, Bale Mountains
Fig. 3 Basella alba

(Basellaceae), young fruits,
Fig. 4 Basella alba

(Basellaceae), mature fruits,
Basella alba: Leaf decoction drunk as galactagogue and to cure abdominal pain, and
constipation after birth. Also used to regularize menstruation (Jeruto et al. 2008;
Kokwaro 2009). The leaves are boiled and eaten for general weakness (Njoroge and
Bussmann 2006), the roots for diarrhea (Njoroge et al. 2004). In Ecuador, the roots
are also used to treat diarrhea (Bussmann and Sharon 2006), in India the leaves are
applied to burns, and taken for blood pressure problems. Other uses include the
treatment of tuberculosis, constipation, vomiting, and intestinal problems (Raj et al.
2018).
Local Food Uses
Basella alba: Eaten as vegetable (Njoroge and Bussmann 2006).

Basella alba L. 199
Basella alba: Leaf decoction given to bloated livestock (Kokwaro 2009). Eaten by
cattle and wildlife (Bussmann et al. 2011).
References
appendices 1–7. www.naturekenya.org/JournalEANH.htm.
Bussmann RW. Vegetation ecology and regeneration of tropical mountain forests. In: Ambasht RS,
Ambasht NK, editors. Modern trends in applied terrestrial ecology. Dordrecht: Kluwer; 2002b.
p. 195–223.
Northern Bengal, India. J Ethnobiol Ethnomed. 2018;141:8. https://doi.org/10.1186/s13002-
018-0208-9.
Berberis holstii Engl.
BERBERIDACEAE

Grace N. Njoroge
Local Names
Berberis holstii: Kikuyu: Mukekenye, Kamukurui, Kamba: Muthundi, Marakwet:

Kipsoroin, Maa: Olekikuuni, Meru: Murage, Shambaa: Kilasho (Beentje 1994;
Botany and Ecology
Berberis holstii Engl.: Glabrous shrub up to 3 m tall, branches purplish when young,
sulcate, with 3-partite spines up to 4 cm long. Leaves usually clustered on short
lateral shoots, almost sessile, apparently simple but in reality 3-foliolate with a
normal, terminal leaflet articulated at its base and a petiole of about 1 mm long
bearing 2 subulate lateral leaflets 1–3 mm long at its apex. The petioles and subulate
leaflets are characteristically persistent on the short shoots, lamina of terminal leaflet
up to 6 2.7 cm, coriaceous, oblong or oblanceolate to obovate, midrib produced as
R. W. Bussmann (*)
G. N. Njoroge

a short prickle, margin prickly dentate or more rarely entire, purplish when young
and somewhat glaucous below. Flowers yellow, in axillary racemes or cymes up to
5 cm long, bracts about 3 mm long, lanceolate-acuminate. Petals somewhat smaller
than the inner sepals, obovate, with two linear glands near the base. Sepals increas-
ing in size towards the interior, smallest about 2 2 mm, largest about 6.5 4 mm,
ovate to broadly ovate, rounded at the apex. Stamens on stout filaments about 2 mm
long, anthers about 2.5 mm long, oblong, opening by a pair of spreading wing-like
valves 1.5 mm long, hinged at the apex of the anther-thecae. Ovary narrowly
ellipsoid, about 4-ovulate, with a broad capitate subsessile stigma. Berry about
1.3 0.7 cm, ellipsoid, dark blue, pruinose, with a persistent stigma. Seed about
6 3 mm, usually solitary, ellipsoid, brown, rugulose. (Polhill 1966). Very common
shrub in the undergrowth of dry, Juniperus procera dominated Afromontane forests
(Bussmann 2002a, b, 2006a, b) (Figs. 1, 2, 3, 4, 5, 6, 7, and 8).
Fig. 1 Berberis vulgaris

(Berberidaceae), Tbilisi,
Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and
Berberis holstii Engl. 203
Berberis holstii: Root decoction as remedy for constipation, as laxative, and for
chest diseases (Kokwaro 2009).
Berberis species are used as medicinal plants worldwide, e.g., in India and
Pakistan where Berberis asiatica fruits are used as mild laxative for children, the

Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and

(Berberidaceae), Cappadokia,
Turkey. (Photo
R.W. Bussmann and

(Berberidaceae), dried fruits
for sale in market,
Cappadokia, Turkey. (Photo
R.W. Bussmann and

(Berberidaceae), fresh fruits in
market, Gori, Georgia. (Photo
R.W. Bussmann and
roots and bark as astringent, stomatic, diaphoretic, and to remedy piles (Malik et al.
2015, Joshi et al. 2010), with similar uses reported by Singh et al. (Singh et al. 2019);
and Berberis lyceum for eye problems and piles (Joshi et al. 2010), as well heal
wounds (Ur-Rahman et al. 2018), conjunctivitis and diabetes (Malik et al. 2015;
Singh et al. 2019), eye infections (Joshi et al. 2013), jaundice, fever and urinary
infections (Sher et al. 2016), for bone fractures, pneumonia, headache, stomachic,
arthritis, wound healing, to speed delivery (Wali et al. 2019a, b), and blood purifi-
cation (Muhammad et al. 2019). In Nepal, B. asiatica is used for eye problems
(Kunwar et al. 2013, 2015), andserves for gastric problems, as anthelminthic, for
diabetes and eye infections (Malik et al. 2015; Joshi et al. 2010; Kunwar et al. 2009;
Kumar et al. 2011, 2013), and also as mild laxative, especially for children (Joshi
et al. 2013).
Berberis buceronis is used in Peru for liver problems and hepatitis (Bussmann
and Sharon 2006, 2015; Monigatti et al. 2013), respiratory and nervous system
disorders, and gynecological problems (Bussmann and Glenn 2010a, b; Bussmann
et al. 2010, 2011a). The plants show antibacterial properties (Bussmann et al.
2010d), and essentially no toxicity (Bussmann et al. 2011b). Packages labeled as
“Berberis vulgaris” are distributed to patients in the Peruvian social security health
system, and Berberis species are widely sold in markets (Bussmann et al. 2007a, b,
2008, 2009).
Local Food Uses
The fruits are eaten fresh, and dried for use in sauces, for making jams and sweets, as
well as spice. (Batsatsashvili et al. 2017; Bussmann et al. 2014, 2016a, b, c, 2017,
2018; Bussmann 2017; Fedorov 1984). The fruits of Berb eris brandisiana are also
eaten (Abbas et al. 2019).
Berrberis holstii has no use among the Samburu (Bussmann 2006). The bark of
many species however yields yellow, golden, dark violet, dark blue and olive dyes
for wool and silk. (Batsatsashvili et al. 2017; Bussmann et al. 2014, 2016a, b, c,
2017, 2018; Bussmann 2017; Fedorov 1984). Berberis lyceum is also used for
forage, and widely sold (Ahmad Jan et al. 2019; Ullah et al. 2019; Wali et al.
2019a, b), equally to Berberis balochistanica (Umair et al. 2019). The leaf and
bark powder of Berberis lyceum is given to livestock for internal wounds) (Ali et al.
2019). The spiny branches of Berberis brandisiana are used as fence around
gardens, house, and hut material. Whole plant is also used as fuel wood (Abbas
et al. 2019).
References
Abbas Z, Alam J, Muhammad S, Bussmann RW, Mulk Khan S, Hussain M. Phyto-cultural diversity
of the Shigar valley Central Karakorum Baltistan, northern Pakistan. Ethnobot Res Appl.
2019;1832. https://doi.org/10.32859/era.18.31.1-18.
Ahmad Jan H, Jan S, Bussmann RW, Wali S, Sisto F, Ahmad L. Complementary and alternative
medicine research, prospects and limitations in Pakistan: a literature review. Acta Ecol Sin.
2019. https://doi.org/10.1016/j.chnaes.2019.12.005.
Ali A, Aldosari A, Tng DYP, Ullah M, Hussain W, Ahmad M, Hussain J, Khan, A, Hussain H,
Sher H, Bussmann RW, Shao J-W. Traditional uses of plants by indigenous communities for
veterinary practices at Kurram District, Pakistan. Ethnobot Res Appl. 2019;1824. https://doi.
org/10.32859/era.18.24.1-19.
Batsatsashvili K, Mehdiyeva N, Fayvush G, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S,
Tchelidze D, Aleksanyan A, Alizade V, Paniagua Zambrana NY, Bussmann RW. Berberis
vulgaris L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer Interna-
tional Publishing; 2017.
2006b;111:41–66.
2017. XXVII, 746 p. ISBN 978-3-319-49411-1.
Bussmann RW, Glenn A. Traditional Peruvian medicine for the treatment of respiratory disorders.
Revista Peruana de Biologia. 2010a;17(2):331–46.
Bussmann RW, Glenn A. Medicinal plants used in northern Peru for reproductive problems and
female health. J Ethnobiol Ethnomed. 2010b;6:30.
Bussmann RW, Sharon D. Traditional plant use in northern Peru: tracking two thousand years of
Bussmann RW, Sharon D. Medicinal plants of the Andes and the Amazon – the magic and
medicinal flora of northern Peru. St. Louis: MBG/William L. Brown Center; 2015. ISBN
978-0-9960231-2-2.
Bussmann RW, Sharon D, Vandebroek I, Jones A, Revene Z. Health for sale: the medicinal plant
markets in Trujillo and Chiclayo, northern Peru. J Ethnobiol Ethnomed. 2007a;3:37.
Bussmann RW, Sharon D, Lopez A. Blending traditional and Western medicine: medicinal plant use
amongst patients at Clinica Anticona in El Porvenir, Peru. Ethnobot Res Appl. 2007b;5:185–99.
Bussmann RW, Sharon D, Ly J. From garden to market? The cultivation of native and introduced
medicinal plant species in Cajamarca, Peru and implications habitat conservation. Ethnobot Res
Appl. 2008;6:351–61.
Bussmann RW, Sharon D, Garcia M. From chamomile to aspirin? Medicinal plant use among
clients at Laboratorios Beal in Trujillo, Peru. Ethnobot Res Appl. 2009;7:399–407.
Bussmann RW, Sharon D, Glenn A. Healing the body, healing the soul. Traditional remedies for
“magical” ailments, nervous system and psychosomatic disorders in northern Peru. Afr J Pharm
Pharmacol. 2010;4(9):580–629.
northern Peruvian medicinal plants. Ethnobot Res Appl. 2011a;9:67–96.
Bussmann RW, Malca G, Glenn A, Sharon D, Nilsen B, Parris B, Dubose D, Ruiz D, Saleda J,
Martinez M, Carillo L, Walker K, Kuhlman A, Townesmith A. Toxicity of medicinal plants used
in northern Peru. J Ethnopharmacol. 2011b;137:121–40.
2014;12:237–313.
Samtskhe-Javakheti, Tusheti, Svaneti, and Racha-Lechkhumi, Republic of Georgia
(Sakartvelo), Caucasus. J Ehnobiol Ethnomed. 2016a;12:43. https://doi.org/10.1186/s13002-
016-0110-2.
(Republic of Georgia), Caucasus. Med Aromat Plants. 2016b;5:266. https://doi.org/10.4172/
2167-0412.1000266.
Batsatsashvili K, Hart RE. Ethnobotany of Samtskhe-Javakheti, Sakartvelo (Republic of Geor-
gia), Caucasus. Indian J Tradit Knowl. 2017;16(1):7–24.
Sakartvelo (Republic of Georgia), Caucasus. Indian J Tradit Knowl. 2018;17(1):7–33.
Bussmann RW, Batsatsashvili K, Kikvidze Z. Berberis jamesiana Forrest & W.W. Sm.; Berberis
vulgaris L. In: Batsatsashvili, K; Kikvidze, Z; Bussmann, RW (Eds.) Ethnobotany of Mountain
Regions Central Asia and Altai. Springer International Publishing International Publishing.
2020. https://doi.org/10.1007/978-3-319-77087-1_26-1.
Fedorov AA, editor. Plant resources of the USSR: flowering plants, their chemical composition,
use. Volume 1. Families Magnoliaceae – Limoniaceae. Leningrad: Akademia Nauk; 1984.
Joshi M, Kumar M, Bussmann RW. Ethnomedicinal uses of plant resources of the Haigad watershed
in Kumaun Himalaya. Med Aromat Plant Sci Biotechnol. 2010;4(special issue 1):43–6.
Kunwa RM, Baral K, Paudel P, Acharya RP, Thapa-Magar KB, Cameron M, Bussmann RW. Land-
use and socioeconomic change, medicinal plant selection and biodiversity resilience in far
western Nepal. PlosONE. 2016;11. https://doi.org/10.1371/journal.pone.0167812.
Kunwar RM, Bussmann RW. Ethnobotany in the Nepal Himalaya: a review. J Ethnobiol Ethnomed.
2008;4:24.
Biodiversität, Naturausstattung im Himalaya, vol. III. Erfurt: Universität Erfurt; 2009, p. 475–
89.
Kunwar RM, Nepal BK, Kshetri HB, Rai SK, Bussmann RW. Ethnomedicine in Himalaya: a case
study from Dolpa, Humla, Jumla and Mustang districts of Nepal. J Ethnobiol Ethnomed.
2006;2:27.
Kunwar RM, Chowdhary A, Bussmann RW. Diversity, utilization and management of medicinal
plants in Baitadi and Darchula districts, farwest Nepal. Initiation. 2008;21:157–64.
Kunwar RM, Upreti Y, Burlakoti C, Chowdhary C, Bussmann RW. Indigenous use and ethnophar-
macology of medicinal plants in Far-west Nepal. Ethnobot Res Appl. 2009;7:5–28.
P. Cross-cultural comparison of plant use knowledge in Baitadi and Darchula districts, Nepal
Kunwar RM, Fadiman M, Hindle T, Suwal MK, Adhikari YP, Baral K, Bussmann
RW. Composition of forests and vegetation in the Kailash Sacred Landscape, Nepal. J For
Res. 2019a. https://doi.org/10.1007/s11676-019-00987-w.
Kunwar RM, Shrestha K, Malla S, Acharya T, Sementelli A, Kutal D, Bussmann RW. Relation of
medicinal plants, their use patterns and availability in the lower Kailash Sacred Landscape,
Nepal. Ethnobot Res Appl. 2019b;187. https://doi.org/10.32859/era.18.6.1-14.
Kunwar RM, Fadiman M, Thapa S, Acharya RP, Cameron M, Bussmann RW. Plant use values and
phytosociological indicators: implications for conservation in the Kailash Sacred Landscape,
Nepal. Ecol Indic. 2020;108:105679. https://doi.org/10.1016/j.ecolind.2019.105679.
Monigatti M, Bussmann RW, Weckerle CS. Medicinal plant use in two Andean communities
located at different altitudes in the Bolivar Province, Peru. J Ethnopharmacol. 2013;145
(2):450–64.
Muhammad S, Hussain M, Abbas Z, Saquib Z, Bussmann RW, Shah GM. An ethnomedicinal
appraisal of Kurram Agency Tribal area Pakistan. Indian J Tradit Knowl. 2019;184:631–47.
Paniagua Zambrana NY, Bussmann RW, Romero, C. Berberis rigidifolia Kunth ex DC. In: Paniagua
Zambrana NY, Bussmann RW (Eds.) Ethnobotany of Mountain Regions – Ethnobotany of the
Andes. Springer International Publishing, Cham. 2020. https://doi.org/10.1007/978-3-319-
77093-2_37-1.
Polhill RM. Flora of tropical East Africa: Berberidaceae. London: Crown Agents for Overseas
Governments & Administrations; 1966. 4 pp. ISBN 978-1-84246-233-1
Singh AP, Kumar M, Nagar B, Pala NA, Bussmann RW. Ethnomedicinal use of plant resources in
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;1814.
https://doi.org/10.32859/era.18.14.1-11.
the southern and tribal regions of Khyber Pakhtunkhwa province, Pakistan. Ethnobot Res Appl.
2019;188. https://doi.org/10.32859/era.18.8.1-20.
10.1186/s13002-019-0285-4.
and ethnomedicinal dermatology practices in Pakistan. Clin Dermatol. 2018;363:310–9. https://
doi.org/10.1016/j.clindermatol.2018.03.018.
Wali R, Rahman K, Raja NI, Qureshi R, Bussmann RW, Mashwani ZUR. A quantitative medico
botanical expedition of Fairy Meadows National Park, Diamir, Gilgit Baltistan, Pakistan.
Ethnobot Res Appl. 2019a;1835. https://doi.org/10.32859/era.18.35.1-30.
Wali S, Ahmad Jan H, Bussmann RW. Quantitative ethnomedicinal study of indigenous medicinal
plants used for digestive disorders of Laspur Valley, Chitral, northern Pakistan. Ethnobot Res
Appl. 2019b;1832. https://doi.org/10.32859/era.18.32.1-18.
Bersama abyssinica Fresen.
FRANCOACEAE

Grace N. Njoroge
Local Names
Bersama abyssinica: Kikuyu: Murumandu, Muthandi, Kipsigis: Cheptoroguet,

Ororuewet, Kisii: Omubamba, Luhya: Shirikamabinga, Shiviambinga-Mekundi,
Maa: Olobaye-Tiongata, Marakwet: Kipset, Kipsagas, Meru: Muthanda-Thandi,
Nandi: Kibuimetiet, Marakwet: Kapsagas, Kipsagas, Ndorobo: Kipteleliet,
Kipuimetuet, Sebei: Morgenet, Sigirwo, Tugen: Kipumetiet, Hehe: Mbatsamwono,
Nyakyusa: Misunangatu, Swahili: Mnangwakwao, Kaguru: Mkururumbi,
Chagga: Mirindwa (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Bersama abyssinicaI Fresen.: Small to medium-sized tree (rarely shrubby) with

smooth to rather rough, grey, brown or mottled bark. Leaves up to 1 m long,
imparipinnate, rhachis wingless to very widely winged, leaflets up to 10-jugate,
lanceolate to oblong or ovate-oblong, glabrous to hairy, often densely so beneath,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Bersama abyssinica

(Francoaceae), leaves,
Kenya. (Photo
R.W. Bussmann)

(Francoaceae), flowering
branch, Bale Mountain
National Park, Demaro forest,
Ethiopia. (Photo
R.W. Bussmann)
3.5–22 cm long and 1–8 cm wide, margins quite entire to sharply and conspicuously
serrate. Flowers creamy-white, yellowish or pink-tinged, especially in bud, in erect
or rather drooping, long, glabrescent to hirsute racemes. Disc Λ-shaped to annular-
pentagonal. Capsules reddish-pink, seeds bright red with yellow aril (Verdcourt
1958). The species is an important component in drier, Juniperus or Cassipourea
dominated Afromontane forests (Bussmann 2002a, b) (Figs. 1, 2, 3, 4, 5, and 6).
Bersama abyssinica: Leaves crushed and used as snuff for colds, chewed as
aphrodisiac. Bark juice used as purgative. Fried powdered bark or bark decoction
as anthelminthic for small children. Juice from young twigs is drunk for diarrhea and
roundworms. Root decoction for epilepsy, hemorrhoids, and for washing wounds. In
Bersama abyssinica Fresen. 213

(Francoaceae), flowers, Bale
Mountain National Park,
Demaro forest, Ethiopia.

(Francoaceae), flowers detail,
Bale Mountain National Park,
Small quantities for syphilis, cancer, and female infertility. To be used with great care
due to toxicity (Kokwaro 2009). The stem bark is used to cure tonsillitis (Giday et al.
2009). The stem bark is boiled, and the extract used to remedy sexually transmitted
diseases (Njoroge and Bussmann 2007; Yineger et al. 2008).
Bersama abyssinica: Wood soft but durable (Beentje 1994). Used to make tools,
especially plows, as well as used as firewood (Bussmann et al. 2011). The leaves are
boiled, and the extract painted over the body of horses for strength and as anti-
parasitic (Yineger et al. 2007).

(Francoaceae), young
inflorescence, Bale Mountain
Ethiopia. (Photo
R.W. Bussmann)

(Francoaceae), flowers detail,
Bale Mountain National Park,
Bersama abyssinica Fresen. 215
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. Journal of East African Natural History.
2002a;91 1/2:27–79, appendices 1-7 @ http://www.naturekenya.org/JournalEANH.htm.
Syst Geogr Plants. 2002b;712:959–74.
Giday M, Asfaw Z, Woldu Z. Medicinal plants of the Meinit ethnic group of Ethiopìa: an
ethnobotanical study. J Ethnopharmacol. 2009;124:513–21.
Verdcourt B. Flora of tropical East Africa: Melianthaceae. London: Crown Agents for Overseas
Governments & Administrations; 1958. 7 pp. ISBN 978-1-84246-450-2.
Yineger H, Yewhalaw D, Teketay D. Ethnomedicinal plant knowledge and practice of the Oromo
ethnic group in southwestern Ethiopia. J Ethnobiol Ethnomed. 2008;4:11. https://doi.org/
10.1186/1746-4269-4-11.
Bidens pilosa L.
ASTERACEAE

Grace N. Njoroge
Synonyms
Bidens pilosa L.: Bidens alausensis Kunth, Bidens alba (L.) DC., Bidens alba var.
radiata (Sch. Bip.) R.E. Ballard ex Melchert, Bidens chilensis DC., Bidens hirsuta
Nutt., Bidens hispida Kunth, Bidens leucantha (L.) Willd. ex Walp., Bidens
leucantha fo. discoidea Sch. Bip., Bidens leucantha var. pilosa (L.) Grieseb., Bidens
leucanthema (L.) Willd., Bidens leucanthema fo. discoidea Sch. Bip.,
Bidens leucanthema var. pilosa (L.) Grieseb, Bidens montaubani Phil., Bidens
odorata Cav., Bidens pilosa fo. discoides Sch. Bip., Bidens pilosa fo. indivisa
Sherf., Bidens pilosa fo. radiata Sch. Bip., Bidens pilosa fo. rubiflora S.S. Ying,
Bidens pilosa var. alausesis (Kunth) Sherf, Bidens pilosa var. discoidea (Sch. Bip.)
J.A. Schmidt, Bidens pilosa var. minor (Blume) Sherf, Bidens pilosa L. var. pilosa,
Bidens pilosa var. radiata (Sch. Bip.) Schmidt, Bidens pilosa var. radiata Sch. Bip.,
Bidens pilosa var. subinternata Kuntze, Bidens reflexa Link, Bidens scandicina
Kunth, Bidens sundaica var. minor Blume, Centipeda minuta (G. Forst.) Benth. ex
C.B. Clarke, Centipeda orbicularis Lour., Coreopsis leucantha L., Coreopsis
leucanthema L., Cotula minuta G. Forst., Kerneria pilosa (L.) Lowe, Kerneria
R. W. Bussmann (*)
G. N. Njoroge

pilosa var. discoides (Sch. Bip.) Lowe, Kerneria tertagona Moench, Myriogyne
minuta (G. Forst.) Less.
Local Names
Bidens pilosa: Kikuyu: Mucege, Muheneje, Meru: Munyugunyugu, Kamba:

Munzee, Luhya: Ologoye, Luo: Onyiego, Nyanyiek-Mon, Iraqw: Qalamir Bo,
Padhola: Sere (Gachati 1989; Kokwaro 2009).
Botany and Ecology
Bidens pilosa L.: Annual herb, erect, 0.1–1.5 m high, stem much branched or
sometimes hardly branched, pale green or sometimes reddish, 4-angled, sparsely
pubescent to glabrous. Leaves pinnately 3–5-lobed or occasionally simple in upper
or lowermost part of stem, up to 15( 20) cm long, segments ovate to ovate-
lanceolate, margins serrate or crenate-serrate, sparsely pubescent to glabrous, termi-
nal segment 5–10 cm long, 2.5–5 cm wide, lateral segments 2–5 cm long, 1–2 cm
wide and asymmetric, often lobed or bilobed, petiole to 70 mm long. Ray florets
white or cream, sometimes absent, neuter or with pistillodes or staminodes, 4–8, rays
(when fully developed) 7–15 3–4.5 mm, tube 0.5–0.8 mm long, disc florets yellow
or yellow-orange, 4 mm long, pubescent at base. Capitula terminal and solitary or
few together, peduncle to 16 cm long, outer phyllaries 7–10, 3–4 mm long, reflexed
at anthesis, inner phyllaries 5–8, yellow-green to pale brown, 3–4.5 mm long with
yellow scarious margins, glabrous but for the apex, receptacle club-shaped in fruit,
palea light brown, striate, 3–5 mm long. Achenes black, 4–6-ribbed, linear-
tetragonal, 4–12 mm long, strigose or verrucose, aristae 2–3( 4), 2–4 mm long,
retrorsely barbed (Beentje et al. 2005). Common in dry Afromontane forests
(Bussmann 2002) (Figs. 1 and 2).
Bidens pilosa: Leaves are squeezed, and juice applied to eyes for conjunctivitis.
Root juice taken for malaria. Crushed leaves mixed with water drunk for stomach-
ache. Leaf decoction as anthelminthic and for constipation, crushed leaves applied to
wounds and snakebites. Root infusion as cough syrup (Kokwaro 2009). In China, the
whole plant is used to treat appendicitis, gastroenteritis, analgia, and is applied
externally to boils (Ma et al. 2019). Used also as smoke for gynecological problems
(Mohagheghzadeh and Faridi 2006) and ingested to increase fertility in women
(Njoroge and Bussmann 2009). Crushed leaves are inserted into the nose to stop
nosebleeds (Njoroge and Bussmann 2006). The species is also used as
Bidens pilosa L. 219
Fig. 1 Bidens pilosa

(Asteraceae), flowering shoot,
Kenya. (Photo R.W:
Bussmann)
Fig. 2 Bidens pilosa

(Asteraceae), fruits,
Kenya. (Photo R.W:
Bussmann)
anthelminthic, coagulant, for diarrhea, and stomach problems (Njoroge et al. 2004).
Bidens macrocarpa is used to treat kidney disease (Yineger et al. 2008). The leaves
and seeds are boiled, and the extract ingested to expel a retained placenta after birth
Bidens pilosa: Sometimes browsed by livestock, but mostly useless (Bussmann

et al. 2006).
References
Beentje H, Jeffrey C, Hind DJN, editors. Flora of tropical East Africa: Compositae (part 3). Kew:
Royal Botanic Gardens; 2005.
Bussmann RW.. Islands in the desert – forest vegetation of Kenya’s smaller mountains and highland
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002;91 1/2:27–79,
appendices 1–7 @ www.naturekenya.org/JournalEANH.htm.
doi.org/10.1186/1746-4269-5-34.
Ma Y, Liu D, Cheng H, Bussmann RW, He H, Guo Z, Liu B. Ethnobotanical study of medicinal
plants used by Miao people in Jijiezi, Yunnan, China. Ethnobot Res Appl 2019;1826. https://doi.
org/10.32859/era.18.26.1-14.
2009;82:262–9.
2008;26:132–53.
Blighia unijugata Bak.
SAPINDACEAE

Grace N. Njoroge
Synonyms
Blighia unijugata Bak.: Phialodiscus unijugatus (Baker) Radkl.
Local Names
Blighia unijugata: Kikuyu: Muikuni (Beentje 1994; Gachati 1989; Kokwaro

2009).
Botany and Ecology
Blighia unijugata Bak.: Tree, probably dioecious, usually 6–9 m but sometimes
reaching 30 m. tall, with a dense crown, bark grey-brown smooth, thin, with warts
and horizontal ridges, slash white, pale red or brown with white streaks, young twigs
R. W. Bussmann (*)
G. N. Njoroge

golden-hairy, glabrescent, dark grey. Leaves reddish when young, drying bright
green or brownish, petiole up to 4 cm long, rhachis 0.6–10 cm long, petiolules 1–
2 mm long, leaflets in (1–)2–4( 5) pairs, elliptic, oblong, obovate or oblanceolate,
the upper pair 3–26 cm long, 1.2–9.5 cm wide (but in West Africa up to 30 13 cm),
the lowest either similar (usually when there are only two pairs) or as small as
2 1.2 cm, acuminate at apex and tapering or sometimes rounded at the extreme tip,
the midrib visible as a raised line on the upper surface, glabrous apart from some
hairs on the veins plus tufts in the nerve axils, lateral veins in 6–10( 12) pairs,
curving and ending well inside the leaflet margin, usually whitish beneath, glabrous.
Inflorescences borne in axils of current leaves, 5–10 cm long, flowers singly inserted
or in cymose groups, pedicels 1–5( 10 in fruit) mm long, pubescent. Flowers
whitish to yellow, sweet-scented, calyx divided halfway, lobes 2 mm long, 1 mm
wide, petals rhomboid in outline, 1.5 mm long, 1.5–2 mm wide, scale bilobed,
shaggy hairy. Stamens 6–8, filaments 4.5 mm long, the basal part strikingly white
hairy. Capsule reddish, (2–)3( 4)-gonous and -locular, 1.7–3 cm long, 1.9–3 cm
wide, glabrescent, the angles acute or with wings up to 2 mm broad, style and sepals
persistent, endocarp pink with yellow margins, glabrous or nearly so. Seeds, dark
brown or black, ovoid 1.5–2 cm long, 6–8 mm wide, aril bright yellow, up to 1 cm
long (Davies and Verdcourt 1998). In the undergrowth of wet, Ocotea and Lovoa
dominated Afromontane forests (Bussmann 2002a, b; Kiefer and Bussmann 2004).
Blighia unijugata: Very good firewood (Kiefer and Bussmann 2008). Sometimes
Blighia species are used as fish-poison (Neuwinger 2004).
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. Journal of East African Natural History
2002a;91 1/2:27–79, appendices 1–7 @ http://www.naturekenya.org/JournalEANH.htm.
Davies FG, Verdcourt B. Flora of Tropical East Africa: Sapindaceae. Rotterdam: Balkema; 1998.
108 pp. ISBN 90-6191-380-2 [Fl. Trop. E. Africa, Sapind.].
2004;62:161–9.
Bothriocline longipes (Oliv. & Hiern.) N.E. Br.
ASTERACEAE

Grace N. Njoroge
Synonyms
Bothriocline longipes (Oliv. & Hiern.) N.E. Br.: Bothriocline eupatorioides

(Hutsch. & B.L. Burtt) Wild. & G.V. Pope, Bothriocline schimperi var. longipes
Oliv. & Hiern, Bothrocline schimperi var. tomentosa Oliv. & Hiern, Bothriocline
tomentosa (Oliv. & Hiern) Willd. & G.V. Pope, Erlangea eupatorioides Hutsch. &
B.L. Burtt, Erlangea longipes (Oliv. & Hiern) S. Moore, Erlangea pubescens
S. Moore, Erlangea spissa S. Moore, Erlangea squarrulosa Chiov., Erlangea
tomentosa (Oliv. & Hiern) S. Moore, Erlangea tomentosa var. acuta R.E. Fr.
Local Names
Bothriocline longipes: Shambaa: Boha, Mtuyu, Luhya: Matendi, Chagga:

Maurie, Samburu: Lodoporo (Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Bothriocline longipes (Oliv. & Hiern.) N.E. Br.: A slender to bushy, long-lived
annual or perennial herb, 0.2–1 m tall. Stems leafy, sparsely to much-branched in the
upper part, becoming stout and woody at the base to 2 cm in diameter, strongly
ribbed at least above, pilose and glandular, the hairs flagelliform, branches spread-
ing, to 40 cm long. Leaves alternate, sessile or subsessile, mostly 3–14 0.8–4 cm,
narrowly oblong to elliptic or ovate-oblong, apex obtuse, base rounded to
sub-cordate or less often cuneate, margins subentire to serrate sometimes coarsely
remote or dentate-lobed, shortly pilose or pubescent, sparsely so on the upper
surface, hairs flagelliform. Capitula many, laxly corymbiform cymose, stalks slen-
der, 0.3–9 cm long. Involucres 5–7 9–15 mm, broadly campanulate. Phyllaries
numerous, lanceolate, the innermost lorate, all tapering acuminate and bristle-
Upped, margins narrowly subhyaline and finely pectinate towards the apex, pilose
and glandular, obscurely 3-nerved, the outer from 2 mm long, the inner to 7 mm
long. Corollas purple to dark-mauve, to 6 mm long, narrowly funnel-shaped, the
lobes with short acicular-setae and a few flagelliform hairs, limb and tube glandular
outside. Achenes numerous, 1.2–1.8 mm long, narrowly oblong-ovoid, 4-ribbed,
rugulose and with globose glandular trichomes sunk in pits between the ribs,
sparsely setulose or glabrous except for glands, pappus of caducous sub-plumose
bristles 3–4 mm long. Very common in the undergrowth of Podocarpus and Olea
dominated Afromontane forests (Bussmann 2006a) (Figs. 1 and 2).
Bothriocline longipes: Roots and leaves are crushed, and the extract drunk as anti-
emetic. Leaf paste is applied to sores caused by maggots. Raw roots are chewed for
sore throat, a leaf decoction drunk for malaria treatment (Kokwaro 2009). A leaf
decoction serves for fever and can also be used as purgative to vomit in cases of
Fig. 1 Bothriochline
longipes (Asteraceae),
Naro Moru, Kenya. (Photo
R.W. Bussmann)
Bothriocline longipes (Oliv. & Hiern.) N.E. Br. 225
Fig. 2 Bothriochline
longipes (Asteraceae),
flowers, Mt. Kenya National
Park, Naro Moru, Kenya.
malaria. It is also used to wash babies to give them strength (Bussmann 2006b). The
roots of Bothriocline schimperi are chewed and applied externally for stomach
problems (Giday et al. 2009).
Bothriocline longipes: The species is eaten by livestock, and the branches are used
in construction of huts, and for firewood (Bussmann 2006b).
References
2006a;111:41–66.
Brachylaena huillensis O. Hoffm.
ASTERACEAE

Grace N. Njoroge
Synonyms
Brachylaena huillensis O. Hoffm.: Brachylaena hutchinsii Hutch.
Local Names
Brachylaena huillensis: Kikuyu: Muhugu, Mugugu, Swahili: Muhuhu, Boni:

Abozi, Avud, Kamba: Muvuuvu, Sanya: Mshenzi, Wato, Taita: Kipungupungu
(Beentje 1994; Gachati 1989).
Botany and Ecology
Brachylaena huillensis O. Hoffm.: Evergreen or deciduous shrub or slender erect

tree (2–)5–40 m tall, with ascending branches and rather light crown, trunk up to
60 cm in diameter, usually very fluted, eventually buttressed, crown thin, confined to
R. W. Bussmann (*)
G. N. Njoroge

upper quarter of stem, bark light or dark grey, longitudinally fissured, fibrous, peeling
in long narrow shreds, wood white, hard, without visible heartwood, rays or rings,
branches porrect, not spreading, young twigs densely shortly white- or grey-
tomentellous, glabrescent and becoming dark purplish, lenticellate and narrowly
sulcate. Leaves crowded at the ends of twigs, coriaceous, shortly petiolate, blade
(narrowly) obovate to elliptic or broadly so, 3–12( 15) 1–3( 5.5) cm, base
cuneate or rarely rounded, margins entire or sometimes remotely denticulate (some-
times spinescent in obovate leaves) and slightly to strongly revolute, apex acute to
obtuse and shortly acutely acuminate, mucronate, strongly discolorous, green and
araneose becoming glabrous adaxially, silvery white or grey with short dense persis-
tent tomentum and glandular abaxially, secondary veins 8–15, petiole densely shortly
white-tomentose, 0.2–1.4 cm long. Capitula numerous in axillary or terminal racemes
or panicles 1.5–4 cm long. Male inflorescence with capitula sessile or shortly stalked
(stalk to 2 mm), 3–5.5 mm high, in glomerules 1 cm in diameter and these in turn
sometimes racemosely arranged, bracts small, narrow, acute, densely tomentose
proximally, capitula with (4–)6–9( 12) florets, involucre cylindrical or campan-
ulate, 2–4 mm long, phyllaries 5–8, densely white-tomentose, narrowly ovate to
elliptic, acute, 1–5 1.5–3 mm, corolla tube 1.3–2 mm long, lobes narrowly oblong,
2–3 0.4–0.6 mm, glandular, with a distal tuft of hair, recurved, anthers 1.3–2.1 mm
long, not or hardly tailed, completely exserted on filaments 0.3–1.3 mm long, style
3.7–7 mm long with linear obtuse branches, the distal mm with short trichomes
abaxially, achene minute, pappus of few bristles, 2–3 mm long and sub-plumose.
Female inflorescence with capitula 2–6 together in axillary clusters or sometimes in
racemes to 7 cm long, the individual capitula stalked (stalk to 2 mm), 3–6-flowered,
8–11 mm high, bracts small, narrow, acute, densely tomentose proximally, involucre
cylindrical or campanulate, 3–6 mm long, phyllaries 5–8, densely white-tomentose,
glabrescent with age, narrowly ovate to elliptic, acute, 1–5.2 1.5–3 mm, corolla
tube slender, cylindrical, 3.5–6.2 mm long, lobes slender, narrowly ovate, 0.5–1 mm
long, style 5–7 mm long, the branches oblong-linear, acute, glabrous, recurved.
Achenes columnar or flattened, 5–8-ribbed, slightly narrowed towards the base,
densely pubescent and glandular, 2.8–4 mm long, pappus of many pale tawny
minutely barbellate setae, 4–6.5 mm long, 2-seriate. Deciduous in dry areas, ever-
green in wetter sites. In the southern part of the distribution area it remains much
smaller (usually up to 7 m, rarely to 25 m), but in Kenya it is usually a tall tree (Beentje
2000). The species was originally the dominant forest tree on the central Kenyan
plateau, forming dry Afromontane forests, often together with Croton species, and is
especially characteristic for plinthosols – soils with layers of iron pellets. Nowadays
most of these forests have been destroyed (Bussmann 2002).
Brachylaena merana serves as aphrosisiac in Madagascar (Rakotoarivelo et al.

2013). Brachylaena ramiflora is often sold in markets in Madagascar, and the
leaves are used for asthenia and to treat stomach ulcers (Randriamiharisoa et al.
2015).
Brachylaena huillensis O. Hoffm. 229
Brachylaena huillensis is used for carvings, as construction timber, and excellent

firewood. Smells of sandalwood (Beentje 1994). Brachylaena merana is used as
firewood, for tools and construction, and for ritual purposes in Madagascar
(Rakotoarivelo et al. 2013).
References
Beentje HJ. The genus Brachylaena (Compositae: Mutisieae). Kew Bull. 2000;55(1):1–41.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi). J East Afr Nat Hist 2002;91 (1/2):27–79,
appendices 1–7 @ http://www.naturekenya.org/JournalEANH.htm.
Randrianasolo A, Bussmann RW. Guide de plantes d’Ambalabe, vol. 1. St. Louis: William
org/10.1186/s13002-015-0046-y.
Brucea antidysenterica J.F. Mill.
SIMAROUBACEAE

Grace N. Njoroge
Synonyms
Brucea antidysenterica J.F. Mill.: Brucea abyssinica Spreng., Brucea erythraeae

Chiov., Brucea ferruginea L’Hér., Brucea salutaris A. Chev., Lussa antidysenterica
Kuntze, Trichilia siderotricha Chiov.
Local Names
Brucea antidysenterica: Kamba: Mukambe, Kikuyu: Mutingombe, Samburu:

Songori, Chagga: Mkamana, Mirikawanda (Beentje 1994).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Brucea antidysenterica J.F. Mill.: Shrub or small tree to 5 m tall, young twigs stout,
pubescent, older twigs with roughly triangular leaf-scars 5–10 4–9 mm, pubescence
golden-brown. Leaves 15–60 cm. long, with 3–5 pairs of leaflets, pubescent above and
below, most densely on the nerves below, rarely almost glabrous, petiole 8–15 cm. long,
leaflet-laminas 4–13 2–6 cm, oblong to narrowly ovate, opposite, apex subacute,
margins entire to shallowly repand, base asymmetric truncate to cuneate, petiolules up to
5 mm long. Inflorescences 12–35 cm long, andromonoecious, of spiciform panicles
bearing distant glomerules (or rarely short spiciform racemes) along the main axis,
axillary, borne towards the end of the young shoots, pubescent, bracts subulate, pubes-
cent. Flowers not opening widely. Sepals 1 mm long, ovate, pubescent at least outside.
Petals up to 2 mm long, similar to the sepals, pubescent at least outside. Stamens 1 mm
long, filaments glabrous. Disk pulviniform, margin 4-lobed. Gynoecium (absent in the
male flowers) in the bisexual flowers consisting of free, ovoid, usually glabrous carpels
pressed together at first with the stigmas bent outwards in a cruciform arrangement. Fruit
developing from 1–4 of the carpels in each flower and consisting of red ovoid drupa-
ceous mericarps 10 6 mm acute at the apex. Brucea antidysenterica occurs widely in
tropical Africa, from Guinea and Nigeria east to Ethiopia and south to Angola, Malawi,
and Zambia. (Stannard 2000). Characteristic component in humid Afromontane forests
dominated by Faurea saligna and Ilex mitis (Bussmann 2002) (Figs. 1, 2, 3, and 4).
Brucea antidysenterica: As its name suggests, the species is used in traditional

medicine for the treatment of dysentery (Beentje 1994). The bark, fruit, and roots are
widely used against dysentery, as an anthelmintic and to treat fever. The bark, fruit,
Fig. 1 Brucea
antidysenterica
(Simaroubaceae). Young
plant, Mt. Kenya National
Brucea antidysenterica J.F. Mill. 233
Fig. 2 Brucea
antidysenterica
(Simaroubaceae). Flowers,
R.W. Bussmann)
Fig. 3 Brucea
antidysenterica
(Simaroubaceae). Young
fruits, Mt. Kenya National
Fig. 4 Brucea
antidysenterica
(Simaroubaceae). Fruits,
R.W. Bussmann)
seeds, leaves, and roots (sometimes boiled) are used as a remedy for diarrhea,
indigestion, and stomach-ache (Kokwaro 2009). The leaves and roots are cooked
with meat or infused with milk (for children) to relieve asthma. Wounds and skin
complaints, such as those caused by leprosy and scrofula, are treated leaves and
twigs mixed with ghee or butter, or from the ripe fruits mixed with honey (Kokwaro
2009). Preparations of the roots are used on sores caused by sexually transmitted
diseases, while the leaves and seeds are used to treat cancerous skin tumors. The bark
is boiled to treat malaria (Bussmann 2006; Mekonen et al. 2015). Crushed leaves
used as wound dressing (Teklehaymanot et al. 2007). The root juice is taken orally
for diarrhea and stomach-ache (Teklehaymanot 2009).
Brucea antidysenterica: In Eritrea, the wood is used as firewood and for roof
construction. The fruits, leaves, and twigs are also used in ethnoveterinary medicine
on skin galls and sores, and the powdered leaves to relieve bloating and colic in
cattle. The roots are used to treat rabies (Kokwaro 2009). The leaves are eaten by
cattle and goats. The timber serves as firewood, and a decoction of the bark is used as
arrow-poison (Bussmann 2006). Smoke smells bad and is used as insect repellent
Brucea antidysenterica J.F. Mill. 235
(Bussmann et al. 2011; Karunamoorthi et al. 2009). Root tied around the neck as
amulet against evil eye (Giday et al. 2007).
References
and Amhara proples in northwest Ethipopia. J Ethnopharmacol. 2007;110(3):516–25.
Karunamoorthi K, Mulelam A, Wassie F. Assessment of knowledge and usage custom of traditional
insect/mosquito repellent plants in Addis Zemen Town, South Gonder, North Western Ethiopia.
Stannard B. Flora of Tropical East Africa: Simaroubaceae. Rotterdam: Balkema; 2000.
Buddleja polystachya Fresen.
SCROPHULARIACEAE

Grace N. Njoroge
Local Names
Buddleja polystachya: Kikuyu: Ruti, Muthimbari, Muchorowe, Maa: Ol-biran,

Marakwet: Musereti, Gelelwa, Nandi: Chorenet, Marakwet / Nandi: Choruet,
Chorua, Ndorobo: Pinet, Samburu: Ngurangura (Beentje 1994; Gachati 1989;
Kokwaro 2009).
Botany and Ecology
Buddleja polystachya Fresen.: Robust shrub, up to 5 m tall, with pale brown bark,
young parts densely flocculent, glabrescent. Leaves shortly petiolate, petiole up to
5 mm long, lamina up to 16 cm long, 4 cm wide, lanceolate, gradually narrowing
above to an acute apex, convergent below, serrate at least in part. Inflorescence a
compound raceme of short sub-glomerulate clusters, lateral racemes up to 15 cm
long and less than 2 cm in diameter, pedicels obscure. Corolla reddish-orange,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Buddleja coriacea

(Scrophulariaceae),
Uchumarca, Peru. (Photo
R.W. Bussmann &
corolla-lobes almost square, with rounded apex. Anthers subsessile, inserted just
within the throat. Ovary narrowly ovoid, shortly hirsute (Bruce and Lewis 1960)
(Figs. 1, 2, 3, 4, 5, 6, 7, and 8).
Buddleja polystachya: A root decoction with fat is taken for sterility in women. The
leaf juice is applied for eye problems (Kokwaro 2009). Used for a variety of ailments
(Bekalo et al. 2009). The species has antimicrobial properties (Geyid et al. 2005).
The fresh leaves of Buddleja americana are used to treat headache and eyesight
(Béjar et al. 2002; Bussmann and Sharon 2006a, 2007a). Buddleja coriacea: Stems
and leaves are used to treat rheumatism, childbed infections, uterine cysts, kidney
infection, and prostate (Bussmann et al. 2016). Buddleja utilis: Used for menstrual
problems, inflammation of womb and ovaries, ovarian cysts, and uterus inflamma-
tion (Bussmann and Sharon 2006b, 2007b; 2015a, b; Monigatti et al. 2013). The
species shows mild antibacterial activity (Bussmann et al. 2009). Buddleja asiatica
is used as abortificant in India (Bhat et al. 2013, 2015).
Buddleja polystachya Fresen. 239

(Scrophulariaceae),
R.W. Bussmann &

(Scrophulariaceae),
R.W. Bussmann &
Fig. 4 Buddleja incana

(Scrophulariaceae),
R.W. Bussmann &

(Scrophulariaceae),
R.W. Bussmann &

(Scrophulariaceae),
R.W. Bussmann &
Buddleja polystachya Fresen. 241
Fig. 7 Buddleja polystachya

(Scrophulariaceae), Bale
Fig. 8 Buddleja polystachya

(Scrophulariaceae), Bale
Local Food Uses
Buddleja alternifolia is used as ornamental plant (Wu and Raven 1996).
Buddleja polystachya: Used as firewood (Bussmann et al. 2011). Buddleja davidii

serves as live fence (Mekonnen et al. 2015).
References
Béjar E, Bussmann RW, Roa C, Sharon D. Medicinal herbs of southern Ecuador – Hierbas
Medicinales del Sur Ecuatoriano. San Diego: Latino Herbal Press; 2002, 340p.
doi.org/10.1186/1746-4269-5-26.
in Kedarnath Wildlife Sanctuary of Garhwal Himalaya, India. J Ethnobiol Ethnomed 2013;9:1.
Bhat J, Malik ZA, Ballabha R, Bussmann RW, Bhatt AB. Ethnomedicinal plants traditionally used in
Bussmann RW, Sharon D. Traditional plant use in Loja province, southern Ecuador. J Ethnobiol
9789962-2-2.
Bussmann RW, Sharon D. Plants of the four winds – the magic and medicinal flora of Peru. Plantas
de los cuatro vientos – La flora mágica y medicinal del Perú. Honolulu: Arogya; 2007b. ISBN
978-0-9789962-3-9.
Bussmann RW, Sharon D, Garcia M. From chamomile to aspirin? Medicinal plant use among
clients at Laboratorios Beal in Trujillo, Peru. Ethnobot Res Appl. 2009;7:399–407.
medicinal flora of Northern Peru. William L. Brown Center, MBG, St. Louis; 2015a. ISBN
978-0-9960231-2-2.
Bussmann RW, Sharon D. Plantas medicinales de los Andes y la Amazonía – La flora mágica y
medicinal del Norte de Peru. William L. Brown Center, MBG, St. Louis; 2015b. ISBN 978-0-
9960231-3-9.
Bussmann RW, Paniagua Zambrana NY, Moya Huanca LA, Hart RE. Changing markets – medic-
inal plants in the markets of La Paz and El Alto, Bolivia. J Ethnopharmacol. 2016;193:76–95.
Mekonnen T, Giday M, Kelbessa K. Ethnobotanical study of homegarden plants in Sebeta-Awas
(2):450–64.
Wu ZY, Raven PH, editors. Flora of China. Volume 15, Myrsinaceae through Loganiaceae. Beijing/
St. Louis: Science Press/Missouri Botanical Garden Press; 1996. 387 pp.
Caesalpinia volkensii Harms
FABACEAE

Grace N. Njoroge
Local Names
Caesalpinia volkensii: Kikuyu: Mubuthi, Muchuthi; Digo: Mburuga; Maa:

Olnkulankulei; Watembe: Omukbare; Swahili: Mkomwe; Shamba: Msoo Miba
(Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Caesalpinia volkensii Harms: Climbing bush or liane. Bracts only shortly exceed-
ing buds, ovate-acuminate, ascending or rarely somewhat spreading in upper part,
4–5( 7) mm long. Racemes axillary, pedunculate, simple or with a few branches
below. Leaves: petiole with rachis 14–50 cm long, armed, pinnae 3–6 pairs,
6.5–16 cm long, leaflets 3–7 pairs per pinna, ovate to ovate-elliptic, (2–)3.2–8 cm
long, (1–)1.6–4.3 cm wide, acuminate or at the obtuse to subacute apex, glabrous or
nearly so, except when very young and for some persistent pubescence on midrib.
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Caesalpinia sp. Bale

Demaro forest. (Photo
R.W. Bussmann)
Pods broadly oblong-to obovate-elliptic, (6–)7–13 cm long, 3.6–6.5 cm wide,

densely spreading, prickly outside. Stems shortly pubescent and also armed with
scattered downwardly hooked or deflexed prickles 2–4 mm long. Seeds subglobose,
hard, brown, 1.7–2.3 cm in diameter, cuticle of testa finely and transversely cracked.
Stipules of 2–3 min subulate inconspicuous points 3 mm long. Petals yellow, about
16 mm long and 3.5–4.5 mm wide. Anthers 1.5–1.75 mm long. Pedicels 4–14 mm long.
Sepals 10–14 mm long, 3.5–5.5 mm wide (Brenan 1967). Important shrubs in the
undergrowth of moist Afromontane forests (Bussmann 2002) (Figs. 1, 2, 3, 4, and 5).
Caesalpinia volkensii: Decoction is used for malaria (Beentje 1994; Kokwaro

2009). Leaf decoction is also applied for pregnancy pain and stomach problems.
Roots chewed or boiled as aphrodisiac. Buds are crushed and applied for eye
problems (Kokwaro 2009). Used as antimalarial (Njoroge and Bussmann 2006a).
The root sap is inhaled in the nostrils to treat nose and throat problems (Njoroge and
Bussmann 2006c, 2009).
Caesalpinia volkensii: The leaves are used in ethnoveterinary medicine (Njoroge

and Bussmann 2006b).
Caesalpinia sp.: The wood is used for tools, especially plows, and also as firewood
Caesalpinia volkensii Harms 245

R.W. Bussmann)

R.W. Bussmann)

R.W. Bussmann)

R.W. Bussmann)
References
Brenan JPM. Flora of Tropical East Africa: Leguminosae, subfamily Caesalpinioideae. London:
Crown Agents for Overseas Governments & Administrations; 1967. 230 pp. ISBN 978-1-
84246-275-1 [Fl. Trop. E. Africa, Legum. (Part 2)].
Caesalpinia volkensii Harms 247

Njoroge, G.N., Ethnobotany 2009. Ethnotherapeutic management of Sexually Transmitted Diseases
STDs and reproductive health conditions in Central Province, Kenya. Indian J Tradit Knowl
82, 262–269.
Njoroge GN, Bussmann RW. Herbal usage and informant consensus in ethnoveterinary
management of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol.
2006b;108:332–9.
Central Kenya. J Ethnobiol Ethnomed. 2006c;2:54.
Calodendrum capense (L. f.) Thunb.
RUTACEAE

Grace N. Njoroge
Local Names
Calodendrum capense: Kikuyu: Murarachii, Muroroa, Mukarage; Kamba: Yangu;

Kipsigis: Kipkaria, Sasuriet; Maa: Ol-Larashi; Meru: Mutimueru, Mujai;
Samburu: Larachi; Taita: Mogorusi (Beentje 1994; Gachati 1989).
Botany and Ecology
Calodendrum capense (L. f.) Thunb.: Tree up to 20 m high, young branches, leaves
and inflorescences densely tomentose with simple, stellate and branched hairs, soon
becoming glabrous. Stamens as long as the petals, anthers 1.3–1.5 mm long,
staminodes longer than the petals, very narrowly oblanceolate, caudate, dotted with
bright crimson glands. Gynophore 4–6 mm long, glandular, ovary 2–3 mm in diam-
eter, style 15–20 mm long. Petals narrowly oblanceolate, 20–35 mm long, 5–7 mm
broad, whitish or mauve, straight or somewhat recurved, tomentose with simple or
R. W. Bussmann (*)
G. N. Njoroge

branched hairs on the outside and simple hairs directed downwards inside. Sepals
ovate, 3–5 mm long, 1.8 mm broad, free or slightly connate at the base, minutely
tomentose. Flowers mauve, bracteoles linear, 1.3–1.5 mm long, caducous, pedicel up
to 20 mm long. Inflorescence up to 15 cm long, with opposite cymose branches. Leaf-
lamina elliptic, 6–14( 18) cm long, 3.5–8( 11) cm broad, acute or slightly apiculate
or rounded at the apex, cuneate to slightly cordate at the base, petiole 2–10 mm long.
Seeds black, semi-spherical to pyramid-like, 10–15 mm in diameter. Fruit 5-lobed,
strongly rugose-warty, 35 mm in diameter, dehiscing from below with the 5 valves
remaining attached to the top of a central column (Kokwaro 1982). Found in the lower
and drier part of Afromontane forests (Bussmann 2002a, b). Easy in germination under
both light and dark conditions (Bussmann and Lange 2000).
Calodendrum capense: The bark is used as emetic, especially in cases of upset

stomach (Muthee et al. 2011).
Calodendrum capense: Wood is used for construction, tool handles, and utensils.
Often planted as an ornamental (Beentje 1994).
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002a;91 1/2:27–79,
Nat Hist. 2000;89(1–2):101–11.
Kokwaro JO. Flora of tropical East Africa: Rutaceae. Rotterdam: Balkema; 1982.
Capparis tomentosa Lam.
CAPPARIDACEAE
Gynandropsis gynandra (L.) Briq.
CLEOMACEAE

Grace N. Njoroge
Synonyms
Capparis tomentosa Lam.: Capparis corymbifera E. Mey. ex Sond.

Gynandropsis gynandra (L.) Briq.: Cleome gynandra L.; Cleome heterotricha
Burch.; Cleome pentaphylla L.; Cleome pentaphulla Willd.; Gynandropsis hetero-
tricha (Burch.) DC.; Gynandropsis pentaphylla (L.) DC.; Gynandropsis pentaphylla
Blanco; Gynandropsis sinica Miq.; Pedicellaria pentaphylla (L.) Schrank.
Local Names
Capparis tomentosa: Swahili: Mbada Paka; Borana: Gorrah-Gel; Giriama:

Mzezagunga; Ilelewa: Gora-nyilo, Gora za jovu, Namwaliko; Kamba: Kitamdanboo,
Mutandambo; Luhya: Shikumbumbu; Luo: Ong’ono; Maa: Olaturdei; Pokomo:
Muchovi; Pokot: Tipuyuo; Somali: Gombor Lik; Taita: Wangombei; Turkana:
Ekorokorait; Seguha: Dunguranassa; Chagga: Iravu (Gachati 1989; Beentje 1994;
Kokwaro 2009); English: Woolly caper-bush, African caper
R. W. Bussmann (*)
G. N. Njoroge

Gynandropsis gynandra: English: Spiderplant, Cat’s whiskers, Spider flower, Bas-

tard mustard; Swahili: Mgagani, Mkabili, Mkabilishemsi, Mwangani mgange
Botany and Ecology
Capparis tomentosa Lam.: A spiny, erect or scrambling shrub 1–3 m tall or climbing
over other vegetation to 6 m or perhaps more, young branches covered with a dense,
grayish or brownish pubescence, shorter side-branches often at right angles to the main
stems. Leaf-lamina 3–6 1.5–3 cm, oblong or elliptic, rounded and mucronulate or
slightly emarginate at the apex, rounded or broadly cuneate at the base, tomentose,
thinly hairy or glabrescent above, tomentose below, petioles 5–10 mm long, densely
pubescent. Flowers scented, in terminal corymbs of 3–15 flowers, the lower pedicels
solitary in the axils of the upper leaves, the upper with linear tomentose bracts 3 mm
long, pedicels up to 3 cm long, densely pubescent. Buds globose. Sepals 8–10 mm
long, orbicular, with a caducous grayish or brownish tomentum outside. Petals pale
yellowish-green, spathulate or cuneiform-truncate at the apex, with white hairs at the
base within. Stamens up to 80 with white or pinkish slender filaments up to 4 cm long.
Gynophore up to 4 cm long, slender. Ovary ovoid, bluntly apiculate at the apex. Fruit
up to 5 cm in diameter, globose, reddish-orange, on a very stout gynophore. Seeds
1.4 0.8 cm when mature, ovoid, smooth, embedded in a pinkish flesh. Capparis
tomentosa is widespread, from Mauritania, Senegal and Gambia eastward to Eritrea
and Ethiopia, and southward to north-eastern South Africa and Swaziland. It also
occurs in southern Arabia (Elffers et al. 1964). In disturbed areas in dry Afromontane
forests (Bussmann 2002; Figs. 1, 2, 3, 4, 5, 6, and 7).
Gynandropsis gynandra (L.) Briq.: Glandular-puberulous or glabrescent annual
herb 60 cm tall, sometimes becoming rather shrubby below. Leaves 5-foliolate or
3-foliolate above, leaflets 3–10 1–4.5 cm, narrowly obovate or oblanceolate,
rounded or acute to acuminate at the apex, cuneate at the base, repand-denticulate
or entire, glabrescent or glabrous, petioles up to 12 cm long, petiolules 2 mm long.
Inflorescence of terminal racemes with sessile or subsessile foliaceous bracts
3-foliolate below simple above, pedicels 1–2 cm long. Sepals 2–4 mm long,
lanceolate to narrowly ovate, acute at the apex, glandular. Petals white or rose-
pink, 1–2 0.3–0.5 cm, with an oblong blade, rounded at the apex and narrowing
abruptly into a claw rather longer than the lamina. Androgynophore 2 cm long,
purplish or pinkish, filaments very slender, purplish or pinkish, 2.5–4 cm long,
anthers 2.5 mm long, linear-oblong, gynophore 0.5–2 cm long, ovary linear-oblong,
glandular, style 1–2 mm long, glabrous, stigma capitate. Capsule 3–15 0.4–
1.2 cm, linear or linear-oblong, compressed, with many longitudinally anastomosing
nerves, glandular or glabrescent, terminated by the persistent style. Seeds brown,
numerous, 1.5 mm in diameter, rugose-reticulate. The origin of Cleome gynandra is
not known. There are claims that it has a southern Asian origin, but others suggest
that it originates from Africa or Central America. Cleome gynandra occurs through-
out the tropics and subtropics. In Africa, it is mainly found near human settlements,
possibly escapes from earlier introductions. It occurs probably in all countries of
tropical Africa (Elffers et al. 1964).
Capparis tomentosa Lam. . . . 253
Fig. 1 Capparis scabrida (Capparaceae), Huanchaco, Peru. (Photo R.W. Bussmann and
Fig. 2 Capparis scabrida

(Capparaceae), Huanchaco,
Peru. (Photo R.W. Bussmann
and N.Y. Paniagua-Zambrana)
Capparis tomentosa: Toxic but widely used in traditional medicine throughout the
distribution area of the species. Root powder and root decoctions are taken to treat
rheumatism, colds, cough, tuberculosis, apoplexy, stomach complaints, diarrhea,


rectal prolapse, schistosomiasis, gonorrhea, syphilis, leprosy, endometritis, menor-

rhagia, sterility and threatened abortion, and as anodyne, diuretic, anthelmintic, and
poison antidote. Also used as emetic, for abdominal pain, cough, and chest pains.
Must be used very carefully because the plant is highly toxic (Beentje 1994;


Kokwaro 2009). Grounded or powdered roots are applied to wounds, swellings,

abscesses, and snakebites, and to treat headache, migraine, skin diseases, and eye
complaints including conjunctivitis, ophthalmia, and cataract. Root decoctions are
added to baths against hemorrhoids. The smoke of burning bark powder is inhaled to

treat tuberculosis, bronchitis, and chest pain. Leaf decoctions are applied to treat skin
diseases and wounds, and are taken to treat hepatitis, malaria, convulsions, angina,
venereal diseases, asthma, and leprosy. Root macerations are given to cattle to treat
diarrhea, whereas a paste made of root ash is applied to udder inflammations. The
roots have been used in the preparation of arrow poison and in meat to poison
animals. Capparis tomentosa is one of the best-known woody species with magico-
medicinal properties, and it is commonly used in ritual ceremonies. Leaf decoction is
used for asthma (Kokwaro 2009). The smoke is used to treat respiratory disorders
(Mohagheghzadeh and Faridi 2006). The roots are burnt to treat postpartum bleeding
and tied around the neck for epilepsy (Giday et al. 2009). The plant is applied
externally for skin diseases (Giday et al. 2003). Chewed leaves are applied to
wounds and burns (Bussmann 2006). Capparis fascicularis is used for toothache
(Wondimu et al. 2007). The stems are also used to treat wounds and external injuries,
and the roots are used to treat stomachache (Teklehaymanot and MirMutse 2010).
Capparis erythrocarpos is applied to wounds (Giday et al. 2010). Capparis spinosa:
A decoction of the floral buds is used three times a day for 15 days to treat typhoid
fever and malaria. The vegetable made of the floral buds is considered one of the
famous gifts of Chitrali people to the rest the country (Bussamnn et al. 2020; Sher
et al. 2016). Capparis sicula: The roots are in folk medicine used against rheuma-
tisms and brucellosis. Fruits are used as compress for lumbago and sciatica. In
Armenian folk medicine, a decoction of the fruit is used for diseases of the thyroid,
goiter, hemorrhoids, scurvy, and against dental pain. Fresh roots are mashed and a
thick mass is applied as poultice for relief of pains and treatment in rheumatism, also
for massaging ill joints and. Immature fruits are used in the same way, mixed with
the fruits of walnut and henna. An infusion of leaves and root is used for bathing of
children with rheumatism. The leaves are prepared as water infusion, and the
decoction is used for the treatment of eczema, wounds, and ulcers. An ointment is
prepared from flowers and rhizome and applied for wound healing. Internal diseases:
The fruits of capers are used in diseases of the heart, thyroid gland, liver, lien, and
gastrointestinal tract, either independently or as infusion in grape vinegar. Leaves
with twigs, together with cherry are marinated in salted water and can be stored and
used for reducing blood sugar in diabetes. Seeds in a water decoction are applied as
antimalarial. An infusion and decoction of leaves, shoots, and roots, as well as fruits,
and their juice, are used as analeptic. The infusion of twigs with leaves with addition
of sour cherry and fennel are used in the treatment of jaundice. The water decoction
of the root is used as laxative. The fresh of the fruits or mature marinated fruits are
used to internally to treat hemorrhoids (Mehdiyeva et al. 2017). Capparis scabrida
fruits are used fresh for inflammation (general), heart palpitation, refresh the liver,
reduces anxiety, also good to increase milk production in cows (Bussmann and
Sharon 2006, 2007, 2015a, b; Bussmann et al. 2010a, 2011a, 2020; Paniagua-
Zambrana and Bussmann 2020; Rodriguez-Rodriguez et al. 2007). The species is
also used for respiratory disorders (Bussmann and Glenn 2010, 2011). Capparis has
slight antibacterial activity (Bussmann et al. 2008, 2009, 2010b, 2011b). Both
species show toxicity (Bussmann et al. 2011b).
Gynandropsis gynandra: Boiled spider-plant leaves are traditionally given to
mothers before and after delivery of a child, and in other situations where blood
has been lost, e.g., to warriors. Similarly, an infusion of the leaves is used to treat
anemia. The leaves and seeds are used medicinally as rubefacient and vesicant, and
to treat rheumatism, externally as well as internally. An infusion of the roots is used
as a medicine for chest pain and the leaves to treat diarrhea.
Local Food Uses
Capparis tomentosa: The leaves are sometimes eaten as a vegetable after cooking in
times of food shortage. The fruits can also be eaten (Bussmann 2006). Capparis
sicula: Buds in salt marinade are used as flavoring agent and appetizers. The young
fruits are used fresh, salted, or dried. The plant is known as spicy seasoning.
Marinated young shoots, buds, and fruits are used in food (Mehdiyeva et al. 2017).
Gynandropsis gynandra: The young leaves, young shoots, and occasionally flowers
are eaten boiled as potherb, relish, stew, or side dish. The leaves are utilized in fresh
form or dried as powder. Sometimes the leaves are bitter and then cooked with milk
and/or with other leafy vegetables such as cowpea leaves, amaranth, nightshades
(Solanum spp.), and Cleome monophylla L. In other areas, the leaves are boiled and
the cooking water is discarded. In several countries, pounded groundnut paste
(peanut butter) is added to improve the flavor. The leaves may be blanched, made
into small balls and sun- or air-dried. This is a popular product in southern Africa,
which finds a ready market when available during the rainy season. These balls or
leaf powder can be stored up to a year and are soaked in water before being used in
cooking. The seeds may be used as a substitute for mustard.
Capparis tomentosa: The foliage is browsed by camels and goats, although it has been
recorded as toxic to most livestock. The fruit has been reported to be edible and even
popular with children, but also to be poisonous to man and many animals. The press-
cake made from the seeds is fed to cattle. Stems and branches are used as firewood.
Cut branches are sometimes planted to make life fences. It is also a decorative plant for
gardens. Also sued to prevent evil eye (Teklehaymanot et al. 2007; Teklehaymanot
2009) and given to livestock against anthrax (Teklehaymanot 2009). The species is
eaten by livestock and used for construction and fences (Bussmann 2006). Capparis
fascicularis is poisonous (Bussmann et al. 2006). Capparis erythrocarpos smoke is
also used for evil eye (Giday et al. 2010). Capparis scabrida is widely used to make
utensils (Rodriguez-Rodriguez et al. 2007). Capparis sicula: Roots are used in
traditional handcrafts as source of dark green dye for silk. Black color is obtained
from the root, dark green from stems, for dyeing silk and wool. The shrub can be
planted on slopes along the road to prevent erosion (Mehdiyeva et al. 2017).
Gynandropsis gynandra: Seeds thrown in water can kill fish, which then float to the
surface. The glands on the stems and leaves have insect repellent properties, cabbage
and related crops intercropped with spiderplant suffer less from diamond back moth
larvae. Similarly, in French bean intercropped with spiderplant, the beans are less
affected by flower thrips and are therefore of better quality for export. The seeds are
used to feed birds. The seed contain an edible polyunsaturated oil, which is extracted
by simple pressing and does not need refining. The seed cake can be used as
animal food.
References
Bussmann RW, Glenn A. Traditional Peruvian medicine for the treatment of respiratory disorders.
Revista Peruana de Biologia. 2010;17(2):331–46.
Bussmann RW, Glenn A. Fighting pain. Traditional Peruvian remedies for the treatment of asthma,
rheumatism, arthritis and sore bones. Indian J Tradit Knowl. 2011;10(3):397–412.
Bussmann RW, Sharon D. Plants of the four winds – The magic and medicinal flora of Peru. Plantas
de los cuatro vientos – La flora mágica y medicinal del Perú. Honolulu: Arogya; 2007. ISBN
978-0-9789962-3-9.
medicinal flora of northern Peru. St. Louis: William L. Brown Center, MBG; 2015a. ISBN
978-0-9960231-2-2.
medicinal del Norte de Peru. St. Louis: William L. Brown Center, MBG; 2015b. ISBN 978-0-
9960231-3-9.
Bussmann RW, Sharon D, Perez F, Díaz D, Ford T, Rasheed T, Silva R. Antibacterial activity of
northern-Peruvian medicinal plants – a low cost laboratory approach to assess biological
activity. Arnaldoa. 2008;15(1):127–48.
Bussmann RW, Sharon D, Diaz D, Cardenas R, Chait G, Castro M, Regalado S, Del Toro CR,
Malca GG, Perez AF, Glenn A. Antibacterial activity of medicinal plant species in Northern
Peru. Arnaldoa. 2009;16(1):93–103.
Bussmann RW, Sharon D, Glenn A. Healing the body, healing the soul. Traditional remedies for
“magical” ailments, nervous system and psychosomatic disorders in northern Peru. Afr J Pharm
Pharmacol. 2010a;4(9):580–629.
Bussmann RW, Glenn A, Sharon D. Antibacterial activity of medicinal plants of northern Peru –
can traditional applications provide leads for modern science? Indian J Tradit Med. 2010b;9
(4):742–53.
northern Peruvian medicinal plants. Ethnobot Res Appl. 2011a;9:67–96.
Bussmann RW, Malca G, Glenn A, Sharon D, Nilsen B, Parris B, Dubose D, Ruiz D, Saleda J,
Martinez M, Carillo L, Walker K, Kuhlman A, Townesmith A. Toxicity of medicinal plants used
in northern Peru. J Ethnopharmacol. 2011b;137:121–40.
Bussmann RW, Batsatsashvili K, Kikvidze Z. Capparis spinosa L. In: Batsatsashvili K, Kikvidze Z,
Bussmann RW, editors. Ethnobotany of mountain regions Central Asia and Altai. Cham:
Springer International Publishing International Publishing; 2020.
Elffers J, Graham RA, DeWolf GP. Flora of tropical East Africa: Capparidaceae. London: Crown
Giday M, Asfaw Z, Woldu Z, Teklehaymanot T. Medicinal plant knowledge of the bench ethnic
doi.org/10.1186/1746-4269-5-34.
Mehdiyeva N, Alizade V, Fayvush G, Aleksanyan A, Paniagua Zambrana NY, Bussmann
RW. Capparis sicula Vieill. subsp. herbacea Willd. Inocencio, D. Rivera, Obón & Alcaraz.
In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer International Publish-
ing; 2017.
Paniagua Zambrana NY, Bussmann RW. Capparis crotonoides Kunth Iltis & Cornejo, Capparis
scabrida Kunth. In: Paniagua Zambrana NY, Bussmann RW, editors. Ethnobotany of Mountain
Regions – Ethnobotany of the Andes. Cham: Springer International Publishing; 2020.
Rodriguez-Rodriguez EF, Bussmann RW, Arroyo Alfaro SJ, López Medina SE, Briceño Rosario
J. Capparis scabrida (Capparaceae) una especie del Norte del Perú y Sur del Ecuador que
necesita conservación urgente. Arnaldoa. 2007;14(2):269–82.
Teklehaymanot T, MirMutse G. Quantitative ethnobotany of medicinal plants used by Kara and

Kwego semi-pastoralist people in lower Omo River valley, Debub Omo zone, southern nations,
nationalities and peoples regional state, Ethiopia. J Ethnopharmacol. 2010;130:76–84.
Carissa spinarum L.
APOCYNACEAE

Grace N. Njoroge
Synonyms
Carissa spinarum L.: Anthura edulis Forssk.; Anthura hadiensis J.F. Gmel.;
Arduina edulis (Forssk.) Spreng.; Carandas edulis (Forssk.) Hiern; Carissa
abyssinica R. Br.; Carissa africana A. DC.; Carissa candolleana Jaub. & Spach.;
Carissa carandas var. congesta (Wight) Bedd.; Carissa carandas var. paucinervia
(A. DC.) Bedd.; Carissa cochinchinensis Pierre ex Pit.; Carissa congesta Wight.;
Carissa cornifolia Jaub. & Spach.; Carissa dulcis Schumach. & Thonn.; Carissa
edulis L.; Carissa paucinervia A. DC.; Carissa pilosa Schinz; Carissa pubescens
A. DC.; Carissa richardiana Jaub. & Spach; Carissa tomentosa A. Rich.;
Jasminonerium africanum (A. DC.) Kuntze; Jasminonerium densiflorum (Baker)
Kuntze; Jasminonerium dulce (Schumach. & Thonn.) Kuntze; Jasminonerium edule
(Forssk.) Kuntze; Jasminonerium madagascariense (Thouars ex Poir.) Kuntze;
Jasminonerium pubescens (A. DC.) Kuntze; Jasminonerium sechellense (Baker)
Kuntze; Jasminonerium tomentosum (A. Rich.) Kuntze
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Carissa edulis: Kikuyu: Mukawa; Swahili: Mtanda-mboo; Borana: Fonkole,

Dagams; Boni: Mulimuli; Duruma: Mulowe, Mulolwe; Gabbra: Dagamsa;
Ilelewa: Kaka-mchangani; Kamba: Kikawam, Mukawa; Luo: Ochuoga; Maa:
Olamuriaki; Marakwet/Tugen: Legatetwo; Nandi/Kipsigit/Tugen: Legetetwa,
Legetetwet; Pokot: Lokotetwo; Samburu: Lmuria, Lmuriel, Lamurei; Sanya:
Gurura; Taita: Kirumba; Turkana: Ekamuria; Myamuha: Kamasai; Hehe:
Mufumbwe; Luganda: Muyonza (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Carissa spinarum: L.: Much-branched spreading or sarmentose shrub up to 6 m tall

or more. Young twigs densely pubescent or puberulent, rarely glabrous, the surface
smooth or very finely rugose, older branches with grayish-brown flaking bark.
Spines simple, very rarely furcate, 0.4–5.5(7) cm long. Leaves moderately coria-
ceous, drying discolorous, brown or green, darker above, petiole 2.5–5 mm long,
pubescent or glabrous. Lamina (1.7)2.6–6.8 (1)1.2–4.6 cm, ovate, elliptic,
obovate-elliptic or subcircular, with apex acute or obtuse, with or without mucro,
and base cuneate to rounded, upper surface glabrous to pubescent, midrib slightly
impressed, other veins obscure, lower surface glabrous to pubescent (hairs most
dense on midrib, here sometimes almost lanate), midrib and other veins slightly
raised, venation easily visible or obscure. Cymes short-stalked, few- to many-
flowered, flowers 5 (occasionally 6)-merous, sweetly scented, the corolla white
inside and pink to crimson outside. Calyx 2–4 mm long, lobes lanceolate-subulate,
glabrous, pubescent or lanate. Corolla tube 9.5–20 mm long, slightly wider in upper
half and contracted at the mouth, glabrous or sparsely pilose without, pubescent
within in upper half, corolla lobes 1/3–1/2 as long as tube, lanceolate, glabrous or
sparsely puberulent near the mouth on upper surface, overlapping to the right.
Stamens inserted near apex of tube so that the anthers reach to within 1–1.5 mm
of the mouth, anthers subsessile, 1.5–2 mm long. Ovary 1 mm long, rounded-
conical, glabrous, style glabrous, stigma reaching to base of anthers. Fruit 6–11
(25) mm long, plum-shaped, green turning red to blue-black, edible, 2–4-seeded.
(Goyder et al. 2012). In the understory of dry Afromontane forests (Bussmann 2002;
Figs. 1, 2, 3, and 4).
Carissa spinarum: Fruits sometimes eaten against diarrhea, root decoction used as
analgesic (Beentje 1994; Bekalo et al. 2009; Lulekal et al. 2008; Wondimu et al.
2007). Roots used for indigestion, lower abdominal pain, malaria, chest pain,
arthritis, rheumatism, gonorrhea, yellow fever, epilepsy, and kidney problems.
Leaf decoction applied for breast cancer (Bussmann 2006; Kokwaro 2009), as
Carissa spinarum L. 263
Fig. 1 Carissa spinarum

(Apocynaceae), Naro Moru,
Kenya. (Photo
R.W. Bussmann)

(Apocynaceae), buds and
flowers, Naro Moru, Kenya.
well as polio, tuberculosis, venereal diseases (Bussmann 2006; Bussmann et al.

2006), and back pain (Bussmann et al. 2006). Used for stomachache and women’s
health (Delbanco et al. 2017), digestion during pregnancy, pelvic pain, backache,
and sexually transmitted diseases (Nankaya et al. 2019). Used to treat snakebites
(Teklehaymanot et al. 2007). Sometimes the plant is used to treat malaria (Tabuti
2008). The roots are used as galactagogue, the stem is used for postpartum pain
(Njoroge and Bussmann 2009). Sometimes used as anti-malarial (Gafna et al. 2017;
Njoroge and Bussmann 2006a). The roots and leaves are used for asthma, common
cold, and chest pain (Njoroge and Bussmann 2006b). The plant is used externally for
rheumatism (Giday et al. 2003). The plant is burnt and the smoke is used against
postpartum bleeding and the leaves tied around the neck against evil eye (Giday et al.
2009). Smoke is also used to treat mood disorders (Mohagheghzadeh and Faridi
2006).

(Apocynaceae), mature
flowers, Naro Moru, Kenya.

(Apocynaceae), fruits, Naro
Moru, Kenya. (Photo
R.W. Bussmann)
Local Food Uses
Carissa spinarum: Fruits are edible (Beentje 1994; Bussmann 2006; Bussmann
et al. 2006, 2011; Luizza et al. 2013).
Carissa spinarum L. 265
Carissa spinarum: Browsed by goats (Bussmann et al. 2011). The root powder with
water is poured in the holes of snakes as repellent (Teklehaymanot 2009). The thorns
are used to pierce the ears (Luizza et al. 2013). Planted as life fence (Mekonnen et al.
2015).
References
doi.org/10.1186/1746-4269-5-26.
appendices 1–7, www.naturekenya.org/JournalEANH.htm.
Delbanco AS, Burgess ND, Cuni_Sanchez A. Medicinal plant trade in northern Kenya: economic
importance, uses, and origin. Econ Bot. 2017;711:13–31.
ethnobotan ical study. J Ethnopharmacol. 2009;124:513–21.
Goyder D, Harris T, Masinde S, Meve U, Venter J. Flora of tropical East Africa: Apocynaceae (part
2). Kew: Royal Botanic Gardens; 2012. p. 115–530. ISBN 978-1-84246-396-3 [Fl. Trop.
E. Africa, Apocyn. (Part 2)].
Res Appl. 2013;11:315–39.
10.1186/1746-4269-4-10.

Njoroge GN, Bussmann RW. Ethnotherapeutic management of sexually transmitted diseases
(STDs) and reproductive health conditions in Central Province, Kenya. Indian J Tradit
Knowl. 2009;8(2):262–9.
Omino EA. Flora of tropical East Africa: Apocynaceae (part 1). Rotterdam: Balkema; 2002.
114 p. ISBN 90-5809-409-X.
Casearia battiscombei R.E. Fr.
SALICACEAE

Grace N. Njoroge
Local Names
Casearia batiscombei: Kikuyu: Muirungi, Muthandi, Muirongi; Luhya:

Namasinzi; Marakwet: Liss; Meru: Munogo; Nandi: Cheptokoriet, Chapchabeyet;
Ndorobo: Chepcharaiyet; Pokot: Sigikawa; Samburu: Iltumbach; Sebei: Seruwa,
Seruondet; Okiek: Poponit (Beentje 1994; Gachati 1989).
Botany and Ecology
Casearia battiscombei R.E. Fr.: Tall tree up to about 40 m tall with rough grayish
bark, young branches brown (or silvery in saplings), puberulous or glabrescent.
Leaves on puberulous or glabrescent petioles up to 6 mm long, leaf-lamina
8–22 3–6 cm, papyraceous, narrowly oblong to oblong, shortly acuminate or obtuse
at the apex, unequal-sided and rounded, slightly cordate or broadly cuneate at the base,
margin entire, undulate or rarely shallowly and irregularly crenulate (the leaves on
R. W. Bussmann (*)
G. N. Njoroge

young saplings often serrate) glabrous on both sides, with circular and linear pellucid
dots, lateral nerves in 12–20 pairs, prominent below, venation laxly reticulate, not at all
prominent, petiole up to 6 mm long, puberulous or glabrous, stipules up to 2 mm long,
triangular, scarious, very caducous. Flowers numerous, greenish, in axillary fascicles
borne on a cushion of minute, densely pubescent bracteoles, pedicels 3–4 mm long,
increasing to 1 cm in fruit, sparsely puberulous or glabrescent. Ovary ovoid, glabrous,
style very short, stigma capitate. Staminal tube exceedingly short, stamens 6–10 on
puberulous filaments 0.8–1 mm long, anthers about 0.5 0.3 mm, ovate-lanceolate,
minutely and sparsely puberulous or glabrescent, connective bluntly apiculate,
staminodes 0.5 mm long, oblong or subquadrate, densely hirsute. Receptacle about
2 mm wide at the mouth, about 1 mm tall, funnel-shaped, puberulous or glabrescent
outside. Sepals about 2 1.5 mm, broadly elliptic or rotund, obtuse at the apex,
margins membranous, puberulous or glabrescent outside. Fruit an ellipsoid capsule,
about 1.3 0.8 cm, splitting from above into 2–4 longitudinal valves, glabrous. Seeds
few, about 6 4.5 mm, ellipsoid, with a smooth pale testa, almost enclosed in a fleshy
aril. Common in Afromontane forests with dense Coffea layer (Bussmann 2002a, b).
Casearia battiscombei: Used as timber (Beentje 1994). Some species are used as
fish poison (Neuwinger 2004). Casearia nigrescens is used as construction timber in
Madagascar (Rakotoarivelo et al. 2013).
References
L. Brown Center, MBG; 2013. ISBN 978-0-9848415-7-8
Cassipourea malosana (Baker) Alston
RHIZOPHORACEAE

Grace N. Njoroge
Synonyms
Cassipourea malosana (Baker) Alston: Cassipourea abyssinica (Engl.) Alston;

Cassipourea avettae (Chiov.) Alston; Cassipourea eickii (Engl.) Alston;
Cassipourea elliottii (Engl.) Alston; Cassipourea gerrardii (Schinz) Alston; Weihea
abyssinica Engl.; Weihea avettae Chiov.; Weihea boranensis Chiov.; Weihea eickii
Engl.; Weihea elliottii Engl.; Weihea gerrardii Schinz; Weihea ilicifolia Brehmer;
Weihea malosana Baker; Weihea salvago-raggei Chiov.
Local Names
Cassipourea malosana: Kikuyu: Muthaithi; Kipsigis: Muangaita, Marakwet:

Tendewa; Meru: Muchanja; Nandi: Martit; Ndorobo/Kipsigit: Mangweta;
Samburu: Lobbobo; Maa: Oliami-Orok, Ol-lorget; Tugen: Maiti; Turkana: Elatha
(Beentje 1994; Gachati 1989); English: Pillarwood, Common onionwood, Bastard
onionwood
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Cassipourea malosana (Baker) Alston: Shrub or tree up to 25 m tall, young stem

appressed-pubescent to glabrescent (first year twigs sparsely hirsute), branches hairy
when young and glabrous with age, bark grayish-yellow, gray or brown. Leaf-lamina
3–10 1–5 cm, oblanceolate, lanceolate, narrowly to broadly elliptic, oblong, ovate
or obovate, apex subacute, obtuse or short- to long-acuminate (rarely acute, rounded
or retuse), margins serrate or rarely entire, base cuneate or obtuse, rarely rounded,
superior surface shining, glabrous, with conspicuous venation, inferior surface mat,
sparsely appressed-pubescent to glabrous (except on the midrib) and with more or
less prominent venation, petiole 2–7 mm long, pubescent to glabrous, stipules
truncate to subrounded at the base, abaxial surface densely appressed-pubescent to
pubescent at the apex only, adaxial surface glabrous, soon caducuous. Inflorescences
1–4(6)-flowered, lax to congested. Flowers 4–5(6)-merous, pedicels 2–5(9) mm
long, pubescent, articulate near the apex, bracteoles 1·5–2·5 mm long, conchiform,
rounded or lacerate at the apex and truncate at the base. Calyx slightly pubescent to
sericeous-pubescent outside and glabrous to slightly puberulous inside, calyx-tube
0·5–2 mm long, calyx-lobes 3–6 mm long, lanceolate, oblong or ovate-lanceolate.
Petals 4–8 mm long, spathulate, laciniate above, glabrous. Stamens 15–20, filaments
3–6 mm long, anthers 1 mm long, narrowly to broadly ellipsoid-oblong. Ovary
1–2 mm long, ovoid to spherical, 3–4-locular, glabrous to densely hirsute, style
Fig. 1 Cassipourea
malosana (Rhizophoraceae),
flowering branch underside,
R.W. Bussmann)
Cassipourea malosana (Baker) Alston 271
3–5 mm long, glabrous to pubescent mainly towards the base, persistent. Fruit
6–10 mm long, more or less ellipsoid, ovoid to subspherical, sericeous-pubescent
to glabrous. Cassipourea malosana occurs from eastern DR Congo and Ethiopia
south to South Africa. One of the slowest growing species in Afromontane forests
(Bussmann 1999). Characteristic component of broadleaves, semi-evergreen
Afromontane forests (Bussmann 2002a, b, 2004, 2006a; Figs. 1, 2, 3, 4, and 5).
Cassipourea malosana: The Loita Maasai in Kenya eat the outer bark boiled with
soup to gain strength. Tea is made from the scraped inner bark and drunk by women
to help to remove the placenta. In South Africa, the bark is used as a skin lightener
and to treat skin ailments and sunburn.
Cassipourea malosana: The flowers are a good source of nectar for bees. Also used
as a shade tree. The wood is traded as pillarwood. It is suitable for construction,
flooring (especially heavy duty or industrial flooring), vehicle bodies, furniture and
Fig. 2 Cassipourea
flowering branch upper side,
R.W. Bussmann)
Fig. 3 Cassipourea
flower, Mt. Kenya National
Fig. 4 Cassipourea
cabinet work, tool handles, ladders, sporting goods, agricultural implements, joinery,
sleepers, poles and piles, toys and novelties, beehives, turnery, and for veneer and
plywood. Its low durability, however, limits its applicability outdoors. Nevertheless,
Cassipourea malosana is an important timber tree in East Africa. The wood is used
for construction and also used as firewood (Beentje 1994). The leaves are eaten by
sheep and goats, and the wood is excellent for containers (Bussmann 2006b). Gives
good firewood (Bussmann et al. 2011). The bark is used to treat malaria (Njoroge
and Bussmann 2006). The root decoction is used externally to treat rheumatism
Cassipourea malosana (Baker) Alston 273
Fig. 5 Cassipourea
References
reference to those of Mount Kenya. J East Afr Nat Hist. 1999;88(1–2):69–78.
Bussmann RW. Succession and regeneration patterns in East African mountain forests – a review.
Bussmann RW. Regeneration and succession patterns in African, Andean and Pacific tropical
mountain forests: the role of natural and anthropogenic disturbance. Lyonia. 2004;61:98–111.
2006a;111:41–66.
Ethnomed. 2006;2:8.
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plants Res. 2008;
26:132–53.
Catha edulis Forssk.
CELASTRACEAE

Grace N. Njoroge
Synonyms
Catha edulis Forssk.: Catha edulis (Vahl.) Endl.; Celastrus edulis Vahl.
Local Names
Catha edulis: Kikuyu: Muirungi, Mailungi; Swahili: Mlonge, Miraa, Murungu;

Maa: Ol-Merra; Marakwet: Liss; Kipsigis/Ndorobo: Tomayot; Samburu:
Mamiraa; Somali: Kat; Shambaa: Muandama; Meru: Miraa, Muraa (Beentje
1994; Gachati 1989; Kokwaro 2009); English: Khat, Arabian tea, Abyssinian tea,
Bushman’s tea
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Catha edulis Forssk.: Tree 2–15( 25) m high, branches pale gray-green and
flattened when young, becoming vinous red, terete. Leaf-lamina dark green or
grayish green and glossy above, paler beneath, oblong to elliptic or obovate,
(3.7–)5.5–11 (0.8–)1.5–4.5( 6) cm, acute to acuminate or more rarely obtuse at
the apex, with margin markedly glandular-crenulate-denticulate, cuneate to attenuate
at the base, coriaceous or subcoriaceous, with densely reticulate venation more
prominent beneath than above, petiole 3–10 mm long. Cymes 1–2.3 cm long, rather
dense, with peduncle 6–12 mm long, pedicels very short or up to 2 mm long, flowers
many in each cyme, 2–3 mm in diameter. Sepals broadly ovate to semicircular,
rounded, 0.5–0.7 mm long, with margin ciliate-fimbriate. Petals elliptic-oblong,
1–1.5 mm long, sessile, with margin ciliolate and paler in color. Stamens shorter
than petals, with filaments 0.7–1 mm long. Disc 1–1.5 mm in diameter. Ovary
broadly ovoid, styles less than 0.5 mm long, slender, stigmas small. Capsule red,
narrowly oblong-3-gonous, 6–10 mm long, pendulous. Seeds dark brown, rugose-
papillose, with narrow wing 2–2.5 times as long. Evergreen submontane or medium
altitude forest, usually near the margins, or in woodland often on rocky hills,
1100–1800( 2400) m. Khat is indigenous to the evergreen montane forests in
eastern Africa, from Eritrea south to South Africa (Cape Province) and Swaziland.
The primary centre of origin is assumed to be in the south-western highlands of
Ethiopia. According to the fourteenth century Arabic chroniclers, khat was culti-
vated extensively in the mountains of Yemen and also near Harar in Ethiopia at that
time. It may have been introduced into Yemen from Ethiopia in the sixth century
AD, some 600 years earlier than coffee (Coffea arabica L.) but was not known to the
West until the end of the eighteenth century. Its regular use as a stimulant is confined
largely to Muslim communities of southern Arabia and eastern Africa. Yemen,
Ethiopia, and Kenya are the main khat growing countries, but it is also collected
from the wild or cultivated in several other eastern and southern African countries
and in Madagascar (Robson 1994; Fig. 1).
Catha edulis: In traditional African and Arabic medicine, the leaves and roots of
khat are considered a panacea against all sorts of ailments and diseases. Used mainly
as stimulant (Kokwaro 2009; Tabuti 2008). Root decoction is used for body illness.
Leaves and roots are boiled for flu. Roots alone are used for stomach problems. Root
and bark in decoction are used for gonorrhea (Kokwaro 2009). The leaf powder is
ingested to treat asthma and cough (Teklehaymanot et al. 2007). Sometimes the plant
is employed as an analgesic (Yonathan et al. 2006). Heavily cultivated in home
gardens and exported as medicinal plant from Ethiopia as an important cash crop
(Giday et al. 2003; Lulekal et al. 2008; Mekonnen et al. 2015).
Catha edulis Forssk. 277
Fig. 1 Catha edulis

(Celastraceae), Naro Moru,
Kenya. (Photo
R.W. Bussmann)
Fresh young leaves, and sometimes the tender shoot tips, are chewed for their
stimulating and mildly intoxicating effects (Beentje 1994). As more leaves are
added, a wad is formed which is stored in the cheek. This is slowly chewed until
all juices are extracted and the residue is expectorated or swallowed. Chewing is
accompanied by drinking large quantities of cold water and sometimes also by
smoking cigarettes or water pipe. Khat chewing is an age-old habit in rural areas
to alleviate fatigue during fieldwork or to enliven religious and family gatherings.
Occupational groups, such as drivers of motor vehicles, merchants, and students,
may use it for its energizing effect and to stay awake. In recent years, khat chewing
has become a major recreational activity during lengthy social gatherings, particu-
larly in Yemen, Ethiopia, Djibouti, and Somalia, resulting in a heavy strain on family
incomes and an appreciable loss of productive labor. Rapid transport by air enables
regular fresh supplies of khat even to emigrant communities from these countries
now resident in Europe and elsewhere in the world. Larger leaves that are too hard
for chewing and leaves that have lost their freshness may be dried and pulverized for
the preparation of a paste with water, sugar, or honey and sometimes also spices. The
paste is chewed and swallowed in a similar manner as described for fresh leaves.
Dried leaves are also used to prepare an infusion in the same way as tea, e.g., in
South Africa, or they may be smoked liked tobacco, e.g., in Arabic countries. When
left to grow into large trees, Catha edulis yields a fine timber for furniture and
building (Beentje 1994), called Chirinda redwood in southern Africa. The wood pulp
makes excellent blotting paper.
References
10.1186/1746-4269-4-10.
Robson N. Flora of tropical East Africa: Celastraceae. Rotterdam: Balkema; 1994.
Yonathan M, Asres K, Asse A, Bucar F. In vivo anti-inflammatory and anti-nociceptive activities of
Cheilanthes farinosa. J Ethnopharmacol. 2006;108:462–70.
Celtis africana Burm. f.
CANNABACEAE

Grace N. Njoroge
Synonyms
Celtis africana Burm. f.: Celtis durandii Engl.; Celtis durandii var. ugandensis
Rendle; Celtis gomphophylla Baker; Celtis kraussiana Burm. f.
Local Names
Celtis africana: Kikuyu/Meru: Murundu; Kipsigit: Chepkeleliet, Nyasiat; Luhya:

Mweyu, Mwenye; Maa: Olmositet; Nandi: Chepkeleriet; Ndorobo: Mase; Sebei:
Mastet, Mastitet, Mosididiet; Tugen: Mususetiet, Tandimu, Tandimwo; Turkana:
Eng’omwo (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Celtis africana Burm. f.: Deciduous tree, 5–35 m tall, with smooth gray bark and
often slight horizontal annular ridges (ring marked). Fruits on 10–25 mm long
pedicels, subglobose, 5 mm long, orange, pubescent. Cymes in the lower leaf axils
and at the nodes below of 3–15 clustered ♂ flowers with 1.5–4 mm long pedicels,
uppermost cymules of 1-several hermaphrodite flowers with long slender 10–17 mm
long pedicels, intermediate ones polygamous. Stipules linear, 4–6 mm long, pubes-
cent. Ovary densely hairy, styles unbranched, 2.5–4 mm long. Sepals 4–5, 1.5–3 mm
long, pubescent. Leaf-blades ovate to ovate-lanceolate, 5–10.2 cm long, 2–5.5 cm
wide, acuminate, rounded and a little unequal-sided at the base, serrate in the upper
two-thirds, pubescent, ultimately subglabrous except on the nerves beneath, sca-
brous, with the basal lateral nerves extending well into the upper half and the upper
prominent lateral nerves 1–2 on each side of the midrib, petiole 1–5 mm long. Young
twigs tawny pubescent to tomentose (Verdcourt 1975). The species is especially
common in Coffea arabica dominated forests in Southern Ethiopia and Northern
Kenya (Bussmann 2002a, b) (Figs. 1, 2, 3, 4, and 5).
Celtis africana: Used to treat rheumatism (Yineger et al. 2008).

Celtis africana and Celtis gomphophylla, are used as raw material for medicines in
Madagascar, especially for diarrhea and digestive system problems (Randrianarivony
et al. 2016, 2017).
Celtis australis: Used as blood purifier and tonic (Muhammad et al. 2019), as well as
against allergies (Ur-Rahman et al. 2018). Celtis caucasica: A decoction of fruits is
Fig. 1 Celtis africana

(Cannabaceae), Bale
Odo Bulu forest, Ethiopia.
Celtis africana Burm. f. 281
Fig. 2 Celtis africana.

(Cannabaceae), Bale
Fig. 3 Celtis africana.

(Cannabaceae), Bale
applied in acute gastric diseases and in diarrhea. The decoction of seed acts as
sudorific (Bussmann 2017; Batsatsashvili et al. 2017).
Local Food Uses
Celtis caucasica: Fruits are edible. Out of flesh is sweet; core of nucleus resembles
the taste of almond. Used in food in the raw form. Porridge is made from hard baked
and milled fruits in flour, and “fruit bread” is baked when 20–30% fruit flour is
mixed with simple flour. Bread with such mix has pleasant taste with high content of
vitamins (Bussmann 2017; Batsatsashvili et al. 2017).
Fig. 4 Celtis gomphophylla

(Cannabaceae), Bale
Harenna forest, Ethiopia.
Fig. 5 Celtis gomphophylla

(Cannabaceae), Bale
Harenna forest, Ethiopia.
Celtis africana: Wood is used for furniture, tool handles, and construction (Beentje
1994). Leaves are given to goats for indigestion (Kokwaro 2009). A good source of
firewood (Kiefer and Bussmann 2004, 2008).
Celtis mildbraedii is used as fish poison (Neuwinger 2004). Celtis caucasica:
Silkworm is fed with young leaves, and they are as well a good feed for cattle,
especially for goats. Suitable for construction of park and gardens in dry regions,
grows big tree with round crown in culture. A dye solution is prepared from leaves
Celtis africana Burm. f. 283
and barks to obtain green and yellow colors. Solution is used for dyeing of wool.
Bark is used for tanning. The wood is heavy firm, solid, durable, worm hole stable,
cleavable, thick, elastic, and well polishing. Highly applied in joinery, turning, and
carving; especially valued in vehicle production – wheel pub and wheel axis, shaft
and other details, wind instruments, cutting for knives, stick, ramrod, and so
on. Owing to its elasticity valued as material for production of hoopla hoops.
Nectareous, produces nectar and pollen. The plant can be used for strengthening
mountain slopes and rockslide especially in dry areas. Iconic, as per beliefs chaplet,
beads, and other items made of its wood protect against evil eye. Piece of wood and
bark is also used against evil eye, which is kept in visible place; object protected
against evil eye or hung on neck. The wood was traditionally used to make spindles
(Bussmann 2017; Batsatsashvili et al. 2017).
References
Batsatsashvili K, Mehdiyeva N, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S, Tchelidze D,
Alizade V, Paniagua Zambrana NY, Bussmann RW. Celtis caucasica Willd. In: Bussmann RW,
editor. Ethnobotany of the Caucasus. Cham: Springer International Publishing; 2017.
Bussmann, R.W. ed. 2017. Ethnobotany of the Caucasus. Springer International Publishing: Cham.
XXVII, 746p. ISBN 978-3-319-49411-1.
the forest expansion in east African montane forest ecosystems – a modeling approach. Lyonia.
2004;6(2):161–9.
appraisal of Kurram agency tribal area Pakistan. Indian J Tradit Knowl. 2019;18(4):631–47.
Randrianarivony RN, Andriamihajarivo TH, Ramarosandratana AV, Rakotoarivony F, Jeannoda
VH, Kuhlman A, Randrianasolo A, Bussmann RW. Value of useful goods and ecosystem
services from Agnalavelo sacred forest and their relationship with forest conservation. Madag
Conserv Dev. 2016;11(2):44–51.
Randrianarivony TN, Ramarosandratana VA, Andriamihajarivo TH, Rakotoarivony F, Jeannoda V,
Randrianasolo A, Bussmann RW. The most used medicinal plants by communities in
Mahaboboka, Amboronabo, Mikoboka, Southwestern Madagascar. J Ethnobiol Ethnomed.
2017. https://doi.org/10.1186/s13002-017-0147-x.
Verdcourt B. Flora of tropical East Africa: Cannabaceae. London: Crown Agents for Overseas
10.1186/1746-4269-4-11.
Cissampelos pareira L.
MENISPERMACEAE

Grace N. Njoroge
Synonyms
Cissampelos pareira L.: Cissampelos acuminata Benth.; Cissampelos acuminata

DC.; Cissampelos argentea Kunth; Cissampelos auriculata Miers; Cissampelos
australis A. St.-Hil.; Cissampelos benthamiana Miers; Cissampelos bovionii Baill.;
Cissampelos bojeriana Miers; Cissampelos caapeba L.; Cissampelos capeeba Roxb.;
Cissampelos canescens Miq.; Cissampelos cocculus Poir.; Cissampelos consociata
Miers; Cissampelos convolvulacea Willd.; Cissampelos cordata Ruiz. ex J.F. Macbr.;
Cissampelos cordifolia Bojer; Cissampelos cimingiana Turcz.; Cissampelos
delicatula Miers; Cissampelos diffusa Miers; Cissampelos discolor DC.; Cissampelos
discolor Miers; Cissampelos discolor var. cardiophylla A. Gray; Cissampelos diversa
Miuers; Cissampelos elata Miers; Cissampelos ellenbeckii Diels; Cissampelos
eriocarpa Triana & Planch; Cissampelos glaucescens Triana & Planch; Cissampelos
gracilis A. St.-Hil.; Cissampelos grallatoria Miers; Cissampelos guayaquilensis
Kunth; Cissampelos haenkeana C. Presl.; Cissampelos hederacea Miers; Cissampelos
hernandifolia Wall.; Cissampelos heterophylla DC.; Cissampelos hirsuta Buch.-Ham.
ex DC.; Cissampelos hirsutissima C. Presl.; Cissampelos hohautiana C. Presl.;
R. W. Bussmann (*)
G. N. Njoroge

Cissampelos limbata Miers; Cissampelos littoralis A. St.-Hil.; Cissampelos longipes

Miers; Cissampelos madagascariensis Miers; Cissampelos mauritiana Thouars; Cis-
sampelos microcarpa DC.; Cissampelos monoica A. St.-Hil.; Cissampelos
nephrophylla Bojer; Cissampelos obteca Wall.; Cissampelos orbiculata DC.; Cis-
sampelos orinocensis Kunth; Cissampelos owariensis P. Beauv ex DC.; Cissampelos
pannosa Turcz.; Cissampelos pareira fo. emarginato-mucronata Chodat & Hassl.;
Cissampelos pareira fo. reniformis Chodat & Hassl.; Cissampelos pareira subvar
crassifolia Engl.; Cissampelos pareira subvar. madagascariensis (Miers.) Engl.;
Cissampelos pareira subvar. usambarensis Engl.; Cissampelos pareira var. australis
(A. St. Hil.) Diels; Cissampelos pareira var. caapeba (L.) Eichler; Cissampelos
pareira var. gardneri Diels; Cissampelos pareira var. haenkeana (C. Presl) Diels;
Cissampelos pareira var. laevis Diels; Cissampelos pareira var. mauritiana (Thouars)
Diels; Cissampelos pareira var. monoica (A. St.-Hil.) Eichler; Cissampelos pareira
var. mucronata (A. Rich.) Engl.; Cissampelos pareira var. nephrophylla (Bojer) Diels;
Cissampelos pareira var. orbiculata (DC.) Miq.); Cissampelos pareira L. var pareira;
Cissampelos pareira var. peltata Scheff.; Cissampelos pareira var. racemifolia
Eichler; Cissampelos pareira var. tamnoides (Willd. ex DC.) Diels; Cissampelos
pareira var. transitoria Engl.; Cissampelos pareira typica Diels; Cissampelos piolanei
Gagnep; Cissampelos salzmannii Turcz.; Cissampelos subpeltata Thwaits ex Miers;
Cissampelos subreniformis Triana & Planch.; Cissampelos tamoides Willd. ex DC.;
Cissampelos testudinaria Miers; Cissampelos testudinum Miers; Cissampelos tetra-
ndra Roxb.; Cissampelos tomentocarpa Rusby; Cissampelos tomentosa DC.; Cis-
sampelos tomentosa Vell.; Cissampelos violifolia Rusby; Cocculus orbiculatus (L.)
DC.; Cocculus villosus Wall.; Dissopetalum mauritianum (Thouars) Miers
Local Names
Cissampelos pareira: Kikuyu: Mubarabaria; Rufiji: Moeru; Sukuma: Msimbasi

(Gachati 1989; Kokwaro 2009).
Botany and Ecology
Cissampelos pareira L.: Shrubby twiners, stems striate, glabrous to tomentose.

Leaves petiolate, obscurely peltate or basifixed, broadly ovate, entire, the apex
obtuse, rarely acuminate, frequently somewhat emarginate, mucronate, the base
truncate or cordate, 2–12 cm long and wide, membranous or papery, palmately 3-
to 7-nerved, in youth more or less sericeous-tomentose on both sides, at length above
more or less pubescent, rarely glabrous, below frequently paler, tomentose or
pubescent, rarely entirely glabrous. Staminate inflorescence multiflowered fascicu-
late dichasia in the leaf axils, sometimes upon secondary axillary branches within the
axils of reduced leaves or bracts, bracts of secondary branches basifixed, sessile or
obscurely petiolate, ovate to subreniform, mucronate, about 1 cm long and wide to
minute, membranous, pubescent to tomentose, bracteoles linear, about 0.5 mm long,
Cissampelos pareira L. 287
pubescent. Staminate flowers: Sepals 4, ovate to obovate, exteriorly pubescent, 1.0–

2.0 mm long, 0.5–1.2 mm wide, corolla campanulate, 0.8–1.8 mm in diameter,
anthers 4, glabrous. Pistillate inflorescence composed of individual flowers fascic-
ulate upon secondary axillary branches within the axils of reduced leaves or bracts,
bracts basifixed, sessile or obscurely petiolate, broadly ovate to reniform, mucronate,
about 1.5 mm long, 2.0 mm wide to minute, membranous, puberulent to tomentose.
Pistillate flowers: Sepal 1, obovate, 1.0–2.0 mm long, 0.5–0.8 mm wide, exteriorly
pubescent, petal 1, broadly obovate to reniform, 0.5–0.8 mm long, 0.7–1.5 mm wide,
glabrous or puberulent, carpel 1, gibbose, densely pubescent. Drupe broadly obovate
or suborbicular, compressed, 4–5 mm long, 3–4 mm wide, pubescent or puberulent,
endocarp verrucose, ribbed (Troupin 1956). A characteristic climber common in dry,
Juniperus procera dominated, Afromontane forests (Bussmann 2002) (Fig. 1).
Cissampelos pareira: Powdered roots in water are used for sore throat, cold, cough,
and snakebites. Leaves and roots are chewed for stomach problems and pregnancy
pains. Burnt roots and leaves are applied to wounds (Bussmann et al. 2006; Kokwaro
2009; Lulekal et al. 2008). In Ethiopia, the species is also used to treat chest pain,
tuberculosis, coughing with blood, flu, boils, swelling, stomachache, diarrhea,
scorpion stings, swelling in stomach, febrile disease, malaria, headache, breast
swelling, ascaris, and intestinal parasites (Teklehaymanot and Giday 2010). The
species has been introduced all over the tropics and is used for the same purposes,
e.g., in Nepal (Kunwar et al. 2015, 2016). The plant juice induces abortion and stops
bleeding after delivery. Leaves and stem parts are used in asthma. Roots are useful in
skin diseases, blood disorders, leukorrhea, diarrhea, dysentery, and indigestion and
have antiperiodic effects. The root paste is applied on snakebites (Kunwar et al.
2009). The species is used for birth issues also in South America, where it is known
as “midwives herb” (Kumar Semwal et al. 2014).
Fig. 1 Cissampelos
mucronata
(Menispermaceae), Mt. Kenya
R.W. Bussmann)
Cissampelos mucronata is used in Ethiopia for abdominal cramps (Bekalo et al.

2009), stomachache, and retained placenta after birth (Giday et al. 2009).
Cissampelos pareira: The species is part of arrow poison mixtures in South America
(Kumar Semwal et al. 2014; Spirova et al. 2019).
References
doi.org/10.1186/1746-4269-5-26.
Kumar Semwal D, Badoni Semwal R, Vermaak I, Viljoen A. From arrow poison to herbal medicine
– the ethnobotanical, phytochemical and pharmacological significance of Cissampelos
Menispermaceae. J Ethnopharmacol. 2014;155:1011–28.
Kunwar RM, Baral K, Paudel P, Acharya RP, Thapa-Magar KB, Cameron M, Bussmann RW. Land-
western Nepal. PlosONE. 2016. https://doi.org/10.1371/journal.pone.0167812.
Lulekal E, Kelbessa E, Bekele, Yineger H. An ethnobotanical study of medicinal plants in Mana
10.1186/1746-4269-4-10.
Spirova EN, Ivanov IA, Kasheverov IE, Kudryavtsev DS, Shelukhina IV, Son LV, Lummis SCR,
Malca-Garcia GR, Bussmann RW, Hennig L, Giannis A, Tsetlin VI. Curare alkaloids from Matis
Dart Poison: comparison with d-tubocurarine in interactions with nicotinic, 5-HT3 serotonin and
GABAA receptors. PLOSOne. 2019;14(1). https://doi.org/10.1371/journal.pone.0210182.
Troupin G. Menispermaceae. Flora of tropical East Africa. London: Crown Agents for Oversea
Governments and Administrations; 1956. 32 pp. ISBN 978-1-84246-286-7.
Clausena anisata (Willd.) Hook. f. ex Benth.
RUTACEAE

Grace N. Njoroge
Synonyms
Clausena anisata (Willd.) Hook. f. ex Benth.: Clausena inaequalis (DC.) Benth.;

Clausena inaequalis var. abyssinica Engl.
Local Names
Clausena anisata: Kikuyu: Muthati, Matathi; Borana: Siskar; Digo: Munyapala;

Kamba: Muthingwa; Luhya: Shingulutsi, Shisimbari, Kisimbari; Maa:
Ol-Matasia; Marakwet: Munyitia, Cheboinoiywa; Meru: Mukibia; Swahili:
Mjavikali; Digo: Munyapala; Luguru: Msogambwa; Zinza: Mtanwa; Luo: Siunya
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Clausena anisata (Willd.) Hook. f. ex Benth.: Shrub or small tree up to 10 m tall,

bark gray, mottled, young branches and leaves pubescent becoming glabrous.
Leaves 6–20(28) cm long, imparipinnate, petiole 1–2.5 cm long, terete or slightly
grooved at the base, leaflets alternate, (4)5–8(9–10) on each side of the rachis,
petiolules slender, 1–3.5 mm long, lamina 1–6(7) 0.6–2.5(3.5) cm, rather smaller
in the proximal than in the distal leaflets, asymmetric, ovate or ovate-oblong or
elliptic to narrowly elliptic, pubescent or glabrous, densely dotted with pellucid
glands, acute or slightly emarginate at the apex, margin crenulate or entire, cuneate
or rounded at the base, lateral nerves 5–8 pairs. Inflorescence of elongate puberulous
panicles, as long as the subtending leaves or a little shorter, peduncle long, bracts and
bracteoles scarious, very small, caducous. Flower-buds 3 mm long, obovoid,
pedicels very slender, 3–4(5) mm long, puberulous. Sepals 4, 0.8 mm long, acute,
slightly joined at the base, pubescent. Petals 4, 4–5 mm long, whitish, imbricate in
bud, elliptic. Stamens 8, 4 mm long, inserted at the base of the disk, filaments
subulate, flattened towards the base, anthers basifixed, 1 mm long, introrse, elliptic,
2-lobed at the base. Ovary 1.3 mm long, sub-ellipsoid, 4(5)-locular with 2 ovules in
each loculus, style stout, stigma broadly obtuse (Kokwaro 1982). Component of
humid semi-deciduous Afromontane forests (Bussmann 2002a, b; Bytebier and
Bussmann 2000).
Clausena anisata: The root decoction is used to remedy whooping cough, malaria,
and body aches (Beentje 1994). Pounded roots in soup given to women to ease
childbirth, and after birth to cleanse the uterus. Also drunk for malaria, flu, headache,
kidney problems, syphilis, and indigestion. In the last case, the roots can also be
chewed. The root decoction also serves as galactagogue. Twigs are chewed as
toothbrush and for toothache (Bussmann 2006; Kokwaro 2009). Smoke is used for
cleaning the body (Mohagheghzadeh and Faridi 2006). The leaf juice is used as
eardrops (Teklehaymanot et al. 2007) and malaria treatment (Teklehaymanot 2009;
Yineger et al. 2007), as well as rheumatism (Yineger et al. 2008).
Clausena anisata: Leaf decoction is given to livestock against endoparasites

(Kokwaro 2009). Leaf juice is inhaled against evil eye (Giday et al. 2010).
Clausena anisata (Willd.) Hook. f. ex Benth. 291
References
Bussmann RW. Succession and regeneration patterns in east African mountain forests – a review.
Bytebier B, Bussmann RW. Vegetation of Mount Nyiru Samburu District, Kenya: A checklist and
syntaxonomical survey. J East Afr Nat Hist. 2000;891(2):45–71.
10.1186/1746-4269-4-11.
Clematis brachiata Thunb.
RANUNCULACEAE

Grace N. Njoroge
Synonyms
Clematis brachiata Thunb.: Clematis incisodentata Rich.; Clematis microcuspis

Baker; Clematis orientalis subps. brachiata (Thunb.) Kuntze; Clematis brachiata
subsp. brachiata (Thunb.) H. Perrier; Clematis oweniae Harv.; Clematis petersiana
Klotzsch; Clematis simensis Fresen.; Clematis stewartiae Burtt Davy; Clematis
thunbergii Steud.; Clematis triloba A. St.-Hil.; Clematis triloba Thunb.; Clematis
viridiflora Bertol
Local Names
Clematis brachiata: Kikuyu: Mugaya-Ng’undu, Mukuyumundu; Ndorobo:

Ilkisuchi, Naitashingisho, Nkolekole; Kamba: Ewalia; Kipsigis: Kisengwet,
Bisinda; Luhya: Lunyili; Luo: Achugo, Achogo; Maa: Olkisushet; Marakwet:
Pising; Samburu: Sigiet; Haya: Minkamba; Acholi: Omwombyer (Beentje 1994;
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Clematis brachiata Thunb.: Woody vine. Branches ca. 8-sulcate, glabrous or

puberulous. Leaves once to twice pinnate, leaflets subcoriaceous, ovate, rarely
lanceolate, 1.4–4( 8) 0.8–3( 5.5) cm, apex acute or acuminate, base truncate,
rounded, or broadly cuneate, margin dentate or denticulate, 3-lobed, sometimes
undivided, on both surfaces subglabrous or puberulous, abaxially not reticulate,
basal veins abaxially slightly prominent, petioles 2–5 cm long. Cymes axillary and
terminal, (3–)5–20-flowered, often panicle-like, peduncles 1.2–3.2 cm long, bracts
narrowly ovate or rhombic, 4–8 mm long, usually 3-lobulate. Flower 2–2.4 cm in
diameter, pedicel 1–1.8 cm long, puberulous. Sepals 4, white, narrowly oblong or
lanceolate-oblong, 9–13( 16) 3–4.5( 6) mm, apex obtuse or slightly acute, on
both surfaces appressed-puberulous, margin velutinous. Stamens 7–10 mm long,
filaments narrowly linear, pubescent, anthers oblong or narrowly oblong, 1.6–3 mm
long, glabrous, apex obtuse. Ovaries densely puberulous, styles 6–7 mm long,
densely villous. Achenes compressed, elliptic, 3–3.5 2–2.5 mm, puberulous,
narrowly rimmed, persistent styles ca. 2.5 cm long, plumose (Milne-Redhead and
Turrill 1952). Widespread in drier Afromontane forests (Bussmann 2002) (Figs. 1, 2,
3, 4, 5, and 6).
Phytochemistry
Alkaloids (clampine), triterpene saponins, coumarins, flavonoids, melissic acid,

myricyl alcohol, caulosapogenin glycoside (Fedorov 1984).
Fig. 1 Clematis brachiata

(Ranunculaceae), Tbilisi,
Georgia. (Photo
R.W. Bussmann and
Clematis brachiata Thunb. 295

Georgia. (Photo
R.W. Bussmann and
Clematis brachiata: Root decoction is used against colds and fever (Beentje 1994).
The leaf infusion is used to treat gonorrhea, syphilis, and sore throat. A root infusion
serves for yaws and malaria. Also acts as purgative. Fresh crushed roots are used for
colds, congestion, headache, and body swellings (Kokwaro 2009). Used for lung
cancer and cancer in general. The plant is crushed and mixed with butter and applied
to wounds, hemorrhoids, and burns (Bussmann et al. 2011). The smoke is used as
analgesic (Mohagheghzadeh and Faridi 2006). The steam of boiled leaves is inhaled
for common cold (Njoroge and Bussmann 2006), and the leaves are applied to leg
swellings (Wondimu et al. 2007). In Ethiopia, Clematis simensis and Clematis
hirsuta have medicinal use (Bekalo et al. 2009). The smoke is used as analgesic
(Mohagheghzadeh and Faridi 2006). Both species have shown antibacterial proper-
ties (Geyid et al. 2005). Clematis hirsuta is used for elephantiasis, trachoma,
cataracts, and hemorrhoids, as well as for wounds (Giday et al. 2007, 2009, 2010).
The leaf juice is applied to the ear canal for earache (Yineger et al. 2007) and also
used for other health problems (Yineger and Yewhalaw 2007). A leaf paste is also
applied externally for leishmania and herpes zoster (Teklehaymanot et al. 2007). In
Madagascar, the leaves of Clematis mauritiana are often sold in markets as

Georgia. (Photo
R.W. Bussmann and
Fig. 4 Clematis vitalba

(Ranunculaceae), Bale
Mountain National Park, Odo
Bulu forest, Ethiopia. (Photo
R.W. Bussmann)
medicine for stomach ulcer, hepatitis, erectile dysfunction, as antiasthmatic, for

rheumatism, cough, bronchitis, and abdominal pains (Randriamiharisoa et al.
2015). Clematis longicauda serves to treat toothache, wounds, and skin itch
(Giday et al. 2010). In Central Middle Asia, the leaf powder of Clematis orientalis

R.W. Bussmann)
is applied to fractures. The extract is used to treat tuberculosis. As poultice applied

externally for itchy eczema. Especially in Tadzhikistan, it is applied to skin problems
and used as antidote for snakebites (Ahmad Jan et al. 2019; Batsatsashvili et al.
2017; Fedorov 1984). Clematis grata is used to treat dermatitis (Ur-Rahman et al.
2018). Clematis buchaniana serves for skin ailments (Bhat et al. 2013), tonsillitis,
and pharyngitis (Ma et al. 2019); Clematis montana is used for diabetes and urinary
system problems (Bhat et al. 2013). A decoction of Clematis grata is used as
anthelminthic (Ali et al. 2019; Bhat et al. 2013) and for dermatitis (Ur-Rahman
et al. 2018). Clematis armandii is used for urinary tract infections, dysuria, nephritis,
and edema (Ma et al. 2019). The smoke of Clematis vitalba is used as analgesic
(Mohagheghzadeh and Faridi 2006).
Local Food Uses
Clematis orientalis: The young leaves are boiled and eaten (Bussmann et al. 2016a, b,
2018; Bussmann 2017).

R.W. Bussmann)
Clematis brachiata: Dried and ground leaves are used as snuff. The plant has also
ritual purpose (Kokwaro 2009). The species, together with Clematis simensis, is
used for ropes (Bussmann 2006; Bussmann et al. 2011). Clematis hirsuta is browsed
by livestock (Bussmann 2006; Bussmann et al. 2011). Clematis angustifolia also
serves for ropes (Hart and Bussmann 2018).
References
medicine research, prospects and limitations in Pakistan: a literature review. 2019. Acta Ecol Sin
https://doi.org/10.1093/ecam/neh088.
Ali A, Aldosari A, Tng DYP, Ullah M, Hussain W, Ahmad M, Hussain J, Khan A, Hussain H,
veterinary practices at Kurram District, Pakistan. Ethnobot Res Appl. 2019;18(24). https://doi.
org/10.32859/era.18.24.1-19.
Batsatsashvili K, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S, Tchelidze D, Paniagua
Zambrana NY, Bussmann RW. Smilax excelsa L. In: Bussmann RW, editor. Ethnobotany of the
Caucasus. Cham: Springer International Publishing; 2017.
doi.org/10.1186/1746-4269-5-26.
Bussmann, R.W. ed. 2017. Ethnobotany of the Caucasus. Springer International Publishing: Cham.
XXVII, 746 p. ISBN 978-3-319-49411-1.
Caucasus. J Ethnobiol Ethnomed. 2016a;12:43. https://doi.org/10.1186/s13002-016-0110-2.
2167-0412.1000266.
Sakartvelo Republic of Georgia, Caucasus. Indian J Tradit Knowl. 2018;17(1):7–33.
Fedorov AA, editor. Plant resources of the USSR: flowering plants, their chemical composition,
use. Volume 1. Families Magnoliaceae – Limoniaceae. Leningrad: Akademia Nauk; 1984.
Geyid A, Abebe D, Debella A, Makonnen Z, Aberra F, Teka F, Kebede, Urga K, Yersaw K, Biza T,
and Amhara peoples in Northwest Ethiopia. J Ethnopharmacol. 2007;110:516–25.
Hart R, Bussmann RW. Trans-Himalayan transmission, or convergence? Stauntonia
Lardizabalaceae as an ethnoveterinary medicine. Med Secoli. 2018;30(3):329–48.
Ma Y, Liu D, Cheng H, Bussmann RW, He H, Guo Z, Liu B. Ethnobotanical study of medicinal
plants used by Miao people in Jijiezi, Yunnan, China. Ethnobot Res Appl. 2019;18(26). https://
doi.org/10.32859/era.18.26.1-14.
Milne-Redhead E, Turrill WB. Flora of tropical East Africa: Ranunculaceae. London: Crown
Agents for the Colonies; 1952.

in the markets of Antananarivo, Madagascar. Randriamiharisoa et al. J Ethnobiol Ethnomed.
2015;11:60. https://doi.org/10.1186/s13002-015-0046.
Clerodendrum myricoides (Hochst.) R. Br. ex
Vatke
LAMIACEAE

Grace N. Njoroge
Synonyms
Cyclonema myricoides (Hochst.) Hochst.; Rotheca myricoides (Hochst.) Steane &

Mabb.; Siphonanthus myricoides (Hochst.) Hiern; Spironema myricoides (Hochst.)
Local Names
Clerodendrum myricoides: Kikuyu: Munjuga-Iriua, munjugu; Borana: Mara-sisa;

Kamba: Mungaya, Muvweia; Kipsigis: Chesamisiet, Obetiot, Chemogong’; Luo:
Shikuma, Kurgweno; Maa: Ol-magotogot; Tugen: Gobetie; Samburu: Makutikuti;
Chagga: Mbale; Nyamwezi: Mpugambu; Hehe: Mupambaduma; Shambaa:
Nunke; Acholi: Okwero; Luhya: Sikuma (Beentje 1994; Gachati 1989; Kokwaro
2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Clerodendrum myricoides (Hochst.) R. Br. ex Vatke: Shrub up to 3 m tall,

irregularly branched, or a small tree up to 10 m tall, older branches with rough
deeply fissured bark, twigs pale brown, ridged or 4-angular, pithy in the center,
glabrous to velvety hairy. Leaves opposite or in whorls of 3–4, lamina 2–16
(19.5) 0.4–6 (10) cm, narrowly to broadly elliptic, ovate-elliptic or oblanceolate,
oblong or obovate, usually small but in cultivation can attain large dimensions, acute
to acuminate at the apex, cuneate to attenuate at the base, entire to coarsely serrate,
glabrous or pubescent to densely velvety hairy, glandular-punctate beneath, sessile
or petiole up to 15 mm long, with unpleasant smell when crushed. Flowers in few-
several-flowered dichasial cymes arranged in unelaborated and lax to quite extensive
and elongate panicles 6.5–15 (30) cm long, peduncles 0–7 cm long, secondary
peduncles up to 4 cm long, apparent stalks 1–2.5 cm long but true pedicels
3–5 mm long. Calyx often entirely purplish or crimson-margined, glabrous to
hairy, tube cupular, c. 2.5 mm long, lobes semicircular to ovate or triangular,
1.2–5 mm long, quite rounded, obtuse or acute. Corolla asymmetrical in bud
expanding abruptly on anterior side, usually greenish with a white to pale blue to
lilac limb, the median lobe dark blue, tube 5–7 mm long, pubescent at the throat,
lobes unequal, (0.6) 1–1.9 (2) cm (1.5) 3.5–7.5 mm, the upper obovate, the lower
one spathulate and much larger than the other four. Stamens and style long-exserted
and curving upwards. Fruit black, 5–6 mm 8–10 mm, subglobose, depressed,
mostly deeply 4-lobed, glabrous (Paton et al. 2009). Characteristic species in dry,
disturbed Afromontane forests (Bussmann 2002).
Clerodendrum myricoides: Root decoction used as emetic to treat malaria and

venereal diseases (Beentje 1994). Roots cooked in soup are used for chest pain,
bronchitis, colds, and against bleeding gum. The root extract serves for indigestion,
sore throat, syphilis, tonsillitis, malaria and rheumatism, for gonorrhea, and as
purgative. Also used to treat mental illness, sterility, amoebas, as anthelminthic,
and for AIDS. As enema, the decoction serves to treat constipation and impotence.
The roots are eaten for skin complaints. The leaves are chewed and applied to
mumps. Used for malaria, cold, polio, gonorrhea, strength, bone pain, and pneumo-
nia (Bussmann 2006; Njoroge and Bussmann 2006a). The decoction also serves to
treat sexually transmitted diseases and serves as aphrodisiac (Njoroge and Bussmann
2007). Clerodendron sp. is used for asthma, whooping cough, and bronchitis
(Bussmann and Sharon 2006, 2015). Clerodendron chinense is applied in Indian
traditional medicine (Reang et al. 2016), Clerodendron viscosum serves for pain and
skin diseases (Raj et al. 2018). Clerodendron acutifolium and Clerodendron
macrocalycinum are used in Madagascar for wound treatments (Rabearivony et al.
2015; Rakotoarivelo et al. 2013).
Clerodendrum myricoides (Hochst.) R. Br. ex Vatke 303
Clerodendrum myricoides: Pounded roots are given to cattle to treat East Coast
Fever (Kokwaro 2009). Eaten by animals and rarely by livestock and used as
firewood (Bussmann 2006; Bussmann et al. 2011). Clerodendron johnstonii is
used for weakness in cattle (Njoroge and Bussmann 2006b).
References
Appendices 1–7 at www.naturekenya.org/JournalEANH.htm.
medicinal flora of Northern Peru. St. Louis: William L. Brown Center, MBG; 2015. ISBN:
978-0-9960231-2-2.
Paton AJ, Bramley G, Ryding O, Polhill RM, Harvey YB, Iwarson M, Willis F, Phillipson PB,
Balkwill K, Lukhoba CW, Otieno DF, Harvey RM. Flora of tropical East Africa: Lamiaceae.
Kew: Royal Botanic Gardens; 2009.
018-0208-9.
L. Brown Center, MBG; 2013. ISBN: 978-0-9848415-7-8.
Aromat Plants. 2016;5:234.
Clutia abyssinica Jaub. & Spach.
PERACEAE

Grace N. Njoroge
Local Names
Clutia abyssinica: Kikuyu: Muthima-Mburi; Kamba: Mutemavengi; Kipsigis:

Kurbanyat; Maa: Olgebarinyei; Marakwet: Chekelel, Kioswa, Sitab oin; Taita:
Mwendinguku; Tugen: Kurrukanya; Turkana: Echiato; Pare: Mruruku; Shambaa:
Muhende; Luhya: Sambukwe (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Clutia abyssinica Jaub. & Spach.: A large erect woody herb, shrub or small tree up to
8 m tall with straight glabrous, pubescent or tomentose branches. Petiole canaliculate,
0.5–2.3 cm long, pubescent beneath or glabrescent, blade ovate, elliptic or lanceolate,
2–17( 20) cm long, 1–5.5( 6) cm broad, subacute or obtuse, tapered, cuneate or
rounded at the base, membranous, lateral nerves 5–12 pairs, cystoliths barely visible,
glabrous, sparingly or evenly pubescent or densely tomentose above and beneath.
Stipules 0. Seeds 3 mm long, 2.5 mm broad. Female flowers: pedicels 5 mm long,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Clutia abyssinica

(Peraceae), Bale Mountains
R.W. Bussmann)
extending to 3 cm in fruit, glabrous or pubescent, sepals ovate-oblong to oblong-

lanceolate, 2–2.5 mm long, 1–1.5 mm broad, often puberulous without at the base,
pale brownish green, each with a 2–3-lobed yellowish green gland at the base,
extending to 3 mm in fruit, petals broadly spathulate, 2 mm long, 1 mm broad,
glabrous, pale dull yellow, eglandular at the base, ovary 1 mm diameter, usually
glabrous, styles 0.75 mm long, free, dull yellow. Fruits 4.5–5.5 mm diameter,
shallowly pustulate, glabrous or rarely very sparingly pilose, endocarp 0.6 mm
thick. Male flowers: pedicels 2–8( 10) mm long, pubescent or glabrescent, distinctly
articulated about half way, often pubescent above the articulation and glabrescent
below it, sepals elliptic-obovate, 2–2.5 mm long, 1–1.5 mm broad, glabrous or
puberulous without at the base, pale green, each with 3 filiform-cylindric hooked
glands at the base, petals deltate, unguiculate, limb 1.5 mm long, 1–1.5 mm broad,
claw 0.5–1 mm long, glabrous, pale greenish yellow or white, each with 1 globose
gland at the base, staminal column 1.5 mm long, filaments 1 mm long, anthers
0.75 mm long, pistillode 1 mm long, thicker than the staminal column. Female
fascicles 1-many-flowered, bracts as in male. Male fascicles dense, few-many-flow-
ered, bracts broadly deltate, 0.5 mm long (Smith 1987; Smith and Carter 1988). The
species is a characteristic component in disturbed Afromontane forests, especially in
drier parts dominated by Juniperus procera (Bussmann 2002a, b, c, 2004; Bytebier
and Bussmann 2000) (Figs. 1 and 2).
Clutia abyssinica: Root decoction used for liver problems (Beentje 1994; Kokwaro
2009). The root decoction is also taken for colds, fever, headache, malaria, stom-
achache, flu, and indigestion (Kokwaro 2009; Njoroge and Bussmann 2006a). The
leaves are applied to treat tonsillitis (Njoroge and Bussmann 2006b) as well as for
Clutia abyssinica Jaub. & Spach. 307
Fig. 2 Clutia abyssinica

(Peraceae), Bale Mountains
R.W. Bussmann)
rheumatism (Yineger et al. 2008). The whole plant is burnt, and the smoke is used to
treat gynecological problems (Mohagheghzadeh and Faridi 2006).
Clutia abyssinica: To most peoples in African Mountain Regions, the species is

virtually useless (Bussmann 2006; Bussmann et al. 2011). However, a leaf decoction
is used to treat diarrhea in livestock (Yineger et al. 2007).
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002b;91 1/2:27–79,
Bussmann RW. Vegetation ecology and regeneration of Tropical Mountain forests. In: Ambasht RS,
Ambasht NK, editors. Modern trends in applied terrestrial ecology. New York: Kluwer; 2002c.
p. 195–223.
Mountain forests: the role of natural and anthropogenic disturbance. Lyonia. 2004;61:98–111.
syntaxonomical survey. J East Afr Nat Hist. 2000;891-2:45–71.
Smith AR. Flora of Tropical East Africa: Euphorbiaceae (Part 1). Rotterdam: Balkema; 1987.
408 pp. ISBN 90-6191-332-2 [Fl. Trop. E. Africa, Euphorb.].
Smith AR, Carter S. Flora of tropical East Africa: Euphorbiaceae (Part 2). Rotterdam: Balkema;
1988. p. 409–597. ISBN 90-6191-338-1.
10.1186/1746-4269-4-11.
Coccinia grandis (L.) Voigt
CUCURBITACEAE

Grace N. Njoroge
Synonyms
Coccinia grandis (L.) Voigt: Bryonia grandis L.; Cephalandra indica Naudin;
Coccinia cordifolia (L.) Cogn.; Coccinia indica Wight & Arn.
Local Names
Coccinia grandis: Kikuyu: Kigerema; Swahili: Ruho; Luo: Mutkuru, Nyathund-Guok

(Gachati 1989; Kokwaro 2009); English: Ivy gourd, Scarlet-fruited gourd, Tindori
Botany and Ecology
Coccinia grandis (L.) Voigt: Climber to 20 m, stem glabrous except at nodes,

becoming white-punctate when older. Leaf-lamina 3.5–11.5 3.5–15.5 cm, broadly
ovate in outline, cordate, glabrous, punctate, palmately 3–5-lobed, lobes shallowly
R. W. Bussmann (*)
G. N. Njoroge

triangular to elliptic, entire or sinuate-dentate, sometimes lobulate. Petiole 1–5 cm

long, glabrous or almost so. Tendrils simple. Male flowers solitary, pedicels 7–
70 mm long. Receptacle-tube 3–7 mm long, obconic, expanded above, lobes 2.5–
6 mm long, triangular to lanceolate or oblanceolate. Corolla campanulate, pale
yellow, green-veined, the lobes 1.5–2 1–1.5 cm. Female flowers solitary, axillary,
pedicels 4–25 mm long, ovary 5–15 1.5–3.5 mm, ellipsoid-cylindrical, receptacle-
tube 2–7 mm long, shortly cylindrical, lobes 2–4 mm long, corolla-lobes 2–
3.2 0.7–1.3 cm. Fruit 3–6.5 1.5–3.5 cm, ellipsoid, obtuse, red. Seeds
6 3 1.5 mm, asymmetrically ovate in outline, compressed, with flat faces and
thick 2-grooved margins. Coccinia grandis occurs wild from Senegal east to Soma-
lia and south to Tanzania, Yemen, and India. It is locally naturalized in Mozambique
and Mauritius and has been introduced in many other tropical and subtropical
regions. In East Africa, especially in Kenya, it is cultivated mainly for consumers
of Indian origin (Jeffrey 1967).
Coccinia grandis: The fruits, stems, and leaves have medicinal uses such as to
reduce high blood pressure and to treat abscesses. Roots are believed to heal illnesses
associated with endocrine system disorders such as diabetes mellitus and are used in
Niger to treat intestinal troubles. Bronchial inflammation, respiratory mucosa, and
skin disorders are also said to be healed by this plant. Pounded leaves with fat are
applied externally for spleen problems, lower abdominal pain, and to lighten the skin
(Kokwaro 2009). The juice is applied to skin infections and wounds (Teklehaymanot
and Giday 2010). Leaves of Coccinia cordifolia are used to increase low blood
pressure, and the leaves of Coccinia indica are eaten for gastroenteritis (Raj et al.
2018).
Local Food Uses
Coccinia grandis: The ripe red fruits of sweet-tasting cultivars are eaten raw, or they
are peeled and cut into pieces and prepared as a stew with onions and tomatoes.
Other vegetables may be added such as garden egg, bottle gourd, ridge gourd, okra,
and pulses. Meat is added when available. The dish may be served with bread,
sorghum, or any other starchy food. The dish is known in East Africa as “tindori.”
Immature green fruits are prepared in soups and curries and widely used in this way
in Ethiopia. Cultivars with bitter fruits are mainly used for their leaves and shoots.
The leaves are also eaten as a vegetable by the Mursi people in Ethiopia. The seeds
are chewed in Ethiopia and Kenya (Kokwaro 2009). Leaves and fruits of Coccinia
cordifolia and Coccinia indica are eaten as vegetable (Raj et al. 2018).
Coccinia grandis (L.) Voigt 311
Coccinia grandis: Fodder for goats, sheep, cattle, donkeys, and camels (Bussmann
2006).
References
Jeffrey C. Flora of tropical East Africa: Cucurbitaceae. London: Crown Agents for Overseas
northern Bengal, India. J Ethnobiol Ethnomed. 2018;141:8. https://doi.org/10.1186/s13002-
018-0208-9.
Coffea arabica L.
RUBIACEAE

Grace N. Njoroge
Synonyms
Coffea arabica L.: Coffea arabica var. stenophyllea Bello; Coffea corymbulosa
Bertol.; Coffea laurifolia Salisb.; Coffea moka Heynh.; Coffea sundana Miq.; Coffea
vulgaris Moench
Local Names
Coffea arabica: Kikuyu: Kahua, Kamba: Kaawa; Swahili: Kahawa; Orma: Buna
(Beentje 1994; Gachati 1989; Kokwaro 2009); English: Coffee
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Coffea arabica L.: Bush or tree 2–7 m tall, young branches glabrous, covered with
light brown shiny bark. Leaf blades elliptic to broadly elliptic or oblong-elliptic
(or lanceolate or round in some cultivated forms), 7–18 cm long, 3–7.5 cm wide,
distinctly acuminate at apex, acute to obtuse at base, thinly coriaceous to coriaceous,
with 7–10 pairs of main lateral nerves, shiny above, margin sometimes undulate,
domatia rather inconspicuous, glabrous, occasionally absent from some leaves,
stipules triangular, 4–8 mm long, acute, usually exceeded by a mucro. Flowers
(4–)5(–6)-merous, 2–20 per axil, borne in 1–3 (–15) fascicles, peduncle 0.5–2 (–3)
mm long in flower (2–4 mm long in fruit), pedicels 1–2 (–3) mm long (calyces
usually well clear of surrounding bracteole), bracteoles frequently with sub-
foliaceous lobes up to 6 mm long. Calyx-tube 1–2 mm long, limb reduced to a rim
or sometimes irregularly toothed, teeth up to 1 (–2) mm long. Corolla-tube (0.5–)
0.9–1.1 cm long, 2–3 mm wide at throat, lobes oblong, 0.9–1.6 cm long, (2–)
4–6 mm wide, rounded. Fruit red (yellow or purple in cultivated forms), oblong-
ellipsoid or sometimes subglobose, 1–2 cm long, 0.9–1.1 cm wide, pedicel
lengthening to 4–8 mm. Seeds fawn or greenish fawn, 9–1.2 (–1.5) cm long, 6–
7 mm wide. Afromontane forest, 1370–1525 m. Often forming very dense stands.
Completely undisturbed areas covered with this interesting forest type are, e.g.,
encountered in large parts of Mt. Marsabit and Kulal in Kenya. The so-called
Coffeo-Rinoreetum represents one of the tallest associations of the lower
Afromontane forests, with the highest trees forming an about 30–40 m high, closed
upper canopy. The most striking feature of these forests is the very dense shrub
stratum, which is particularly formed by large bushes of wild coffee (Coffea arab-
ica). Besides Coffea, a high number of other shrubs are encountered, namely,
Rinorea convallarioides, Chionanthus battiscombei, Pavetta abyssinica, Rytigynia
neglecta, Erythrococca fischeri, Monodora grandidieri, and Vepris eugeniifolia,
making the shrub stratum nearly impenetrable in some places. Toe fleshy cactacean
Rhipsalis baccifera is dangling from many branches. Toe ground is often covered by
dense mats of the grass Oplismenus hirtellus and Acanthaceans as Justicia betonica
and Isoglossa lactea. Of special interest is the occurrence of Ocotea kenyensis,
Casearia battiscombei, Celtis africana, and C. gomphophylla, all of them often
growing to huge trees, up to 40 m tall, and also regenerating very well in the shrub
layer. In the lower canopy and the dense understorey, Strychnos henningsii and
S. mitis grow regularly, and are joined by Flacourtia indica, Turraea holstii,
Erythrococca bongensis, Meyna tetraphylla, and Cordia monoica (Bussmann
2002). Many of these species indicate links of these forests to the coffee-rich forests
of Southern Ethiopia, particularly to the Bale region (Bussmann 1997). Large trees
like Strombosia schefjleri, Apodytes dimidiata, Albizzia gummifera, Premna max-
ima, as well as Croton macrostachyus and especially Aningeria adolfi-friedericii are
encountered in the upper canopy and in the shrub layer, together with Ritchiea
albersii. The occurrence of these tree species in Harenna as dominating species is
very remarkable, as they are otherwise unknown to the Ethiopian flora, and their next
Coffea arabica L. 315
Fig. 1 Coffea arabica

(Rubiaceae), Bale Mountains
National Park, Harenna forest,
Ethiopia. (Photo: R. W.
Bussmann)

(Rubiaceae), Bale Mountains
National Park, Harenna forest,
Ethiopia. (Photo: R. W.
Bussmann)
populations are found about 1000 km south in Central Kenya, and there especially
around Nairobi (Bekalo et al. 2009; Bussmann 2002, 2006) (Figs. 1, 2, 3, and 4).
Coffea arabica: Leaf decoction drunk for the treatment of hepatic diseases
(Kokwaro 2009). Sometimes the species is used as analgesic (Bussmann and Sharon
2006, 2015). Roasted and powdered coffee is pasted with honey and taken orally for
the treatment of diarrhea and amoebiasis (Gedif and Hahn 2003; Giday et al. 2009;
Luizza et al. 2013; Lulekal et al. 2008). The leaves are boiled to treat headache

(Rubiaceae), fresh coffee
beans, Bale Mountains
National Park, Goba market.

(Rubiaceae), freshly roasted
coffee beans, Bale Mountains
(Giday et al. 2010). The roots can be boiled to treat cough (Njoroge and Bussmann
2006), as well as back pain in women (Njoroge and Bussmann 2009). In Madagas-
car, the leaves are boiled to treat malaria (Razafindraibe et al. 2013). The leaves of
Coffea perrieri are boiled to ease pregnancy pains (Randrianarivony et al. 2016,
2017).
Coffea arabica L. 317
Local Food Uses
Coffea arabica: Coffee cultivation may have started in the sixth century in Yemen,
its use being for spiritual purposes. Coffee became a popular drink in Europe from
the seventeenth century onwards, being imported from plantations established first
by the Dutch in Ceylon (now Sri Lanka), Java, and later from plantations in Brazil
and the West Indies established in the eighteenth century. Much of the world’s
Arabica coffee is produced in Latin America. The stimulating effects of coffee are
largely due to the alkaloid caffeine contained in the seeds. As well as a beverage,
coffee is used for flavoring foods and confectionery. The beans are also a commercial
source of caffeine, a by-product of making decaffeinated coffee. In some countries,
coffee leaves are used to make a hot drink, like tea. Of course, coffee ranks among
the world’s most used “magic” plants (Bussmann 2016). In Ethiopia, the seeds are
used of course to prepare coffee, and the leaves are roasted to make tea (Bussmann
et al. 2011; Mekonnen et al. 2015).
Coffea arabica: Coffee wood, from the main trunk, is used locally in construction.
David Livingstone, the nineteenth century explorer and missionary, reported seeing
coffee trees being used to make huts in his travels in southern Africa. The timber is
straight, dense, strong, and partially resistant to termites. The wood is also used for
furniture and as fuel wood. Dried roasted berries are pounded, and boiled, and the
decoction given to camels to treat trypanosomiasis (Kokwaro 2009).
References
doi.org/10.1186/1746-4269-5-26.
Bussmann RW. Toe forest vegetation of the Harenna Escarpment (Bale Province, Ethiopia) –
syntaxonomy and phytogeographical affinities. Phytocoenologia. 1997;27(1):1–23.
2006;111:41–66.
Bussmann RW. Magic plants. In: Albuquerque U, Alves R, editors. Introduction to ethnobiology.
Heidelberg: Springer; 2016. p. 163–9.
978-0-9960231-2-2.
Gedif T, Hahn H-J. The use of medicinal plants in self-care in rural Central Ethiopia.
doi.org/10.1186/1746-4269-5-34.
Res Appl. 2013;11:315–39.
Lulekal E, Kelbessa E, Bekele YH. An ethnobotanical study of medicinal plants in Mana Angetu
District, southeastern Ethiopia. J Ethnobiol Ethnomed. 2008;4:10. https://doi.org/10.1186/1746-
4269-4-10.
Njoroge GN, Bussmann RW. Traditional management of ear, nose and throat (ENT) diseases in
Knowl. 2009;82:262–9.
Randrianarivony NT, Randrianasolo A, Andriamihajarivo T, Ramarosandratana AV, Jeannoda VH,
Rakotoarivony F, Bussmann RW. Useful plants and tradition for pregnancy, child delivery and
for postpartum care used by people living around Analavelona forest located in Southwest
Madagascar. Indian J Tradit Knowl. 2016;151:68–78.
2017; https://doi.org/10.1186/s13002-017-0147-x.
2013;9:73.
Cola greenwayi Brenan
MALVACEAE

Grace N. Njoroge
Synonyms
Cola microcarpa Brenan
Local Names
Cola greenwayii: Kikuyu: Muthakai, Mwangao (Beentje 1994; Gachati 1989).
Botany and Ecology
Cola greenwayi Brenan: Tree 3–20 m high, monoecious or dioecious, young

branchlets russet to light cinnamon brown or grey tomentose at first, mixed with
stellate or bunched hairs which fall readily. Stipules very soon falling, subulate-
lanceolate to linear, 3–6 mm long, dark to light-brown or grey tomentose. Leaves
simple, entire, blade elliptic to narrowly obovate-elliptic, narrowing to base and
R. W. Bussmann (*)
G. N. Njoroge

apex, 4–15 cm long, 1.4–5 cm broad, glabrous at maturity, midrib slender, prominent
on both surfaces, with usually 7–18 main lateral veins, distinct to prominent on
lower surface, petiole 0.5–5.5 cm long with a thickened pulvinus at the apex,
tomentose throughout, with some long hairs as well, becoming glabrous except for
the pulvinus which is persistently tomentose. Inflorescence 1-flowered, axillary,
solitary or fascicled and crowded on branchlets between the leaves, flowers unisex-
ual, rarely some bisexual, apparently dioecious (may be monoecious), arising from
sessile imbricating bracts which are 1.5–3.5 mm long, densely stellate-tomentose
without subpersistent, pedicels about 7–20 mm long, (sometimes obscurely articu-
late), about midway, densely stellate-pubescent, hairs dark to light brown, short or
long (as long as the pedicel is broad). Calyx 4- to 6-lobed almost to the base at
flowering time, dorsally stellate-pubescent, lobes from about 5–10 mm long,
2.5–3.5 mm broad, inner face stellate-pubescent at least in upper half lepidote
below. Petals 0. Male flower with staminal tube about 2.5–4.5 mm long, densely
pubescent to subglabrous, anthers up to 8 in one row, sessile in a ring around the
apex of the staminal-tube, rudimentary carpels and styles in the centre of the ring.
Female flower with ovary sessile, densely stellate-pubescent, about 2 or 3 mm diam.,
with a ring of rudimentary stamens around the base, carpels 4–5 (3?), styles arising
laterally with clavate recurved, papillose stigmas. Carpels 4–5 or less by abortion,
orange yellow to deep orange with touch of vermilion when ripe, obliquely sub-
globose obliquely ellipsoid or obovate, 1 1.3–1.8 2 cm rounded on top with the
remains of the style-base lateral and obscure, or forming a mucro, stellate pubescent
in parts (rubs off easily) with dark to light brown hairs, long bristles, rind at maturity
rather thin and brittle, seeds 1–2 (coat like that of fruit), cotyledons 2 (one smaller
than the other, reddish pink tinge between them in fresh fruit), plumule stellate-
pubescent. Common undergrowth tree in Croton dominated Afromontane forests
(Bussmann 2002).
Cola greenwayi: Sometimes used as firewood
References
Commelina africana L.
Commelina benghalensis L.
COMMELINACEAE

Grace N. Njoroge
Synonyms
Commelina africana L.: Dirtea africana (L.) Raf.; Hedwigia africana (L.) Medik.;
Stickmannia africana (L.) Raf.
Commelina benghalensis L.: Commelina canescens Vahl; Commelina cavaleriei
Lév.; Commelina cucullata L.; Commelina delicatula Schltdl.; Commelina mollis
Jacq.; Commelina nervosa Burm. f.; Commelina procurrens Schltdl.; Commelina
turbinata Vahl.
Local Names
Commelina africana: Kikuyu: Mukengeria; Swahili: Kongwa; Digo: Dzedza

Chetu; Kipsigis: Loblobityet; Samburu: Naitetyiai; Luhya: Nioloniolo (Beentje
1994; Gachati 1989; Kokwaro 2009); English: Yellow commelina, Wandering Jew,
Dayflower.
R. W. Bussmann (*)
G. N. Njoroge

Commelina benghalensis: Kikuyu: Mukengeria; Swahili: Kongwa, Kafula,

Mpovupov; Kipsigis: Loblobityet; Taita: Mkengera; Luo: Odielo (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: Blue commelina, Venus’ bath, Benghal
dayflower, Tropical spiderwort.
Botany and Ecology
Commelina africana L.: Perennial herb with tuberous fusiform fleshy roots, stem
creeping or straggling. Leaves arranged spirally, simple, leaf sheaths 0.8–3 cm long,
with purple tinge, ciliate along the free margins, blade generally lanceolate,
6–11.5 1.2–2.2 cm, apex acute, glabrous except for the ciliate margins, rarely
sparsely hairy, veins parallel. Inflorescence a leaf-opposed cyme, peduncle 8–40 mm
long, spathe 0.9–2.4 cm long, margins free, glabrous except for the ciliate margins.
Flowers bisexual, zygomorphic, yellow, rarely protruding from the spathe, lower
petal linear-lanceolate, c. 5 2 mm, paired petals with claw 3–4 mm long, lamina
broader than long, c. 4 6 mm, upper three stamens sterile, with cross-shaped
anthers, medial stamen with filament 4–6 mm long and anther 1.5–2 mm long, the
two lateral (lower) stamens with smaller anthers. Fruit a capsule 5–6 mm long,
3-celled, 3–5-seeded, the 2 ventral locules each 1( 2)-seeded (by abortion of the
lower ovules) and dehiscent, the dorsal locule 1-seeded, indehiscent. Seeds variable
in size, cylindrical-rectangular in outline, 2.2–3.5 1.3–2 mm, dark brown, testa
rough (farinose granules) and pitted, hilum small, round. Commelina africana is a
variable species, in which many varieties are distinguished. The typical variety, var.
africana, is a cultivated plant grown as a vegetable. Commelina africana is easily
distinguished from the other Commelina species by its yellow instead of blue,
purplish, or pink flowers. Commelina africana is indigenous and widespread in
Africa, occurring from Senegal to Ethiopia, and south to South Africa. It occurs also
in Saudi Arabia, Yemen, and Australia (Faden 2012) (Figs. 1, 2, 3, 4, 5 and 6).
Fig. 1 Commelina africana

(Commelinaceae), Sekenani,
Maasai Mara, Kenya. (Photo
R.W. Bussmann)
Commelina africana L. . . . 323
Fig. 2 Commelina
benghalensis
R.W. Bussmann)
Commelina benghalensis L.: Perennial herb with tuberous fusiform fleshy roots,
stems erect or creeping-ascending, often with subterranean cleistogamous flowers.
Leaves arranged spirally, simple, sheath 0.5–3 cm long with purple veins, ciliate
with purple 3–7 mm long bristles along free edges, rarely over the surface, otherwise
glabrous or with hooked hairs, blade generally broadly lanceolate, 9–12 3–5 cm,
pseudo-petiole 3–15 mm long with a few purple or white bristles 3–7 mm long, apex
acute or acuminate, pubescent or glabrous, veins parallel. Inflorescence a leaf
opposed, falsely terminal cyme, peduncle up to 6 mm long, spathe funnel-shaped
1–2 cm long, margins completely fused, pubescent with short hooked hairs and
sparsely pilose with straight hairs. Flowers bisexual, zygomorphic, protruding from
the spathe when flowering, with pedicel 4–7 mm long, reflexed and enclosed in the
spathe when fruiting, petals blue, lower petal smaller than the 2 upper petals, these
with claw 4–5 mm long, lamina 8 10 mm, upper three stamens sterile, with cross-
shaped, yellow anthers, medial stamen with filament 5–7 mm long and fertile, blue
anther c. 2 mm long, the two lateral (lower) stamens with smaller fertile, blue anthers
c. 1.4 mm long. Fruit a capsule 4.5–5.5 mm long, 3-celled, 3–5-seeded, the 2 ventral
Fig. 3 Commelina
benghalensis
R.W. Bussmann)
locules each 1( 2)-seeded, dehiscent, the dorsal locule 1-seeded, delayed or imme-
diately dehiscent. Seeds variable in size, cylindrical-rectangular in outline,
2–5.5 1.5–2.2 mm, brown, testa reticulate with tuberculate ridges, provided
with farinose granules, hilum normally linear, 1–3 mm long, not reaching the
ends. Blue-flowered Commelina species such as Commelina benghalensis and
Commelina diffusa Burm.f. are easily confused. Both species are common through-
out Africa. The properties attributed to one species may therefore also be valid for
the other. In East Africa, Commelina imberbis Ehrenb. ex Hassk. and Commelina
latifolia Hochst. ex A.Rich. are also used as a vegetable. Commelina zambesica C.B.
Clarke is used as a vegetable on Pemba Island (Tanzania). Commelina benghalensis
is originally an Old-World species and is naturalized in the Americas and Hawaii. It
occurs throughout tropical Africa from Cape Verde and Senegal to Ethiopia, and
south to South Africa; it also occurs in Madagascar and the Mascarene Islands.
(Faden 2012). Often in the undergrowth of disturbed Afromontane forests
(Bussmann 2002a, b).
Fig. 4 Commelina
benghalensis
R.W. Bussmann)
Fig. 5 Commelina
sp. (Commelinaceae),
Sekenani, Maasai Mara,
Kenya. (Photo
R.W. Bussmann)
Fig. 6 Commelina
sp. (Commelinaceae),
Kenya. (Photo
R.W. Bussmann)
Commelina africana: In Kenya, an infusion of the plant is used as a wash to reduce

fever, and pounded stalks are used to treat colds and coughs in children. Fluid from
the spathes is applied locally to cure eye diseases. The Zulu of South Africa bathe the
body, especially of a child, with a cold infusion in cases of restless sleeping.
Similarly, an infusion of the leaves is sprinkled over the resting place of a restless
child in Zimbabwe. The Sotho in southern Africa take a decoction of the plant with
Tephrosia capensis Pers. for treatment of a “weak heart” and nervousness. In DR
Congo, the root is used for the same purpose. The plant cooked with Haplocarpha
scaposa Harv., Helichrysum pilosellum (L.f.) Less. or the root of Cotyledon
decussata Sims is given by Sotho as medicine to young women to cure infertility.
Also, an infusion of the plant is drunk, and its ash is rubbed over the loins as a
fertility charm. In Zimbabwe and South Africa, a concoction of the root is used as
treatment for venereal diseases and to treat women with menstrual cramps. This
preparation is also used for pelvic pains and bladder complaints. Latex from flower
buds is used as antidote to latex burns in eyes. Infusion to reduce fever. Juice is also
applied to eyes and earache. A decoction of branches with milk given to children for
cold and cough. Dried leaves are powdered and applied to wounds or used for
massages (Kokwaro 2009; Njoroge et al. 2004; Teklehaymanot 2009). Boiled in
milk to treat cough in children (Bussmann 2006; Nankaya et al. 2019). Used also for
allergies and ear infections (Teklehaymanot et al. 2007).
Commelina benghalensis: In southern Nigeria, the plant is used as a poultice for
sore feet. In East Africa, the sap of leaves and stems is used to treat ophthalmia, sore
throat, and burns, and the liquid contained in the flowering spathe is used to treat eye
complaints in Zanzibar. In Uganda and Tanzania, the sap is used topically against
thrush in infants, and in Tanzania, a solution of pounded leaves soaked in warm
water is used to treat diarrhea. In southern Africa, Commelina benghalensis is used

to counter infertility in women, and a decoction of the root is used for the relief of
stomach disorders. Flower juice for eye problems and sore eyes. Stem sap used for
colds, earache, sore throat, burns, and thrush (Kokwaro 2009). Boiled in milk to treat
cough in children (Bussmann 2006; Nankaya et al. 2019). The leaf paste is also
applied to wounds (Njoroge et al. 2004). Used also for allergies and ear infections
(Teklehaymanot et al. 2007).
Commelina madagascarica is used as abortifacient and against acne
(Randriamiharisoa et al. 2015). Leaves of Commelina foliacea are used to treat
headache in children (Yineger et al. 2008).
Local Food Uses
Commelina africana: In Kenya, Uganda, and Tanzania, the leaves are cooked and
eaten as a vegetable. They are chopped and boiled in water or in fresh or sour milk.
Sesame seeds and groundnut paste are added for flavor and consistency. This
vegetable is eaten with the staple food as a substitute for more preferred vegetables.
It used as famine food (Balemie and Kebebew 2006; Bekalo et al. 2009).
Commelina benghalensis: In parts of West Africa, e.g., Côte d’Ivoire and Ghana,
the leaves of Commelina benghalensis are cooked and eaten as a vegetable. The
leaves are mucilagenous. In Kenya, young leaves are eaten as a relish, and older
leaves are regarded as too acidic and bitter to use. In Kenya, Uganda, and Tanzania,
the leaves and stems are chopped and cooked alone or with other vegetables such as
Bidens pilosa L. or Cleome hirta Oliv. It is also reported as a vegetable in Ethiopia.
In Indonesia, the leaves and young tops are occasionally steamed and eaten as a
vegetable, and in the Philippines, they are eaten cooked. It is used as famine food
(Balemie and Kebebew 2006; Bekalo et al. 2009).
Commelina africana: In Kenya and Tanzania, the leaves are fed to livestock,
especially pigs and rabbits. The flowers provide bee forage. Used to prevent evil
spirits from entering the boma (Kokwaro 2009). Eaten by livestock (Bussmann
2006; Bussmann et al. 2011). Used by specialized old men to put protective magic
on places where cows pass (Bussmann et al. 2006). Leaf paste applied to wounds,
especially on feet (Giday et al. 2007).
Commelina benghalensis: In Sudan and eastern Africa, the plants are grazed by
domestic stock, at the same time providing part of the cattle’s need for water. In
northern Ghana, it is a favorite feed for pigs and poultry; in Tanzania, it is given to
animals, especially pigs and rabbits. The flowers provide bee forage. In southern
Africa, however, its use as pig feed is restricted to times of scarcity as it is thought to
cause a sort of ‘measles’ in the animals. There may be edaphic or genetic differences
causing such differences in properties, but these may also originate from
misidentifications, because most Commelina species resemble each other very

closely. Used to prevent evil spirits from entering the boma, and also in weddings
as fertility symbol (Kokwaro 2009). Eaten by livestock (Bussmann 2006; Bussmann
et al. 2011). Used by specialized old men to put protective magic on places where
cows pass (Bussmann et al. 2006). Leaf paste applied to wounds, especially on feet
(Giday et al. 2007), and also applied to skin infections (Giday et al. 2009). Plant is
used in bronchial infection and asthma. Latex is applied on warts (Kumar et al.
2011).
References
Balemie K, Kebebew F. Ethnobotanical study of wild edible plants in Derashe and Kucha districts,
doi.org/10.1186/1746-4269-5-26.
Ambasht NK, editors. Modern trends in applied terrestrial ecology. Dordrecht: Kluwer; 2002b.
p. 195–223.
Bussmann RW, Swartzinsky P, Worede A, Evangelista P. Plant use in Odo-Bulu and Demaro, bale
Faden R. Flora of tropical East Africa: Commelinaceae. Kew: Royal Botanic Gardens; 2012.
doi.org/10.1186/1746-4269-5-34.
Ethnomedicinal knowledge. Sustainability. 2019;11:5530. https://doi.org/10.3390/su11195530.
2015;11:60. https://doi.org/10.1186/s13002-015-0046.
ailments at Bale Mountains National Park, southeastern Ethiopia. J Med Plant Res.
2008;26:132–53.
Commiphora africana (A. Rich.) Engl.
Commiphora rostrata Engl.
Commiphora schimperi (O. Bern.) Engl.
BURSERACEAE

Grace N. Njoroge
Synonyms
Commiphora africana (A. Rich.) Engl.: Commiphora cakiicola Engl.;

Commiphora pilosa Engl.; Heudelotia africana A. Rich.
Commiphora rostrata Engl.: Commiphora reflexa Chiov.; Commiphora robecchii
Engl.; Commiphora rostrata var. reflexa (Chiov.) J. B. Gillett
Commiphora schimperi (O. Bern.) Engl.: Commiphora betschuanica Engl.
Local Names
Commiphora africana: Kikuyu: Mukungugu; Swahili: Mbambara; Borana:

Hames, Amess; Gabbra: Hammeesa; Kamba: Mutungu, Kitungu; Luo: Arupien,
Arupiny; Marakwet: Chotwa; Oromo: Komper; Samburu: Icheni-Nzito,
Lecheningiro; Somali: Dabba’Un’Un, Hammes sagara; Taita: Mwagari; Turkana:
Ekadeli; Swahili: Mbambara, Kkorogo; Rangi: Mpome; Sumbua: Msangacha;
Giriama: Tola (Beentje 1994; Gachati 1989; Kokwaro 2009)
R. W. Bussmann (*)
G. N. Njoroge

Commiphora rostrata: Kikuyu: Mukungugu; Borana: Diraa, Tirai; Ilelewa:

Choneh; Oromo: Udesi; Somali: Janau, Danu, Danu-Sagar; Turkana: Lokimeta;
Tharaka: Mutunkuri (Beentje 1994; Gachati 1989; Kokwaro 2009)
Commiphora schimperi: Kikuyu: Mukungugu; Borana: Masakayia; Kamba:
Mutungu; Samburu: Lachimi; Somali: Hargoi, Horgoi; Turkana: Ekwangorom
(Beentje 1994; Gachati 1989; Kokwaro 2009)
Botany and Ecology
Commiphora africana (A. Rich.) Engl.: Shrub or small tree 2–5 m tall, bark
smooth, dark green or yellowish, peeling in yellowish strips, young branches
densely pubescent or tomentose with short whitish hairs or sometimes yellowish
longer hairs, spiny. Leaves 3-foliolate, petiole up to 4.5 cm long, but usually less,
pubescent or pilose, terminal leaflet up to 8 5 cm, often smaller, obovate, apex
acute or obtuse, base gradually cuneate, lateral leaflets about 1/2–3/4 the size of the
terminal one, elliptic to rotund, apex acute or rounded, margins (of all leaflets)
coarsely crenate or crenate-serrate, base rounded or broadly cuneate, both surfaces
pubescent or occasionally glabrous above. Flowers appearing before the leaves in
axillary abbreviated clusters, often borne on the spines, pedicels up to 2 mm long,
pubescent or glabrous. Calyx 2 mm long, campanulate, lobed to about 1/3-way,
glabrous or pubescent. Petals 3–5 mm long, stamen-filaments subterete, slender
above but broadening somewhat near the base. Disk-lobes 4, bifid at the apex.
Fruit 1.2 cm in diameter subglobose, pseudaril apparently absent or occasionally
irregularly lobulate and covering about 3/4 of the endocarp, endocarp 1 cm in
diameter, subglobose but with one face more nearly hemispherical than the other,
rugose, the two faces separated by a narrow wing-like rim (Gillett 1991). The wood
of Commiphora pterocarpa and that of various other Commiphora species is known
as “arofy” in Madagascar. It is locally much used for construction, joinery,
shuttering, and vehicle bodies. A decoction of the bark of Commiphora pterocarpa
is used for treating of ulcerated wounds (Figs. 1 and 2).
Commiphora rostrata Engl.: Spiny shrub or small tree, up to 4 m tall. Glabrous,
bark smooth, gray to purplish black. Exudate very liquid and aromatic. Petiole
1–8 mm long, leaves with entire margin, elliptic, 10–45 mm long and 8–35 mm
wide. Male inflorescence a pedunculate cyme, up to 3 cm long, 8–20 flowered. Petals
dark red, 4–6 mm long. Female inflorescence 1-2 flowered, up to 2 cm long, very
slender. Fruits beaked, 13–18 mm long and 1–4 mm wide, stone 8–10 mm long.
Commiphora schimperi (O. Bern.) Engl.: Shrub or small bushy tree, spiny, quite
glabrous except for a few brown hairs at the apex and base of the petioles, bark
smooth, peeling in yellowish strips. Leaves 3-foliolate, petiole up to 2.5 cm long,
terminal leaflet up to 2.6 1.4 cm, narrowly obovate or obovate, apex acute (often
truncate to the north of our area), base gradually cuneate, lateral leaflets about half
the size of the terminal ones, broadly obovate or ovate, apex acute, base broadly
cuneate, margin of all leaflets coarsely crenate in the upper half. Flowers appearing
before the leaves in axillary abbreviated clusters, often borne on the spines, pedicels
Commiphora africana (A. Rich.) Engl. . . . 333
Fig. 1 Commiphora
sp. (Burseraceae). Laikipia
plateau, Kenya. (Photo: R. W.
Bussmann and N. Y.
Paniagua-Zambrana)
Fig. 2 Commiphora
sp. (Burseraceae). Laikipia
plateau, Kenya. (Photo: R. W.
Bussmann and N. Y.
Paniagua-Zambrana)
1 mm long. Calyx 1.5 mm long, lobed to 1/3-way or less. Petals 3.5 mm long.
Stamen-filaments subterete. Disk-lobes 4, bifid at the apex. Fruits 1.3 0.8 cm,
ellipsoid to globose, slightly apiculate at the apex, pseudaril apparently absent,
endocarp 0.5 0.5 cm, ellipsoid, lumpy-rugose, with one face very deeply convex
and the other shallowly so (Gillett 1991).
Commiphora species are commonly found in the forest-savannah transition zone
at the border of dry Afromontane forests (Balemie et al. 2006; Bussmann 2002,
2006b).
Commiphora africana: Roots are boiled and the decoction is used for fever, leprosy,
swollen testicles, and stomach problems. Steam baths of bark and roots are used for
colds and fevers. Barks are chewed with tobacco and are applied to snakebites. Resin
is used for infected wounds. The fruits are eaten for typhoid and chewed for
toothache and bleeding gums (Kokwaro 2009). Used as toothbrush (Bussmann
2006; Lulekal et al. 2008).
Commiphora rostrata: Bark chewed, and decoction drunk for chest pain, coughs,
and colds. Fresh juice is applied to eye diseases (Kokwaro 2009). Used as toothbrush
(Bussmann 2006; Lulekal et al. 2008; Wondimu et al. 2007).
Commiphora schimperi: Bark decoction is given to children for diarrhea (Kokwaro
2009).
Commiphora resinifera is used as anthelminthic in Ethiopia (Desta 1995). Used
as toothbrush (Bussmann 2006; Lulekal et al. 2008; Wondimu et al. 2007).
Commiphora baluensis and Commiphora boranensis are used for treating
cough, stomach complaints, diarrhea, skin diseases, and as toothbrush (Lulekal
et al. 2008; Wondimu et al. 2007).
Some Commiphora species are used for their hypoglycemic effect (Kunwar et al.
2010). Commiphora eminii serves as antimalarial (Njoroge and Bussmann 2006).
Commiphora mukul smoke serves to purify the air, as febrifuge, and for respiratory
diseases (Mohagheghzadeh and Faridi 2006).
Local Food Uses
Commiphora rostrata: Leaves are eaten as vegetable (Beentje 1994)

In Ethiopia, a large number of Commiphora sp., including Commiphora rostrata
and Commiphora schimperi, contain resin that is used as chewing gum, and the
trunks are chewed to quench ones’ thirst. The fruits of many are eaten (Lulekal et al.
2008).
Commiphora africana: Bark decoction is given to cattle with foot and mouth
disease (Kokwaro 2009).
Commiphora schimperi: Bark decoction is given to goats for diarrhea (Kokwaro
2009). Also sometimes used as firewood (Tian 2017).
The most commonly used species is Commiphora myrrha, widely employed as
incense (Bussmann and Sharon 2006, 2015). Commiphora boiviniana is used as fish
poison (Neuwinger 2004). Commiphora fraxinifolia and Commiphora grandifolia
are used in Madagascar for construction, for live fences, and to make traditional
musical instruments (Randrianarivony et al. 2016). Commiphora erythraea and
Commiphora incisa are used as tick repellents in livestock (Wanzala 2017). In
Ethiopia, a large number of Commiphora species, including Commiphora rostrata
Commiphora africana (A. Rich.) Engl. . . . 335
and Commiphora schimperi are used to make glue, for mending tires, for camel
bells, milk container, spoons, combs, firewood, dry and live fencing, are also
browsed by camels, used to make stools and incense, for construction and as
firestick, to make expensive incense, and to make ink. Also used to make sticks to
castrate goats. Commiphora erlangiana bark is used to make hunting poison
(Lulekal et al. 2008).
References
2006a;111:41–66.
978-0-9960231-2-2.
Gillett JB. Flora of Tropical East Africa: Burseraceae. Rotterdam: Balkema; 1991.
Kunwar RM, Burlakoti C, Chowdhary CL, Bussmann RW. Medicinal plants in farwest Nepal: their
indigenous uses and pharmacological validity. Med Aromat Plant Sci Biotechnol. 2010;4
(1):28–42.
10.1186/1746-4269-4-10.
Ethnomed. 2006;2:8.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016. ISBN: 978-0-9960231-4-6.
10.1186/s13002-016-0131-x.
Conyza volkensii O. Hoffm.
ASTERACEAE

Grace N. Njoroge
Synonyms
Conyza volkensii O. Hoffm.: Conyza steudelii Sch. Bip. ex A. Rich.
Local Names
Conyza volkensii: Kikuyu: Muturia-Thoongo (Beentje 1994; Gachati 1989).
Botany and Ecology
Conyza volkensii O. Hoffm.: Annual or biennial herb, becoming woody at base,

0.3–2 m high, stems erect or scandent, branched, green, cylindrical or angular, pubes-
cent and glandular. Florets whitish, cream or pale yellow, the marginal florets many
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Conyza cf. volkensii

forest. (Photo
R.W. Bussmann)
(>100), the tube 0.7–1.5 mm long, minutely toothed at apex, style 2.6–3.3 mm long,
central florets 6–9, tube 2.1–2.7 mm long, lobes 0.5–0.8 mm long, glabrous
or puberulous, anthers 1–1.4 mm long with narrowly triangular appendages, style
2.6–3.2 mm long, with flattened triangular and papillose branches 0.7–0.8 mm long.
Achenes obovoid, compressed or some of the central ones cylindrical and angled, 0.8–
1 mm long, 3–4-ribbed, glabrous or sparsely pubescent, pappus of many whitish to pale
brown setae 3–4.5 mm long, inserted on a ring. Leaves pale green, (at least the
proximal) spatulate with an ovate expanded part, the more distal ones less spatulate,
1.5–14 cm long, 0.5–6 cm wide, at the base with a winged pseudopetiole with the very
base semi-amplexicaul to auriculate, the two auricles together 0.5–1.5 cm wide, the
margins dentate, the apex acute, sparsely to densely hispidulous to pubesent, sometimes
sparsely glandular and aromatic. Capitula 3–7 mm long, up to 15 together to form
subglobose cymes, several cymes together forming the terminal leafy panicles or
corymbs, stalks of individual capitula 2–6 mm long, pubescent and glandular, phylla-
ries about 30, 3–4-seriate, green tipped with yellow or red, lanceolate, the outermost
smallest, 2–5 mm long, 0.5–1 mm wide, margins scarious and pale, apex acute,
pubescent to glabrescent, receptacle toroid, with some small teeth around the areoles
(Beentje 2000, 2002; Beentje and Jeffrey 2005). Often found in Afromontane forests
(Bytebier and Bussmann 2000). While very common, Conyza volkensii and other
species of the genus have very limited value for the local population and are thus an
excellent example of the fact that the flora of African Mountain Regions is much less
used than the plant resources of similar areas worldwide (Figs. 1 and 2).
Conyza volkensii O. Hoffm. 339
Fig. 2 Conyza sumatrensis

forest. (Photo
R.W. Bussmann)
Conyza pyrifolia is chewed for toothache, and Conyza sumatrensis is applied for
back pains, diarrhea, dysentery, pimples, postpartum pains, stomachache, and tooth-
ache (Njoroge et al. 2004; Tabuti 2008). The species is also used to treat a variety of
sexually transmitted diseases and back pain in women (Njoroge and Bussmann
2009) and to treat pimples (Njoroge and Bussmann 2007). Conyza canadensis
sometimes serves as stimulant, astringent, diuretic, aphrodisiac, and homeostatic
(Ahmad Jan et al. 2019; Muhammad et al. 2019). Conyza subscaposa serves for
treating obesity, breast cancer, and tonsillitis in parts of Kenya (Jeruto et al. 2008).
Conyza bonariensis to treat diarrhea in children (Bekalo et al. 2009; Giday et al.
2009) and is applied externally for skin problems (Ali et al. 2019).
Conyza pyrrhopappa is planted as an ornamental (Mekonnen et al. 2015).

References
2019. https://doi.org/10.1093/ecam/neh088.
Ali A, Aldosari A, Tng DYP, Ullah M, Hussain W, Ahmad M, Hussain J, Khan A, et al. Traditional
uses of plants by Indigenous communities for veterinary practices at Kurram District, Pakistan.
Ethnobot Res Appl. 2019;18(24):1. https://doi.org/10.32859/era.18.24.1-19.
Beentje H, editor. Flora of tropical East Africa: Compositae (Part 1). Rotterdam: Balkema; 2000.
Beentje H, Jeffrey C, Hind DJN, editors. Flora of tropical East Africa: Compositae (Part 3). Kew:
doi.org/10.1186/1746-4269-5-26.
appraisal of Kurram Agency Tribal area Pakistan. Indian J Tradit Knowl. 2019;18(4):631–47.
Central Kenya. Journal of Ethnopharmacology. 2007;111:303–307.
(STDs) and reproductive health conditions in Central Province, Kenya. Indian Journal of
Traditional Knowledge. 2009;8(2):262–269.
Njoroge GN, Bussmann R, Gemmill B, Newton LE, Ngumi VW. Antimalarial ethnophy-
totherapeutic resources in Central Kenya: An important component for homegardens and
agroforestry systems in a changing society. Abstract, 9th ISE Congress, Kent, p. 88; 2004.
Cordia africana Lam.
Cordia monoica Roxb.
Cordia sinensis Lam.
BORAGINACEAE

Grace N. Njoroge
Synonyms
Cordia africana Lam.: Calyptracordia abyssinica (R. Br.) Friesen; Cordia

abyssinica R. Br.; Cordia amplifolia A. DC.; Cordia harara Beck; Cordia
unyorensis Stapf; Gerascanthus africanus (Lam.) Borhidi; Gerascanthus holstii
(Gürke) M. Kuhlm. & Mattos; Lithocardium abyssinicum Kuntze; Lithocardium
amplifolium (A. DC.) Kuntze; Varronia abyssinica (R. Br.) A. DC.
Cordia monoica Roxb.: Cordia bakeri Britten; Cordia diversa Thwaites; Cordia
kabarensis De Wild.; Cordia ovalis R. Br. ex A. DC.; Cordia polygama Roxb.;
Cordia quarensis Gürke; Cordia rubra Hochst. ex A. Rich.; Gerascanthus bakeri
(Britten) Borhidi; Gerascanthus dioicus (Bojer ex A. DC.) Borhidi; Lithocardium
leucocomum Kuntze; Lithocardium monoicum Kuntze; Lithocardium ovale (R. Br.
ex A. DC.) Kuntze
Cordia sinensis Lam.: Cordia angustifolia Roxb.; Cordia cuneata B. Heyne ex
A. DC.; Cordia gharaf Ehrenb. ex Asch.; Cordia rothii Roem. & Schult.; Cordia
subopposita A. DC.; Gerascanthus gharaf (Ehrenb. ex Asch.) Borhidi;
R. W. Bussmann (*)
G. N. Njoroge

Gerascanthus sinensis (Lam.) Borhidi; Lithocardium rothii (Roem. & Schult.)

Kuntze; Quarena sinensis (Lam.) Raf.
Local Names
Cordia africana: Kikuyu: Muringa; Swahili: Makobokobo, Mringamringa;

Luhya: Mukumari; Borana: Waddessa; Kamba: Mvutu; Kisii: Mukobokobo;
Meru: Muzigio, Muringa; Nandi: Samutet; Samburu: Lboringo; Sebei:
Mugengere, Mugunguret; Taita: Muringaringa; Tugen: Samuet; Swahili:
Makobokobo; Luganda: Mukebu (Beentje 1994; Gachati 1989; Kokwaro 2009);
English: Large-leaved cordia, East African cordia, Sudan teak.
Cordia monoica: Kikuyu/Meru: Mukuo, Muthigi; Swahili: Msasa; Borana: Qotte,
Mader; Kamba: Muthii, Muthia; Luo: Oseno; Meru: Ikuo; Kipsigit: Nogirwet;
Oromo: Araba; Pokot: Toporerwo; Samburu: Lmantume, Seeki; Somali: Marer
giorgir, Marer gom; Turkana: Etuntun, Elkaisekiseki; Sukuma: Mnyage, Membu
Cordia sinensis: Borana/Orma: Mader; Gogo: Mudawi
Botany and Ecology
Cordia africana Lam.: Small to medium-sized deciduous tree, up to 24 m tall, trunk

usually forking a few meters from the base, bark gray to dark brown, shallow
fissured, crown spreading, dome-shaped, branchlets glabrous, powdery or pubes-
cent, sometimes hairy. Leaves alternate or sometimes almost ternate, petiole (1.5)
2.5–10.5 cm long, canaliculate, glabrous or tawny-powdery and sometimes also
tawny-pubescent, lamina 6–21 cm 4.0–16.5 cm, ovate or broadly ovate, some-
times subcircular or elliptic, glabrous or on the youngest leaves scabrous or minutely
scaly above, powdery and often tawny-hairy on the nerves and veins below, usually
rounded and suddenly shortly acuminate, sometimes acute at the apex, subacute to
cordate and often asymmetrical at the base, with entire or shallowly crenate margins,
usually stiffly coriaceous, with 5–7 secondary nerves on each side of the midrib,
tertiary nerves perpendicular to the secondary ones, both prominent below. Cymes
arranged in terminal leafy usually ample panicles, rachis and branches tawny or
brown, minutely tomentose. Flowers hermaphrodite, on pedicels up to 1 mm long.
Calyx 7–9 mm long and 4–7 mm wide at the mouth, tubular, with 10–12 well
marked ridges, coriaceous, minutely tomentose outside, glabrous inside, opening by
an operculum and later splitting into 3–5 teeth, sometimes becoming bilabiate.
Corolla 17–21 (24) mm long, funnel-shaped, white, tube 15–19 (22) mm long,
lobes 1.5–3.0 10–13 (15) mm, retuse and with a pubescent mucron 1 mm long,
margin undulate. Stamens enclosed, inserted at 3–5 mm from the base of the corolla,
anthers 2.0–2.5 mm long, filaments 7–10 mm long, slender, with a few pellucid hairs
at the base. Ovary 2.0 1.5 mm, ovoid, glabrous, style 13–18 mm long, first forked
at 7–12 mm from the base, stigmatic branches 1.5 mm long, more or less clavate.
Cordia africana Lam. . . . 343
Fig. 1 Cordia africana

(Boraginaceae), Mt. Kenya
National Park, Kenya. (Photo:
R. W. Bussmann)
Fruit 12 8 mm, ovoid, ellipsoid or obovoid, apiculate, glabrous, dark brown,

surrounded at base by the widened cup-shaped calyx, pyrene sub-quadrangular in
cross section. Seeds 1–2, very rarely 3–4. Cordia africana is widespread, from
Guinea east to Eritrea, Ethiopia, and Kenya, and south to Angola, Zimbabwe,
Mozambique, and northern South Africa. It is also found in Saudi Arabia and
Yemen, and has been planted in many tropical countries, including Réunion and
Mauritius (Verdcourt 1991). Characteristic for dry, often Croton megalocarpus,
dominated Afromontane forests (Bussmann 2002) (Figs. 1, 2, 3, 4, 5, and 6).
Cordia monoica Roxb.: Much-branched spreading shrub or small tree branched
from the base, 1.5–8 (–15) m tall, crown usually spreading, bark rather rough, gray,
but sometimes smooth and flaking in long strips, branches longitudinally striate,
with ferruginous indumentum and often velvety when young, later glabrescent,
with short internodes and prominent reniform or semicircular petiole-scars. Leaves
alternate, blades elliptic, ovate, obovate or rounded, (1–) 2.5–8 (–13) cm long,
(1–) 2–9 cm wide, rounded, apiculate or often emarginate at the apex, less often
acute, cuneate to rounded at the base, entire, denticulate or rarely distinctly repand-
dentate, rugose and mostly very scabrid above with minute tubercle-based hairs
together with sparse branched hairs, densely softly white pubescent beneath,
venation reticulate beneath, often reddish, petiole (0.5–) 1.2–5 cm long, brownish
tomentose. Flowers fragrant, hermaphrodite or unisexual in terminal or axillary
panicles of cymes 2–6 cm long, few-flowered, the axes fulvous-tomentose, pedicels
articulate at the apex, 0.2–0.8 (–3) mm long. Male flowers: filaments either 6 mm
long or 2–3 mm long in what appears to be different forms, corolla and stamens 5 (–
6)-merous, gynoecium absent. Female and hermaphrodite flowers: filaments 2–3 mm
long, corolla 4–5-merous, ovary ovoid, 3–3.5 (–5) mm long, 1.5–1.8 mm wide,
glabrous, style exserted, 6–10 mm long, stigmatic branches 3–5 mm long, recurved.
Calyx tubular-campanulate, 6–8.5 mm long, 5–8 mm wide, tomentose outside with
simple and branched hairs, glabrous or sparsely pubescent inside, 3–5-lobed or

R. W. Bussmann)

R. W. Bussmann)
Fig. 4 Cordia
sp. (Boraginaceae),
Kenya. (Photo: R. W.
Bussmann)
Fig. 5 Cordia
sp. (Boraginaceae),
Bussmann)
sometimes 2-lipped or irregularly torn. Corolla greenish white to pale yellow,

1–1.2 cm wide, tube cylindric, 4–7 mm long, lobes 3–5, oblong to narrowly obovate,
3.5–5.5 (–6.5) mm long, 1–2.5 mm wide, spreading. Fruits yellow or orange but also
reported to be olive-black to black, ovoid or obovoid, 0.9–2 cm long, 0.6–1.2 cm
wide, sitting in the persistent 1 cm wide accrescent calyx, 3–4-locular but usually
1-seeded by abortion. Very catholic, from quite wet evergreen forest to Acacia
woodland, Acacia-Commiphora bushland, and Acacia-Euphorbia thicket in grass-
land, coastal thicket, etc., often riverine, 0–1825 m (Verdcourt 1991). Characteristic
for dry, often Croton megalocarpus, dominated Afromontane forests (Bussmann
2002).
Cordia sinensis Lam.: A shrub usually up to 4 m or a bushy tree up to 8 m high,
branchlets angular, fulvous-tomentellous to subglabrous when young, glabrescent,
often cream becoming gray or brown and lenticellate with age. Leaves opposite,
Fig. 6 Cordia
sp. (Boraginaceae),
Bussmann)
subopposite or alternate, petiole 0.3–1.0 (1.5) cm long usually with some long pale
hairs, lamina 3–9 cm 1.0–4.5 cm, narrowly obovate to elliptic, rarely oblanceolate,
with minute bulbous-based appressed or slightly ascending hairs and sometimes
with scattered white bristles to subglabrous but often dotted with groups of cystoliths
above, with minute bulbous-based hairs appressed in all directions or pubescent to
subglabrous but usually with dense long hairs on the axils of the secondary nerves
and along the midrib (sometimes not much evident) below, obtuse to rounded and
sometimes retuse at apex, acute to obtuse and often slightly asymmetrical at base,
margins entire to crenate or serrate to the upper half, papery to leathery, grayish-
green or yellowish-green, with 5–7 secondary nerves on each side of the midrib.
Panicle 1.5–6.0 cm long, little-branched, terminal or axillary, rachis and branches
minutely fulvous-tomentose with scattered long pale hairs to pubescent, often
glabrescent. Flowers male on pedicels 0.5–1.5 mm long. Calyx 4–5 mm long,
narrowly campanulate, irregularly 3–5-toothed, pubescent or minutely fulvous-
tomentose outside, strigose inside. Corolla white, glabrous, tube 2.5–4.0 mm long,
cylindrical, lobes 4, 3.0–4.0 mm 1.2–2.0 mm, oblong to narrowly obovate,
rounded at apex, reflexed. Stamens 4, inserted at the corolla-throat, filaments
2–3 mm long, glabrous, anthers 1.5 mm long, oblong. Ovary 2 mm long, ovoid,
glabrous, style 5–7 mm long, first-forked at 2.0–2.5 mm and with stigmatic branches
2–3 mm long, clavate. Fruit 12–15 mm 8–11 mm, ovoid or ellipsoid, apiculate,
glabrous, yellow, orange or reddish when ripe, surrounded at the lower third by the
widely enlarged, usually cream, campanulate calyx, mesocarp fleshy, pyrene

c. 8 7 mm, rhomboidal or quadrangular in cross section, 1–3 seeded (Verdcourt
1991). Characteristic for dry, often Croton megalocarpus, dominated Afromontane
forests (Bussmann 2002).
Cordia africana: Leaf decoctions are administered to treat headache, nose bleeding,
dizziness, and vomiting during pregnancy, wounds, and worms. Fresh bark is
applied to fractures and bark extracts and taken against fatigue. Root decoctions
are drunk to treat jaundice, venereal diseases, and schistosomiasis (Kokwaro 2009).
Wood ash is applied to skin diseases. Used as vitamin supplement and to treat
measles (Muthee et al. 2011).
Cordia monoica: Leaves are used for earache. Pounded roots are applied to wounds.
Steam baths are used to remedy leprosy, and the leaf paste is also applied as poultice.
The root decoction is used for malaria and vomiting in children. The leaf extract is
given to expel the placenta (Kokwaro 2009). Used in fractures as splint (Bussmann
2006). Used for back pain and gonorrhea (Muthee et al. 2011). The species is used in
Ethiopia to treat the febrile condition “mich” and for eye diseases (Giday et al. 2003).
Cordia sinensis: Roots are boiled with milk and drunk for malaria. Larger amounts
of chewed roots are used for abortion. Bark and root decoction are used to remedy
stomachache (Kokwaro 2009). The roots are boiled for chest pain (Lulekal et al.
2008; Teklehaymanot and Giday 2010).
Cordia ovalis is used in Ethiopia to treat the febrile condition “mich” (Wondimu
et al. 2007).
Local Food Uses
Cordia africana: The fruit is edible (Beentje 1994)

Cordia monoica: The fruit is edible (Beentje 1994; Bussmann 2006)
Cordia africana: The wood is commonly used for joinery, interior trim, paneling,
furniture, cabinet work, drums, beehives, boxes, mortars, and canoes. The gum is
used as glue (Beentje 1994). It is suitable for light construction, ship building,
vehicle bodies, toys, novelties, vats, draining boards, food containers, matches,
veneer, plywood, hardboard, particle board, and pulp for paper making. It is also
used as firewood. The leaves serve as fodder for livestock. Cordia africana is planted
as a shade tree in coffee plantations and as roadside tree, ornamental shade tree, and
boundary marker. The leaves make a good mulch. The flowers provide nectar for
honeybees. Sold as expensive timber species (Balemie and Kebebew 2006; Bekalo
et al. 2009).
Cordia monoica: Leaves are used as sandpaper (Beentje 1994). The leaf extract is
given to livestock to expel the placenta. Used in rituals (Luo, Samburu) (Kokwaro
2009). Eaten by livestock, used for construction, fences, firewood, and ceremonial
(Bussmann 2006). In Madagascar, it is used for building, construction of carts, and
for blessings during circumcision (Randrianarivony et al. 2016).
Cordia sinensis: Root decoction to treat conjunctivitis in cattle. The bark infusion is
dripped in infected eyes of cattle. Ash from twigs is applied to infected udders with
mastitis (Kokwaro 2009). To treat corridor disease in livestock, the roots are boiled
and the extract is given to the animal (Kioko et al. 2015).
Cordia curassavica leaves are used as tick repellent in livestock (Wanzala 2017;
Wondimu et al. 2007)
References
doi.org/10.1186/1746-4269-5-26.
10.1186/1746-4269-4-10.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016. (ISBN: 978-0-9960231-4-6).
Verdcourt B. Flora of Tropical East Africa: Boraginaceae. Rotterdam: Balkema; 1991.

Crotalaria agatiflora Schweinf.
Crotalaria fascicularis Polhill
Crotalaria incana L.
Crotalaria natalitia Meisn.
FABACEAE

Grace N. Njoroge
Synonyms
Crotalaria agatiflora Schweinf.: Crotalaria dawei Baker f.; Crotalaria engleri

Harms ex Engl.; Crotalaria erlangeri (Baker f.) Harms. ex Hutch. & Bruce;
Crotalaria imperialis Thunb.; Crotalaria megistantha Taub.
Crotalaria incana L.: Chrysocalyx schimperi Hochst. ex A. Rich.; Crotalaria affinis
DC.; Crotalaria criocaula S. Schauer; Crotalaria cubensis DC.; Crotalaria
cytisifolia Hoffmanns. ex Steud.; Crotalaria diffusa Link; Crotalaria diffusa Vell.;
Crotalaria eriocaula S. Schauer; Crotalaria glabrescens Andersson; Crotalaria
glabrescens Benth.; Crotalaria herbacea Schweigger ex Schrank; Crotalaria hirta
Lag.; Crotalaria hirta Roth; Crotalaria incana fo. glabrescens R. Wilczek;
Crotalaria incana fo. microphylla Chodat & Hassl.; Crotalaria incana subsp.
purpurascens (Lam.) Milne-Redh.; Crotalaria incana var. australis Grieseb.;
Crotalaria megapotamica Burkhart; Crotalaria montana A. Rich.; Crotalaria
picensis Phil.; Crotalaria pubescens Hort. ex Steud.; Crotalaria pubescens Moench;
R. W. Bussmann (*)
G. N. Njoroge

Crotalaria purpurascens Lam.; Crotalaria radiata Merr.; Crotalaria schimperi

A. Rich.; Crotalaria setifera DC.; Lupinus rotundifolius Sessé & Moc.
Crotalaria natalitia Meisn.: Crotalaria gymnocalyx Baker; Crotalaria
kilimanscharica Taub. ex Engl.
Local Names
Crotalaria agatiflora: Kikuyu: Mwethia, Muchingiri; Kamba: Iviinzi; Kipsigis:

Kipkururiet; Maa: Olontwalan; Meru: Muchuguchugu (Beentje 1994; Gachati
1989; Kokwaro 2009).
Crotalaria fascicularis: Kikuyu: Muchingiri; Samburu: Letwalan; Turkana:
Emaret (Beentje 1994; Gachati 1989).
Crotalaria incana: Kikuyu: Muchingiri; Luhya: Lisilinyende (Beentje 1994;
Crotalaria natalitia: Kikuyu: Muchingiri; Maa: Olontwalan; Samburu: Celiuk;
Iraqw: Qooli (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Crotalaria agatiflora Schweinf.: Woody bushy herb, shrub or small tree, 1–10 m
tall, much branched. Seeds oblique-cordiform, 6–7(–9) mm long, smooth, pale
brown. Pod subcylindrical, gradually narrowed into the 1.5–2.5 cm long stipe,
7.5–10 cm long, 1.2–2 cm across, glabrous, many-seeded. Standard ovate, lemon-
yellow to greenish-yellow, glabrous or medially pubescent outside, wings half to
two-thirds as long as the keel, keel broadly rounded, with a relatively short pro-
jecting often greenish or purplish untwisted beak, 35–45(–55) mm long. Calyx
18–30 mm long, basally narrowed into a prominent receptacle, glabrous to pubes-
cent outside, sometimes puberulous on the lobes inside, upper and lateral lobes on
each side joined almost to the tips, twice as long as the tube. Stipules linear,
4–12 mm long, caducous, or often absent. Racemes stoutly pedunculate, up to
40 cm long, many-flowered, bracts subulate to suborbicular, acuminate to long
caudate, 5–38 mm long, 0.7–30 mm wide, caducous, bracteoles inserted on the
pedicel, filiform to attenuate-lanceolate or slightly hastate, 0.5–29 mm long, up to
8 mm wide. Twigs glabrous or pubescent to tomentose. Leaves 3-foliolate, leaflets
elliptic-lanceolate to elliptic-ovate, up to 3.8–6.7 cm long, 2–3.5 cm wide, glabrous
to pubescent above, glabrous to tomentose beneath, petiole up to 12 cm long (Brenan
1967) (Figs. 1, 2, and 3).
Crotalaria fascicularis Polhill: Shrub or bushy perennial herb, up to 1–2 m tall.
Branches tomentose, with yellowish only slightly spreading hairs, ultimately
glabrescent, bark with peeling epidermis. Leaves, except on the youngest branches,
mostly crowded on very short lateral shoots, appearing fasciculate, 3-foliolate,
leaflets oblanceolate, oblanceolate-elliptic or obovate, up to 15–37 mm long,
5–18 mm wide, glabrous above, thinly to rather densely appressed pubescent
Crotalaria agatiflora Schweinf. . . . 353
Fig. 1 Crotalaria
cf. agatiflora (Fabaceae),
Sirimon, Kenya. (Photo
R.W. Bussmann)
Fig. 2 Crotalaria
cf. agatiflora (Fabaceae),
Sirimon, Kenya. (Photo
R.W. Bussmann)
beneath, petiole 10–42 mm long. Stipules linear, 2–4(–16) mm long, rather persis-
tent. Racemes up to 4–13 cm long, with few to numerous fairly closely arranged
flowers, bracts linear, 2–5(–9) mm long, bracteoles inserted on the pedicel, filiform,
up to 1–5 mm long. Calyx 5–7 mm long, slightly protracted on the lower side,
appressed pubescent, upper lobes triangular, as long as the tube. Standard broadly
elliptic to suborbicular, yellow, becoming suffused with red or brown, glabrous
Fig. 3 Crotalaria cf. incana

(Fabaceae), Mt. Kenya
National Park, Sirimon,
Kenya. (Photo
R.W. Bussmann)
outside or puberulous along the midvein, wings nearly as long as the keel, keel
abruptly rounded below the middle, with a well-developed twisted beak,
(12–)13–16 mm long. Pod shortly stipitate, oblong-clavate, 25–30 mm long, 6 mm
across, appressed puberulous, with apparently rather few seeds matured from the 20–
24 ovules. Seeds oblique-cordiform, 3–3.5 mm long, smooth, brown. Margins of
upland rainforest and dry evergreen forest, upland grassland and evergreen bushland,
usually in drier, more exposed often rocky positions, 1950–2950 m, rarely extending
down to 1300 m along rivers (Brenan 1967).
Crotalaria incana L.: Erect or spreading usually rather bushy herb, up to 1.5(3.5) m tall,
variously hairy. Leaves 3-foliolate, leaflets up to 2.5–5 1.7–4.5 cm, elliptic-obovate to
obovate or subcircular, rounded to retuse at apex, usually thinly pilose beneath, petioles
mostly longer than leaflets, stipules 2.5–12(20) mm long, filiform. Racemes usually lax,
few-many-flowered, bracts 1–10 mm long, subulate or linear, bracteoles curved-
ascending from just below calyx, 2–6 mm long, linear. Calyx 7–11 mm long, sub-
glabrous to spreading pilose, lobes narrow, twice as long as tube. Standard elliptic,
yellow, veined reddish-brown or purple, glabrous or medially pubescent outside, wings
nearly as long as keel, keel 8–11.5 mm long, bent at right-angles in lower half, lanate-
pilose on upper margin. Pod 3–4.5 cm long, fusiform-clavate, subsessile, somewhat
pointed, pilose. Seeds 2.5–3 mm long, oblique-cordiform, smooth or obscurely papil-
lose, pale brown, mottled darker, or olive-green (Brenan 1967).
Crotalaria natalita Meisn.: Woody herb or small shrub, 1–2.6 m tall, with 1-several
thin stems and rather strictly ascending pubescent branches mostly towards the top,
Crotalaria agatiflora Schweinf. . . . 355
rarely somewhat procumbent in burned and heavily grazed places. Leaves mostly in
tufts, 3-foliolate, leaflets mostly (1.5)2–4.5 0.4–1.5 cm, linear-oblanceolate to
oblanceolate-elliptic, appressed pubescent beneath, petioles mostly a little shorter
than leaflets, stipules (3)5–15 1–4 mm, shortly stalked, linear-lanceolate to
oblong-falcate, very unequal-sided. Racemes terminal, up to 10–22 cm long, with
few-many fairly closely to rather laxly arranged flowers, bracts 2.5–6 mm long,
linear to elliptic-caudate, caducous, pedicels rather stiffly erect. Calyx 6–9(10) mm
long, upper lobes narrowly attenuate-triangular, as long as to a little longer than the
tube. Standard subcircular, yellow, sometimes reddish-brown tinged outside, fading
orange-red, glabrous outside, wings broad, exceeding the keel, keel 11–14 mm long,
abruptly rounded about the middle, with a rather short slightly incurved beak, lanate
towards the upper margin. Pod 3.4–4(4.5) 1.1–1.5 cm, broadly cylindrical,
abruptly contracted to a 4–5 mm long stipe, glabrous, often mottled dark green to
blue-black before ripening, 25–40-seeded. Seeds 3.5–4 mm long, oblong-reniform,
with a small aril, granulate or smooth, dark brown (Brenan 1967).
Crotalaria agatiflora: Root decoction is used for gonorrhea (Beentje 1994;

Kokwaro 2009) and other sexually transmitted diseases (Njoroge et al. 2004).
Used to expel retained placenta after childbirth (Yineger et al. 2008). The juice is
applied to the ear for otitis media (Njoroge and Bussmann 2006).
Crotalaria incana: Roots are chewed by expectant mothers to remedy placenta
position problems. Ash of burned roots is applied to mouth rashes. Leaf paste is
applied to wounds (Kokwaro 2009). Used for strength, coughs, and chest pain
(Bussmann 2006).
Crotalaria natalita: Root bar is chewed to treat boils (Kokwaro 2009). Used for
strength, coughs, and chest pain (Bussmann 2006).
Crotalaria pallida is used for bone injuries (Reang et al. 2016). Crotalaria pallida is
one of the most commonly used medicinal species in parts of Madagascar
(Randrianarivony et al. 2017). Crotalaria coursii and Crotalaria retusa are given
to mother for strength (Randrianarivony et al. 2016a). Crotalaria coursii is also used
for fever, yellow fever, malaria, colic, epilepsy in infants, to treat mothers after
childbirth, swelling of the nerves, and as an aphrodisiac (Randrianarivony et al.
2016b). Crotalaria xanthoclada is a remedy for wound problems, diarrhea, and
intestinal parasites (Rabearivony et al. 2015). Crotalaria rosenii is used to expel
retained placenta after childbirth (Yineger et al. 2008). Crotalaria laburnifolia helps
to treat fungal skin infections (Wondimu et al. 2007).
Crotalaria aculeata smoke is used in social settings (Mohagheghzadeh and Faridi

2006). Crotalaria rosenii is a preferred forage species for sheep and goats in
Ethiopia (Bussmann et al. 2011).
References
Brenan JPM. Flora of tropical East Africa: Leguminosae, subfamily Caesalpinioideae. London:
Crown Agents for Overseas Governments & Administrations; 1967.
Madagascar. Indian J Tradit Knowl. 2016a;15(1):68–78.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016b. ISBN: 978-0-9960231-4-6
2017. https://doi.org/10.1186/s13002-017-0147-x.
Aromat Plants. 2016;5(2):234.
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plants Res. 2008;
2(6):132–53.
Croton dichogamus Pax
Croton macrostachyus Hochst ex Delile
Croton megalocarpus Hutch.
EUPHORBIACEAE

Grace N. Njoroge
Synonyms
Croton macrostachyus Hochst ex Delile: Croton acuminatus R. Br.; Croton

butaguensis De Wild.; Croton guerzesiensis Beille; Croton guerzesiensis Beille ex
A. Chev.; Croton macrostachyus var. mollissimus Chiov; Oxydetes macrostachya
(Hochst. ex Delille) Kuntze; Rottlera schimperi Hochst. & Steud.
Local Names
Croton dichogamus: Kikuyu: Kererwa, Kireru, Mukunduri; Borana: Mokhof;

Gabbra: Mokofe; Kamba: Muthiani; Kipsigis: Kelelwet; Luo: Angin, Rachar;
Maa/Samburu: Ol-logerdangai; Mijikenda: Mwarula; Samburu: Lageridingai;
Somali: Gobole; Turkana/Pokot: Kekelwa; Swahili: Msenefu; Mbulu: Girigirimu
(Beentje 1994; Gachati 1989; Kokwaro 2009); English: Croton
R. W. Bussmann (*)
G. N. Njoroge

Croton macrostachyus: Luhya: Musudzu, Musutsu; Kikuyu/Meru: Mutundu;

Hehe: Muwulugu; Luganda: Musogasoga; Borana: Mukanisa; Kamba: Kitundu;
Kipsigis/Nandi/Tugen: Tebeswet; Luo: Ngong’Ngo; Luhya: Musutsu; Maa:
Orkeparlu; Marakwet: Taboswa, Toboswa; Taita/Taveta: Mifirifiri (Beentje
1994; Kokwaro 2009).
Croton megalocarpus: Kikuyu: Mutundu-wa-Njora; Meru: Mutundu; Swahili:
Msenefu; Borana: Mukanisa; Kamba: Kitundu, Muthulu; Kipsigis/Nandi/
Tugen: Tebeswet; Luo: Ngong’Ngo’; Luhya: Musutsu, Musine, Mutsuitsui;
Maa: Orkeparlu, Ol-Mergoit; Marakwet: Taboswa, Toboswa; Taita/Taveta:
Mfirifiri; Chagga: Lalei; Luguru: Mbali; Luganda: Nkulumire (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: Croton
Botany and Ecology
Croton dichogamus Pax: A monoecious pyramidal or sometimes straggling shrub

or small tree up to 7.5 m tall, but more usually 2–5 m tall, with symmetrical, frequent
branching. Young twigs densely silvery or brown lepidote, later becoming grayish.
Peltate scales dissected or fimbriate. Petioles 0.5–2( 3) cm long, silvery or brownish
lepidote, leaf-blade ovate-lanceolate to elliptic-lanceolate, more rarely ovate, 2–12
( 16) cm long, 1–5( 7.5) cm wide, usually acuminate, often acutely so, less often
obtuse, rounded at the base, or sometimes subcuneate or truncate-subcordate, with
basal glands subsessile or shortly stipitate, fairly prominent, entire, firmly membra-
nous, lateral nerves 9–15 pairs, slightly prominent beneath, midrib impressed above,
prominent beneath, glabrous and dark green above except for a strip of sparse,
adpressed scurfy stellate hairs toward the margin when young, shiny silvery lepidote
and sometimes brown flecked beneath. Stipules triangular-lanceolate, 1 mm long,
somewhat obscured by the scales, caducous. Axillary buds not perulate. Racemes
terminal, up to 6 cm long, but more often 2–4 cm long, female in the lower third,
male in the upper two-thirds, sometimes all male, occasionally mostly female, axis
brown lepidote, bracts triangular-lanceolate, 1.5 mm long, lepidote with fimbriate
scales, ciliate. Male flowers: pedicels 3–5 mm long, slender, yellow-brown lepidote,
sometimes bibracteolate, sepals 5 or 6, broadly ovate, 2 mm long, 2 mm wide,
obtuse, yellow-brown lepidote without, glabrous within, margin finely ciliate, pale
yellow-green, petals 4 or 5, oblanceolate-oblong, 2 mm long, 0.5 mm wide, margin
finely ciliate, otherwise glabrous, disc pentagonal, stamens (13–)15( 20), pale
orange, filaments 2.5 mm long, glabrous above, pubescent below, anthers 0.5 mm
long, receptacle pilose. Female flowers: pedicels 2–4 mm long, stouter than in the
male, yellow-brown lepidote, sepals as in the male, petals 0 or, if present (5 or
fewer), then linear, not more than 0.5 mm long, ciliate and readily caducous, disc
annular, ovary 3( 4)-lobed, 2 mm diameter, densely brown lepidote, styles
3, spreading, 2 mm long, glabrous, deeply bipartite. Fruit 3( 4)-lobed, often some-
what parallel-sided, 6–7 mm long, 6–7 mm diameter, septicidal, evenly brown
lepidote. Seeds oblong-ovoid, 5 mm long, 3 mm wide, brown, with a convex
caruncle 1.5 mm wide. Dry forest, bushland and thicket, on rocky ground, lava
Croton dichogamus Pax. . . 359
Fig. 1 Croton
cf. dichogamus
(Euphorbiaceae), Sekenani,
R.W. Bussmann)
Fig. 2 Croton macrostachyus (Euphorbiaceae), interior of Croton macrostachyus dominated

forest, Karura forest, Nairobi, Kenya. (Photo R.W. Bussmann)
Fig. 3 Croton megalocarpus

(Euphorbiaceae), exterior
aspect of Croton
megalocarpus dominated
forest, Mathews Range,
Kenya. (Photo
R.W. Bussmann)

(Euphorbiaceae), Mathews
Range, Kenya. (Photo
R.W. Bussmann)
and porous soils, 550–2000 m (Smith 1987; Smith and Carter 1988) (Figs. 1, 2, 3, 4,
and 5).
Croton macrostachyus Hochst ex Delile: A dioecious or sometimes monoecious
shrub or spreading tree up to 25 m. tall, although more commonly 6–12 m tall. Male

(Euphorbiaceae), Mathews
Range, Kenya. (Photo
R.W. Bussmann)
flowers: pedicels 4–10( 14) mm long, densely stellate-pubescent, sepals 5, ovate,

3 mm long, 2.5 mm broad, obtuse, stellate-pubescent without, pubescent within at
the middle and towards the apex, pale green, petals 5, oblanceolate-oblong, 3.5 mm
long, 1.5–2 mm wide, obtuse, glabrous without, densely villous within and on the
margin, pale yellow, disc-glands rounded, hairy, stamens 15–20, filaments 4 mm
long, villous in the lower half, anthers 1 mm long, receptacle densely villous.
Racemes terminal, (7–)15–32 cm long, up to 3 cm wide, either all male or all female,
or mostly male with a few female flowers at the base, or sometimes half male and
half female or mostly female with a few male flowers at the top, the male flowers in
clusters along the axis, the female’s usually solitary within the bracts, or with
1 female flower accompanied by 1 or more male flowers per cluster, axis densely
stellate-strigose, or evenly or sparingly stellate-pubescent, bracts linear, 2–3( 5)
mm long, sparingly pubescent or subglabrous. Fruit subtrilobed, rarely 4-lobed,
often more particularly so at the apex, 8–9 mm long, 8–10( 15) mm wide, loculi-
cidal, evenly stellate-pubescent-sublepidote. Female flowers: pedicels 2–5( 8) mm
long, stouter than the male pedicels, scarcely elongating in fruit, densely fulvous
stellate-pubescent, sepals 5, lanceolate or ovate-lanceolate, 3 mm long, 1.5 mm
broad, obtuse, evenly stellate-pubescent without, sparingly stellate-pilose within,
petals 0, or, if present, linear and smaller than in the male flowers, staminodes 0–1
( 2), filament-like, disc-glands free, truncate, ovary trigonous, 2 mm diameter,
densely fulvous adpressed stellate-pubescent-sublepidote, styles 3( 4), spreading,
2.5 mm long, pubescent at the base, otherwise glabrous, deeply bipartite, the lobes
linear, sometimes bifid at the apex, 2 mm long. Older twigs grayish brown. Young
twigs sparsely, evenly or densely grayish adpressed stellate-pubescent, the hairs
often with a projecting central ray. Stipules linear or linear-subulate, 5–14 mm long,
entire or with a few subulate lateral lobes, sparingly stellate-pubescent or sub-
glabrous, caducous. Petioles 1.5–11 cm long, usually sparingly stellate-pubescent,
sometimes either densely so or almost glabrous, leaf-blade broadly ovate to elliptic-
ovate, 4, 15( 20) cm long, 2.5–10( 13) cm wide, obtusely or subacutely acumi-
nate, rounded or cordate, with 2–4( 8) subsessile or stipitate discoid basal glands
visible or not from above, crenulate-serrulate to subentire, thinly to firmly
chartaceous, 5–7( 9)-nerved from the base, remaining lateral nerves 4–8( 11)
pairs, usually fairly prominent beneath, densely fulvous (above) and grayish
(beneath) stellate-pubescent at first, later becoming evenly or sparingly adpressed
stellate-pubescent and dark green above, and evenly or sparingly grayish hirsute-
stellate-pubescent beneath. Seeds ellipsoid, 7 mm long, 4 mm wide, longitudinally
rugulose, gray, somewhat shiny, with a large waxy caruncle 4.5 mm long and 4 mm
wide (Smith 1987; Smith and Carter 1988). Easy to germinate under light conditions
but does not store well (Bussmann and Lange 2000). Common in drier, broad-leafed
Afromontane forests, dominated by Cassipourea malosana. Ilex mitis, a character
species, often reaching high up into the uppermost tree layer, together with Olea
capensis ssp. hochstetteri. and with Teclea nobilis in the lower tree and shrub stratum
(Bussmann 2002, 2004, 2006a).
Croton megalocarpus Hutch.: A large monoecious or occasionally dioecious tree
up to 35 m tall, though commonly 15–25 m, with a clear bole up to 20 m high and
1 m diameter and a spreading crown, bark pale gray-brown, closely longitudinally
fissured. Young twigs densely fulvous lepidote to fulvous or ferruginous stellate-
lepidote. Older twigs grayish brown, tardily glabrescent. Petioles 2–8( 11) cm long,
densely fulvous lepidote to fulvous or ferruginous stellate-lepidote, leaf-blade ovate,
elliptic-ovate, elliptic-lanceolate or oblong-lanceolate, (4–)7–14( 19) cm long,
(1.5–)3–7( 11) cm wide, shortly and acutely or subacutely acuminate, cuneate-
rounded, rounded or shallowly cordate, with 2( 4) sessile or stalked basal glands
visible or not from above, entire, firmly chartaceous to subcoriaceous, lateral nerves
15–25 pairs, not or slightly impressed above, slightly prominent beneath, midrib
impressed above, prominent beneath, evenly minutely stellate-puberulous and
medium to dark green above, densely silvery lepidote or rarely stellate-lepidote
and brown flecked beneath, the scales fimbriate. Stipules linear-filiform, sometimes
somewhat laciniate towards the base, 5–10 mm long, sparingly scurfily stellate-
lepidote, subpersistent. Racemes terminal, 7.5–30 cm long, all male or with a few
female flowers at the base, axis angular, densely fulvous lepidote to fulvous or
ferruginous stellate-lepidote, bracts linear-lanceolate to elliptic-lanceolate,
2–4.5 mm long, densely stellate-lepidote, soon caducous. Male flowers: pedicels
7–15 mm long, densely silvery lepidote, often brown flecked, calyx 5-lobed, the
lobes triangular-ovate, 4.5 mm long, 3.5 mm wide, subacutely acuminate, densely
silvery lepidote and brown flecked without, densely pubescent within, somewhat
villous on the margin at the apex, greenish yellow, petals 5, obovate, 5 mm long, 3.5 mm
wide, densely silvery lepidote and sparingly light-brown flecked without, pubescent
within, somewhat villous on the margin, pale straw-yellow, disc-glands free, truncate,
stamens 25–30( 40), filaments 5–7( 8) mm long, glabrous above, sparingly pilose
below, anthers 1.5–2 mm long, receptacle pubescent. Female flowers: pedicels 4–12 mm
long, up to 2 cm long in fruit, stouter than in the male, calyx-lobes 3.5 mm long, 2 mm
wide, otherwise as in the male, petals 5, linear, 5 mm long, 0.5 mm wide, lepidote
without in the lower part, otherwise subglabrous, pubescent within and villous on the
margin, disc shallowly 5-lobed or annular, ovary subglobose-subtrilobed, 4 mm
diameter, densely silvery lepidote and brown flecked, styles 3, spreading, 1.5 mm
long, subglabrous above, densely stellate-lepidote beneath, shortly bipartite, the seg-
ments 2–4-lobulate. Fruit ellipsoid-ovoid to subglobose, not lobed, 3–4.5 cm long,
2.5–3( 4) cm diameter, loculicidal from the apex, lepidote, endocarp woody, 3–
5.5 mm thick, with three double rows of pits outside corresponding to the septa.
Seeds ellipsoid-ovoid or oblong-ellipsoid, (1.8–)2.2–2.4 cm long, 1.2–1.4 cm wide,
somewhat ridged and rugulose, white when fresh, gray-brown when dry, with a minute
caruncle. Evergreen forest, (700–)1200–2400 m. Croton megalocarpus occurs from
eastern DR Congo east to Kenya and south to Malawi, Zambia, and Mozambique
(Smith 1987; Smith and Carter 1988). Easy to germinate under light conditions but does
not store well (Bussmann and Lange 2000). Croton megalocarpus forms very specific
forests especially on the foot slopes of mountains in Ethiopia and Northern Kenya, e.g.,
at the lower extensions of Mathews Range, from 1500 to 2200 m. In these semi-
deciduous forests, Croton megalocarpus forms an open canopy. Due to the high amount
of light reaching the ground, a very diverse herbal flora is encountered in these forests,
which also have a dense shrubby undergrowth. In the dry season, when Croton sheds its
leaves, these forests are flooded with light, with Panicum monticola and Setaria
plicatilis are then forming dense soft carpets on the forest floor. The shrub layer is
dominated by Erythrococca bongensis, Flacourtia indica, Schrebera alata, and
Phyllanthus fischeri. These forests are extremely peculiar and rich in endemic species.
For example, in Mathews Range in Kenya, the most striking species in Croton
megalocarpus dominated forests is Encephalartos tegulaneus. Growing up to 8–10 m
tall, the palm-like Encephalartos gives these forests a primeval appearance (Bussmann
2002, 2004, 2006a, b).
Croton dichogamus: Roots and sometimes leaves are used in traditional medicine to
treat colds, fever, tuberculosis and syphilis, as a tonic, for stomachache and for
cough (Beentje 1994; Kokwaro 2009). Used to treat malaria, stomach problems, and
chest problems (Bussmann 2006a, b). Used as toothbrush (Bussmann et al. 2006).
Used in Ethiopia to treat malaria, rabies, gonorrhea, and wounds (Giday et al. 2007;
Lulekal et al. 2008), as well as snakebites (Giday et al. 2009).
Croton macrostachyus: Leaves are boiled and drunk for cough. The root decoction
is used as anthelminthic and as purgative. Leaf juice is applied to stop hemorrhages,
and the boiled leaf juice is used for malaria and venereal diseases. Newborn babies
are bathed in a bark decoction against skin rashes. A bark decoction is used for flu
(Kokwaro 2009). Used for wound treatment, sores, warts, and especially ringworm
(Njoroge and Bussmann 2007). Used for the treatment of scabies, hepatitis, jaundice,
and diarrhea (Teklehaymanot et al. 2007, 2009; Wondimu et al. 2007; Yineger et al.
2007, 2008).
Croton megalocarpus: A maceration or decoction of the bark is taken as a vermi-
fuge and to treat whooping cough, pneumonia, stomachache, fever, including
malaria, and abdominal complaints associated with gall bladder and spleen prob-
lems. Sap from leaves and young twigs are applied to wounds (Beentje 1994;
Kokwaro 2009). Also used as anthelminthic and for whooping cough. Root decoc-
tion is used to treat pneumonia (Kokwaro 2009). An infusion of the powdered bark
with potash is given to goats as a conditioner. It is one of the most popular medicinal
species in parts of Ethiopia (Bekalo et al. 2009; Giday et al. 2003). Used as
antimalarial (Njoroge and Bussmann 2006a). The roots and bark are used to treat
cough and tonsillitis (Njoroge and Bussmann 2006b), and for wounds (Njoroge and
Bussmann 2007). Also used to treat back pain in women (Njoroge and Bussmann
2009). Sometimes used for respiratory conditions (Muthee et al. 2011).
Local Food Uses
Croton dichogamus: The roots and stems are used to flavor food and drinks, and the
stems are used to ferment traditional beer (Beentje 1994). Sometimes dried leaves
are chewed or smoked like tobacco (Kokwaro 2009). Used to flavor local beer and to
give meat more “strength” (Bussmann et al. 2006).
Croton macrostachyus: Used as firewood, for household tools, and as shade species
(Mekonnen et al. 2015). Widely used in veterinary medicine (Wondimu et al. 2007).
Latex sometimes used as fish poison (Neuwinger 2004).
Croton dichogamus: The wood is used in construction, for tools, and ax handles
(Beentje 1994). Used as firewood (Bussmann 2006a, b). Latex is sometimes used as
fish poison (Neuwinger 2004).
Croton megalocarpus: The wood is used for construction, flooring, stools, mortars,
beehives, veneer, and plywood. It is suitable for joinery, interior trim, ship building,
vehicle bodies, furniture, cabinet work, railway sleepers, and agricultural imple-
ments. It is also used as firewood and for charcoal production (Beentje 1994). Bark
decoction is used to treat anthrax and East Coast Fever (Kokwaro 2009). Although
trees are usually not browsed by livestock, leafy twigs may serve as forage for goats.
The seeds can be used to dye wool yellowish. The flowers provide nectar for
honeybees, and the honey produced is dark and has a strong flavor. The species is
planted in hedges, live fences, shelterbelts, and windbreaks, and as an ornamental
shade tree. It may serve as shade tree for coffee plantations. The foliage provides
good mulch. The fruit shells are used as mulch in vegetable gardens and as a
component of potting mixtures. Sometimes used as firewood (Kiefer and Bussmann
2008). Latex is sometimes used as fish poison (Neuwinger 2004).
References
Bekalo TH, Demissew Woodmata S, Asfaw WZ. An ethnobotanical study of medicinal plants used
by local people in the lowlands of Konta Special Woreda, southern nations, nationalities and
peoples regional state, Ethiopia. J Ethnobiol Ethnomed. 2009;5:26. https://doi.org/10.1186/
1746-4269-5-26.
Mountain Forests: The role of natural and anthropogenic disturbance. Lyonia. 2004;6(1):98–111.
2006b;111:41–66.
Nat Hist. 2000;891–2:101–11.
Sekenani Valley, Maasai Mara, Kenya. Econ Bot. 2018;722:207–16.
10.1186/1746-4269-4-10.
Njoroge GN, Bussmann RW. Ethnotherapeutic management of sexually transmitted diseases STDs
and reproductive health conditions in Central Province, Kenya. Indian J Tradit Knowl.
2009;82:262–9.
Smith AR. Flora of tropical East Africa: Euphorbiaceae (Part 1). Rotterdam: Balkema; 1987.
1988.
10.1186/1746-4269-4-11.
Cucumis dipsaceus Ehrenb. ex Spach
CUCURBITACEAE

Grace N. Njoroge
Synonyms
Cucumis dipsaceus Ehrenb. ex Spach: Cucumis ambigua Fenzl ex Hook. f.;

Cucumis bardana Fenzl ex Naudin; Cucumis dasycarpa Hochst. ex A. Rich.
Local Names
Cucumis dipsaceus: Pokot: Alaskau; Sukuma: Limbiligulu; Swahili: Mtango;

Mwitu, Luo: Nyabuth-Mwok.
Botany and Ecology
Cucumis dipsaceus Ehrenb. ex Spach: Fibrous-rooted annual climbing or trailing

herb, stems angular, rather gray-green, hispid with stiff bristly spreading hairs
mainly on the ridges. Leaf-blade ovate- or reniform-cordate in outline, rounded,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Cucumis dipsaceus

(Cucurbitaceae), Pusac,
R.W. Bussmann and
regularly sinuate-toothed, scabrid hairy above and especially on the nerves beneath,
18–95 mm long, 27–102 mm wide, unlobed or shallowly and broadly 3-lobed, with
the lobes rounded, petiole roughly spreading-hairy with broad-based pointed hairs,
17–115 mm long. Male flowers 1–4, on 7–13 mm long pedicels, receptacle-tube pale
green, 3.5–5 mm long, lobes subulate-filiform, 1.5–4 mm long, petals yellow with
pellucid green veins, 5–10 mm long, 2.5–5 mm broad, united below. Female flowers
on 5–10 mm long stalks, ovary ellipsoid, bristly, 9–17 mm long, 4–8 mm broad,
receptacle-tube green, 3–4 mm long, lobes subulate, spreading, 2.5–5 mm long,
petals 6–15 mm long, 3–8.5 mm broad, rounded and apiculate. Male flowers 1–4, on
7–13 mm long pedicels, receptacle-tube pale green, 3.5–5 mm long, lobes subulate-
filiform, 1.5–4 mm long, petals yellow with pellucid green veins, 5–10 mm long,
2.5–5 mm broad, united below. Female flowers on 5–10 mm long stalks, ovary
ellipsoid, bristly, 9–17 mm long, 4–8 mm broad, receptacle-tube green, 3–4 mm
long, lobes subulate, spreading, 2.5–5 mm long, petals 6–15 mm long, 3–8.5 mm
broad, rounded and apiculate. Fruit on a stout 8–11 mm long stalk, ellipsoid,
43–66 mm long, 28–40 mm across, densely and softly spiny, with each spine ending
in a hyaline bristle, pale green, yellow when ripe. Seed elliptic, rather pointed at each
end, 4.5 2 1 mm. Bushland, woodland and wooded grassland, also a weed of
cultivation, 425–1800 m (Jeffrey 1967) (Fig. 1).
Cucumis dipsaceus: Fruits are crushed and boiled for stomach pain. Pounded roots
and leaves are applied as poultice to wounds. The fruit decoction is used as purgative
in case of poisoning. The leaf decoction is drunk to speed up childbirth (Kokwaro
2009). Used also to treat dandruff, hair loss (prevention), and stopping baby from
breastfeeding (Bussmann and Sharon 2006, 2015). In Ethiopia, it is used to treat
snake- and hyaena bites, fox bites, cough, and as diuretic (Lulekal et al. 2008).
Cucumis ficifolius fruits are crushed and applied to wounds, and a root decoction
is drunk for sudden collapse and stomach problems (Bussmann et al. 2011). Also
Cucumis dipsaceus Ehrenb. ex Spach 369
used for chest pain (Giday et al. 2003). Employed for meningitis, epistaxis, rabies,
stomachache (Teklehaymanot et al. 2007), as well as rabies, warts, amoebas, and
external injury (Teklehaymanot 2009). Cucumis aculeatus is used as an antimalarial
(Njoroge and Bussmann 2006a) and for general weakness (Njoroge and Bussmann
2006b). Cucumis prophetarum is used to remedy gonorrhea (Wondimu et al. 2007).
Cucumis hartwegii is used for fever and urinary problems (Malik et al. 2015).
Cucumis dipsaceus: Used for adding shine and beauty to hair (Bussmann and
Sharon 2006, 2015). Browsed by livestock (Lulekal et al. 2008). Cucumis ficifolius
is used to treat blackleg (Yineger et al. 2007). The extract is applied to nasal bots
(Wanzala 2017).
References
medicinal flora of Northern Peru. St. Louis: William L. Brown Center, MBG; 2015. ISBN
978-0-9960231-2-2.
10.1186/1746-4269-4-10.
livestock industry with focus on Acari ticks. Evid Based Complement Alternat Med.
2017;2017:Article ID 8647919, 33 pp. https://doi.org/10.1155/2017/8647919.
Cupressus lusitanica Mill.
CUPRESSACEAE

Grace N. Njoroge
Synonyms
Cupressus lusitanica Mill.: Callitropsis lusitanica (Mill.) D.P. Little; Cupressus

benthamii Endl.; Cupressus benthamii var. knightiana (Perry ex Gordon) Mast.;
Cupressus benthamii var. lindleyi (Klotzsch ex Endl.) Mast.; Cupressus glauca
Lam.; Cupressus knightiana Perry ex Gordon; Cupressus lindleyi Klotzsch ex
Endl.; Cupressus lindleyi var. hondurensis (Silba) Silba; Cupressus lusitanica var.
benthamii (Endl.) Carrière; Cupressus lusitanica var. hondurensis Silba; Cupressus
lusitanica var. knightiana Rehder; Cupressus lusitanica var. lindleyi (Klotzsch ex
Endl.) Franco; Hesperocyparis lusitanica (Mill.) Bartel
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Cupressus
sp. (Cupressaceae), Tbilisi,
Georgia. (Photo
R.W. Bussmann and
Local Names
Cupressus lusitanica: Kikuyu: Mutarakwa (Gachati 1989).
Botany and Ecology
Cupressus lusitanica Mill: Evergreen tree, up to 35 m high, with a dense, conical

crown. Branches spread out widely but terminate in pendulous branchlets. Trunk
short, 70 cm in diameter. Bark reddish-brown, exfoliating in long, narrow strips,
eventually becoming roughened by the development of many short cracks. Branchlets
covered with small, decussate, adpressed, acute, sessile, scale leaves with flexed tips.
The distinctly bluish-green foliage is ovate, closely pressed, usually with long, pointed
apex. Male flowers small, oblong or cylindrical; female subglobose, very small,
composed of 6–14 fertile decussate scales with several ovules each. Cones ellipsoid;
bluish-green when young turning reddish-brown when mature; 13–25 mm in diame-
ter; composed of 6–12 woody, shield-like scales, each bearing 8–10 seeds; male cones
appear to be fat tips to branchlets and produce clouds of yellow pollen; female cones
rounded, scales with central pointed projections. Seeds brown, with resin glands, up to
4 mm long, with a narrow wing (Macbride and Weberbauer 1936–1995; Wu et al.
1994–2013). Very common plantation tree around the globe (Figs. 1 and 2).
Cupressus lusitanica: Green parts are used to treat vaginal hemorrhages, general
bleeding, acne, and hair loss (Bussmann and Sharon 2006). Fresh leaves are boiled,
and the extract is drunk to treat diarrhea (Yineger and Yewhalaw 2007).
Cupressus lusitanica Mill. 373
Fig. 2 Cupressus
sp. (Cupressaceae), Tbilisi,
Georgia. (Photo
R.W. Bussmann and
Cupressus lusitanica: The leaf smoke is used as insect repellent (Karunamoorthi

et al. 2009; Tabuti 2008), and it is also used as incense in social settings
(Mohagheghzadeh and Faridi 2006). It is used as live fence and multipurpose timber
tree (Mekonnen et al. 2015).
References
Karunamoorthi K, Ilango K, Endale A. Ethnobotanical survey of knowledge and usage custom of
traditional insect/mosquito repellent plants among the Ethiopian Oromo ethnic group.
Macbride JF, Weberbauer A. Flora of Peru. Chicago: Field Museum; 1936–1995.
Wu Z, Raven PH, Hong D, editors. Flora of China. St. Louis: Science Press, Beijing & Missouri
Botanical Garden Press; 1994–2013.
Cussonia holstii Harms ex Engl.
ARALIACEAE

Grace N. Njoroge
Local Names
Cussonia holstii: Kikuyu: Murogorogo, Muroha; Meru: Morogorogo; Borana:

Abratu; Kamba: Malende, Maende; Kipsigis: Sopwek, Lulukwet; Maa: Ol-Orur;
Samburu: Olbolorio; Somali: Wadad; Taita: Muaya; Tugen: Soya (Beentje 1994;
Botany and Ecology
Cussonia holstii Harms ex Engl.: A tree to 20 m tall with a straight bole sometimes
exceeding 1 m in diameter and 10 m or more tall in well grown specimens, bark
rather fissured, shed in oblong papery scales. Flowering spikes up to 30 together,
usually less than 15, up to 25 cm long, dense or somewhat lax, floral bracts scale-
like, with or without an apiculum, or peg-like, nearly glabrous or densely pubescent.
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Cussonia holstii (Araliaceae). Bale Mountain National Park, Demaro forest, Ethiopia.
Fruits 4–6 mm long, glabrous or puberulous, sometimes galled. Leaves digitately

compound, petiole up to 41.5 cm long, glabrous or virtually so, lamina up to 36 cm
wide by about half as long, leaflets 3–7, chartaceous, ovate, up to 18.5 cm long by
9 cm wide, acuminate to caudate, with a cuneate to cordate, symmetric or asymmet-
ric base, with serrate to crenate or entire margin, glabrous or puberulous, often with
very fine silky-white scattered hairs and sometimes a few short crisped hairs,
petiolules up to 7 cm long, juvenile leaves simple, palmately lobed (Tennant
1968). Understory tree in drier Afromontane forests (Bussmann 2002; Bytebier
and Bussmann 2000) (Figs. 1 and 2).
Cussonia holstii: Bark decoction is used to expel the placenta and clean the uterus
after birth (Beentje 1994; Kokwaro 2009). Leaves are boiled and the extract is
ingested for abdominal pain (Muthee et al. 2011). The bark is pounded and chewed
to treat cough (Njoroge and Bussmann 2006). Crushed leaves are applied to burns
The bark extract also serves to treat filariasis (Desta 1995).
Cussonia holstii Harms ex Engl. 377
Fig. 2 Cussonia holstii (Araliaceae). Bale Mountain National Park, Demaro forest, Ethiopia.
Local Food Uses
Cussonia holstii: The roots are boiled for a strengthening soup (Bussmann 2006).
Cussonia holstii: Soft wood is used for doors and beehives (Beentje 1994;
Bussmann et al. 2011). Leaves are browsed by livestock (Bussmann 2006).
References
Tennant JR. Flora of tropical East Africa: Araliaceae. London: Crown Agents for Overseas
Governments & Administrations; 1968. 22 pp. ISBN: 978-1-84246-231-7.
Cyathea manniana Hook.
CYATHEACEAE

Grace N. Njoroge
Synonyms
Cyathea manniana Hook.: Cyathea engleri Hieron. ex Brause; Cyathea

laurentiorum Christ; Cyathea manniana var. preussii Tardieu; Cyathea preussii
Diels; Cyathea sellae Pirotta; Cyathea usambarensis Hieron. ex Brause
Local Names
Cyathea manniana: Kikuyu: Ruthirathiru, Rusirusiru; Kipsigis: Biriruop cheretet

R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Cyathea manniana Hook: Stem up to 10 cm in diameter, and up to 6 m tall, erect,

slender, occasionally sending down lateral “prop” stems which are eventually
creeping and may form a new upright caudex. Frond arching to horizontal, firmly
chartaceous. Stipe about 30 cm long, light- to very-dark-purplish-brown, sharply
spinose, the spines at the base up to 4 mm long, and with a thin rufous tomentum
when young. Lamina up to 2.4 1 m, 3-pinnate, ovate in outline; pinnae up to
52 20 cm, oblong, acute, pinnate into very narrowly shortly attenuate pinnate
pinnules; pinnule segments very narrowly oblong, somewhat falcate, acute, sub-
entire to crenate-dentate; ventral surface glabrous except for dense stiff curved pale-
brown hairs along the costa and costules; dorsal surface often glaucous and with
often imbricate lanceolate lacerate brown scales up to 3 mm long along the costules
and costae of the pinnae segments; rachis light-brown with scattered small prickles,
glabrous at maturity. Sori up to 9 per pinnule segment, about 0.8 mm in diameter;
indusium unequally cupuliform wet shaded areas in forest 1400–2300 m (Edwards
2005). A characteristic component of the wettest part of lower Afromontane forests
(Bussmann 2002) (Figs. 1, 2, 3, 4, 5, and 6).
Fig. 1 Cyathea manniana

(Cyatheaceae), Mt. Kenya
R.W. Bussmann)
Cyathea manniana Hook. 381
Fig. 2 Cyathea pallescens

(Cyatheaceae), Laguna El
Jardín, Leymebemba, San
Martín, Perú. (Photo
R.W. Bussmann)
Fig. 3 Cyathea pallescens (Cyatheaceae), Laguna El Jardín, Leymebemba, San Martín, Perú.
Cyathea manniana: The stem mucus is used as anthelminthic (Kokwaro 2009).

Cyathea marattioides is used to relief back pain (Rabearivony et al. 2015).
Cyathea manniana Hook. 383
Fig. 6 Cyathea pallescens

(Cyatheaceae), Laguna El
Jardín, Leymebemba, San
Martín, Perú. (Photo
R.W. Bussmann)
References
Edwards PJ. Flora of tropical East Africa: Cyatheaceae. Kew: Royal Botanic Gardens; 2005. 15 pp.
ISBN: 1-84246-114-1
Cyathula cylindrica Moq.
Cyathula polycephala Bak.
AMARANTHACEAE

Grace N. Njoroge
Synonyms
Cyathula cylindrica Moq.: Achyranthes cylindrica Bojer; Pupalia alopecurus Fenzl

ex Drège
Local Names
Cyathula cylindrica: Kikuyu: Githengenya, Maramata, Magwata-Ng’ondu;

Shambaa: Mwilangoto; Kipsigis: Ng’atumyat (Gachati 1989; Kokwaro 2009).
Cyathyla polycephala: Kikuyu: Githengenya, Maramata, Magwata-Ng’ondu;
Kipsigis: Namgueit, Ng’atumyat; Iraqwi: Nuhururi; Luhya: Shinamundu (Gachati
1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Cyathula cylindrica: Moq. Perennial herb, very variable in habit from bushy and
0.6–1 m high to sprawling or decumbent and rooting at the lower nodes, or
subscandent to 6 m or more. Ovary obovoid, 1 mm, rather firm. Style slender,
2–3 mm Capsule ovoid, 2–3 mm, membranous save for the firm apex. Seed ovoid,
1.75–2.75 mm, brown, shining, almost smooth. Stems and branches terete and striate
in the older parts, becoming bluntly tetragonous and finally sharply tetragonous-
sulcate above, glabrous or moderately to densely (especially upwards) furnished
with long spreading or upwardly appressed fuscous bristly multicellular hairs, nodes
distinctly swollen in life; in dried material, the stem and branches commonly
shrunken just above the nodes. Leaves very variable in size and shape, small and
roundish to large and broadly oblong- or elliptic-ovate, 1–14 0.7–6 cm, subcordate
to attenuate at the base, rounded to acuminate at the apex, glabrous to densely
furnished with long appressed multicellular hairs on both surfaces, more rarely
tomentose, petiole distinct, up to 2.5 cm long. Inflorescences terminal on the stem
and branches, spiciform, 1.5–2(–2.5) cm in diameter, in robust plants elongate-
cylindrical and up to 18 cm long, or sometimes scarcely longer than broad, “spikes”
formed of densely congested (more rarely a few of the lower distant) shortly
pedunculate cymose clusters composed mostly of triads of fertile flowers each
subtended by 1–2 modified flowers, bracts elliptic-oblong, 7–8.5 mm, stramineous
or silvery, glabrous or furnished with long multicellular hairs about the tip, aristate
with the excurrent midrib, the arista usually bent but not sharply uncinate, bracteoles
broadly ovate, acuminate, 4.5–9 mm, glabrous or furnished with long multicellular
hairs about the tip, aristate with the excurrent midrib, the arista usually bent but not
sharply uncinate. Tepals narrowly lanceolate-oblong, 4.5–7.5 mm, 3(–5)-nerved,
outer 2 tepals rather feebly nerved, gibbous dorsally at the base, broadly hyaline-
margined, glabrous or almost so, acute to rather blunt, the midrib excurrent in a short
mucro, inner 3 tepals progressively more strongly nerved, more narrowly hyaline-
margined and blunter, the innermost obtuse and often minutely lacerate-dentate at
the apex with the midrib ceasing below the tip, all 3 moderately to densely furnished
with long white multicellular barbellate hairs. Modified flowers with a few narrow
lanceolate bracteoliform processes with uncinate tips, simple hooks, and shorter
membranous scales within. Filaments slender, 3.5–5 mm, the pseudostaminodes
cuneate-obovate, 1/4 the length of the filaments, fringed above, frequently with a
filiform dorsal scale (Townsend 1985). Common in the undergrowth especially of
Croton dominated Afromontane forests (Bussmann 2002; Bytebier and Bussmann
2000) (Figs. 1, 2, and 3).
Cyathula polycephala Bak: Perennial herb, erect and little branched to trailing with
the lower nodes rooting, or semi-scandent or scandent, 0.6–2 m or probably more
when scrambling over tall bushes. Seed ovoid, 1.75 mm, brown, almost smooth.
Stem and branches rather weak, terete and striate in the older parts, becoming
tetragonous and finally more clearly angled and sulcate above, densely pilose with
Cyathula cylindrica Moq. . . . 387
Fig. 1 Cyathula cylindrica

(Amaranthaceae), Mt. Kenya
forest, Kenya. (Photo
R.W. Bussmann)

R.W. Bussmann)
patent or deflexed, whitish or yellowish, multicellular hairs, the older parts more thinly
hairy and occasionally glabrescent, nodes distinctly swollen, the stem and branches
shrunken above the nodes when dry. Leaves broadly ovate to lanceolate-ovate or

R.W. Bussmann)
oblong-lanceolate, 4–10(–16) (2.25–)3.5–7.5 cm, shortly cuneate to subcordate at

the base, acute to shortly or more longly acuminate at the apex, moderately to
densely furnished on the upper surface with appressed multicellular barbellate,
hairs, more densely hairy on the lower surface with the venation sometimes densely
velutinous, petiole of larger leaves 0.5–1.7 cm Style slender, 2–3 mm Capsule ovoid,
2 mm, membranous with a firm flattish top. Inflorescences terminal on the stem and
branches, each a spike-like thyrse formed of opposite or subopposite (occasionally
clusters of more than 2) sessile or shortly (to 7 mm) pedunculate globose condensed
cymes 2–2.5 cm in diameter, the entire thyrse (4–)6–20 cm long with the lowest
cymes increasingly distant, peduncle (3–)5–18 cm, both it and the inflorescence-axis
densely yellowish or whitish pilose, bracts ovate or oblong-ovate, 4.5–5.5 mm,
brownish or silvery membranous, whitish pilose at least along the midrib, which is
Fig. 4 Cyathula polycephala

(Amaranthaceae), Bale
excurrent in a short mucro, bracteoles ovate or deltoid-ovate, 4.5–7 mm, similarly

membranous, thinly to moderately long-pilose at least along the midrib, which is
excurrent into a distinct uncinate arista, ultimate divisions of lateral cymes formed of
a central fertile flower subtended on each side by a triad of 1 fertile and 2 lateral
modified flowers. Filaments delicate, 2–3 mm, pseudostaminodes 0.75–1 mm,
cuneate-oblong, fimbriate. Ovary obovoid, 1 mm. Outer 2 tepals lanceolate-
subnavicular, 5–7 mm, both with an uncinate awn formed by the excurrent midrib,
1-nerved, with long whitish barbellate hairs along the central dorsal surface, inner
3 tepals shorter, 3.5–5 mm, oblong-lanceolate (broader than those of C. uncinulata),
with 3–4 slender nerves along the greenish center, not or obscurely mucronate,
margins hyaline, the apical half densely pilose with whitish barbellate hairs. Mod-
ified flowers of 2 lanceolate uncinate-tipped bracteoliform processes and a few
uncinate spines of variable length (Townsend 1985). Common in the undergrowth
especially of Croton dominated Afromontane forests (Bussmann 2002; Bytebier and
Bussmann 2000) (Figs. 4, 5, 6, and 7).

Cyathula cylindrica: Pounded root bark is boiled in water, and the decoction used
for malaria and as emetic and purgative, as well as for leprosy. The leaf extract is
used to treat hookworms (Kokwaro 2009). Used for malaria, as emetic and purgative
(Jeruto et al. 2008).
Cyathula polycephala: Root extract is used for malaria and fever, as well as emetic
and purgative. The root tubers are eaten for stomach problems (Kokwaro 2009). The
leaves are boiled and the extract drunk to strengthen the bones (Njoroge and
Bussmann 2006) and for urinary system problems (Njoroge et al. 2004).
Cyathula capitata is used as emetic and abortifacient; Cyathula tomentosa is used
as emetic (Bhat et al. 2013) and for snakebites and skin ailments (Malik et al. 2015).
Cyathula schimperiana serves as antimalarial, for diarrhea, and against fungal
infections (Jeruto et al. 2008). Cyathula uncinulata also has medicinal uses

(Yineger et al. 2007, 2008). In Madagascar, the species is used to treat stomach
ulcers, hepatitis, diabetes, and cardiac problems, and is often sold in markets
(Randriamiharisoa et al. 2015).
Cyathula cylindrica: Crushed roots are used against rabies, eaten by cattle
(Bussmann 2002).
Cyathula polycephala: Eaten by livestock (Bussmann 2006; Bussmann et al. 2011)
and used as veterinary medicine (Yineger et al. 2007).
Cyathula uncinulata pods can cause eye damage (Bussmann et al. 2006) but is eaten
by cattle, and the crushed roots are used against rabies (Bussmann et al. 2011; Luizza
et al. 2013).

References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002;91(1/2), 27–79,
appendices 1–7.
Res Appl. 2013;11:315–39.
Randriamiharisoa MN, Kuhlman AR, Jeannoda V, Rabarison H, Rakotoarivelo N, Randrianarivony T,
Raktoarivony F, Randrianasolo A, Bussmann RW. Medicinal plants sold in the markets of
Antananarivo, Madagascar. J Ethnobiol Ethnomed. 2015;11:60. https://doi.org/10.1186/s13002-
015-0046-y.
Townsend CC. Amaranthaceae. In: Polhill RM, editor. Flora of tropical East Africa. Rotterdam/
Boston: A.A.B.alkema; 1985. 136 pp. ISBN: 90-6191-313-6. www.naturekenya.org/
JournalEANH.htm
10.1186/1746-4269-4-11.
Cynodon dactylon (L.) Pers.
POACEAE

Grace N. Njoroge
Synonyms
Cynodon dactylon (L.) Pers.: Agrostis bermudiana Tussac ex Kunth; Agrostis

filiformis J. Koenig ex Kunth; Capriola dactylon (L.) Hitchc.; Capriola dactylon
(L.) Kuntze; Capriola dactylon var. maritima (Kunth) Hitchc.; Chloris cynodon
Trin.; Chloris paytensis Steud.; Cynodon aristiglumis Caro & E.A. Sánchez;
Cynodon aristulatus Caro & E.A. Sánchez; Cynodon dactylon fo. viviparus Beetle;
Cynodon dactylon subsp. glabratus (Steud.) Chev.; Cynodon dactylon var. elegans
Rendle; Cynodon dactylon var. glabratus (Steud.) Chiov.; Cynodon dactylon var.
maritimus (Kunth) Hack.; Cynodon erectus J. Presl.; Cynodon glabratus Steud.;
Cynodon maritimus Kunth; Cynodon occidentalis Willd. ex Steud.; Cynodon
pascuus Nees; Cynodon polevansii Stent; Cynodon portoricensis Willd. ex Steud.;
Cynodon tenuis Trin. ex Spreng.; Cynodon umbellatus (Lam.) Caro; Cynosurus
dactylon (L.) Pers.; Cynosurus uniflorus Walter; Dactilon officinale Vill.; Digitaria
dactylon (L.) Scop.; Digitaria glumaepatula (Steud.) Miq.; Digitaria littoralis
Salisb.; Digitaria littoralis Stent; Digitaria maritima (Kunth) Spreng.; Digitaria
stolonifera Schrad.; Fibichia dactylon (L.) Beck; Fibichia umbellata Koeler; Milium
R. W. Bussmann (*)
G. N. Njoroge

dactylon (L.) Moench; Panicum dactylon L.; Panicum glumaepetalum Steud.;

Paspalum dactylon (L.) Lam.; Paspalum umbellatum Lam.; Phleum dactylon Pall.
ex Georgi
Local Names
Cynodon dactylon: Kikuyu: Igoka; Luganda: Kalandalugo (Gachati 1989;

Kokwaro 2009).
Botany and Ecology
Cynodon dactylon (L.) Pers.: Perennial, stoloniferous, also with slender scaly
rhizomes, sward forming. Culms slender, 10–40 cm tall. Leaf sheaths bearded at
mouth, otherwise glabrous or thinly pilose, leaf blades linear, short and narrow, 1–
12 cm, 1–4 mm wide, usually glabrous, apex subacute, ligule a line of hairs. Racemes
digitate, (2–)3–6, 2–6 cm, straight or gently curved, rather stiff, spreading, spikelets
overlapping by 1/2–2/3 their length. Spikelets 2–2.7 mm, rachilla extension ca. 1 mm,
sometimes with minute rudimentary floret at apex, glumes linear-lanceolate, often
purplish, usually more than half as long as floret, 1.5–2 mm, 1-veined, keel scabrous,
thickened, lemma as long as spikelet, silky villous along keel, hairs straight, other-
wise glabrous or lateral veins thinly villous, apex subacute, palea glabrous, keels
scaberulous. Anthers more than 1 mm. Caryopsis subterete, scarcely laterally com-
pressed. Flowering and fruiting nearly all the year (Wu et al. 1994–2013).
Cynodon dactylon: A leaf decoction is used for liver problems (Kokwaro 2009). In
Latin America, it is used to treat cysts of the ovary, cysts of the uterus, inflammation
of the kidneys, inflammation (general), uterus problems, fibroids, and uterus pro-
lapse (Bussmann and Sharon 2006). The species has low toxicity (Desta 1995). Used
to treat fever and internal injuries (Kumar et al. 2011). Effective to treat sprains and
earache (Kunwar et al. 2009, 2010). In Madagascar, it is applied for intoxication,
gonorrhea, pneumonia, tonsillitis, measles, and tension (Razafindraibe et al. 2013),
also sold in markets to treat kidney stones and syphilis (Randriamiharisoa et al.
2015). The species serves for pregnancy issues, postpartum recovery, and uterus
treatments (Randrianarivony et al. 2016). The leaves are chewed for toothache
(Debbarma et al. 2017). Leaf paste is applied to bleeding wounds, and the extract
is used to treat diarrhea (Muhammad et al. 2019). Also employed to treat cuts and
wounds, bleeding, vomiting, nose bleeding, to control vomiting, skin disease,
leprosy, piles, asthma, arthritis, burning urination, indigestion, cancer, eye and
mouth problem, dysentery, as blood purifier, to control nose bleeding, as antiseptic,
to treat snakebite, and to stop miscarriages (Raj et al. 2018). Sometimes used to treat
Cynodon dactylon (L.) Pers. 397
stomachache, bladder stones, eye inflammations, high blood pressure, and itching
Cynodon dactylon: Eaten by livestock (Bussmann 2006). Also used to increase

lactation in livestock (Raj et al. 2018).
References
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in Far-West Nepal: a
RW. Indigenous uses of ethnomedicinal plants among forest dependent communities of North-
ern Bengal, India. J Ethnobiol Ethnomed. 2018;14(1):8. https://doi.org/10.1186/s13002-018-
0208-9.
org/10.1186/s13002-015-0046.
Madagascar. Indian J Tradit Knowl. 2016;15(1):68–78.
2013;9:73.
10.1186/s13002-019-0285-4.
Cyphostemma maranguense (Gilg.) Desc.
VITACEAE

Grace N. Njoroge
Local Names
Cyphostemma maranguense: Kikuyu: Munyenyenga (Gachati 1989).
Botany and Ecology
Cyphostemma maranguense (Gilg.) Desc.: Robust climber, 1.5–20 m long, tendrils

branched once, leaves shiny, 3 foliate, leaflets ovate with crenate – serrate margins.
Flowers creamy-white, sometimes with reddish tips, in open cymes. Petals 4–5 mm
long. Fruits reddish purple, 6–9 10–15 mm. Common climber in Afromontane forests
(Bussmann 2002a, b, 2006a, b; Bytebier and Bussmann 2000) (Figs. 1, 2, and 3).
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Cyphostemma
cf. maranguense (Vitaceae).
Bale Mountains National
Park, Demaro forest, Ethiopia.
Cyphostemma maranguense is used as an antimalarial (Njoroge et al. 2004;

Njoroge and Bussmann 2006a).
Cyphostemma maranguense, Cyphostemma bambuseti, and Cyphostemma
kilimandscharicum are used to treat bone pain and tuberculosis (Bussmann 2006a, b).
Local Food Uses
Cyphostemma maranguense, Cyphostemma serpens, Cyphostemma bambuseti,

and Cyphostemma kilimandscharicum: Fruit and leaves are eaten (Bussmann
2006a, b; Bussmann et al. 2006).
Cyphostemma maranguense latex is used against anaplasmosis in livestock

(Njoroge and Bussmann 2006b; Njoroge et al. 2004).
Cyphostemma maranguense (Gilg.) Desc. 401
Fig. 2 Cyphostemma
Fig. 3 Cyphostemma
Many Cyphostemma species are regarded as poisonous in Ethiopia (Bussmann

et al. 2011). Cyphostemma cyphopetalum is used as remedy for rabies, snakebites,
and leishmania (Teklehaymanot et al. 2007).
References
2006b;11(1):41–66.
ment of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol.
2006b;108:332–9.
Dactyloctenium aegyticum (L.) Willd.
Dactyloctenium giganteum B.S. Fischer &
Schweick.
POACEAE

Grace N. Njoroge
Synonyms
Dactyloctenium aegyticum (L.) Willd.: Aegilops saccharinum Walter; Cenchrus

aegypticus L. ex P. Beauv.; Chloris mucronata Michx.; Chloris prostrata (Willd.)
Poir.; Cynosurus aegypticus L.; Cynosurus carolinianus Willd. ex Steud.;
Cynosurus distachyos Rottler ex Steud.; Dactyloctenium aegyptiacus Willd.;
Dactyloctenium aegypticum fo. viviparum Beetle; Dactyloctenium aegypticum var.
mucronatum (Michx.) Lanza & Mattei; Dactyloctenium distachyum Trin.;
Dactyloctenium figarii De Not. Dactyloctenium meridionale Ham.; Dactyloctenium
mucronatum (Michx.) Willd.; Dactyloctenium mucronatum var. erectum E. Fourn.;
Dactyloctenium prostratum Willd.; Eleusine aegyptica (L.) Desf.; Eleusine
aegyptica (L.) Pers.; Eleusine aegyptica (L.) Roberty; Eleusine aegyptica (L.)
Roxb.; Eleusine aegyptica Raf.; Eleusine aegyptica (L.) Desf.; Eleusine cruciata
Elliott; Eleusine cruciata Lam.; Eleusine egyptia Raf.; Eleusine mucronata (Michx.)
Hornem.; Eleusine mucronata Michx.; Eleusine mucronata Stokes; Eleusine
pectinata Moench; Rabdochloa mucronata (Michx.) P. Beauv.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Kikuyu: Mukinda (Gachathi 1989).
Botany and Ecology
Dactyloctenium aegyticum (L.) Willd.: Slender to moderately robust spreading

annual; culms up to 70(–100) cm high, usually geniculately ascending and rooting
at the lower nodes, frequently shortly stoloniferous and mat-forming, less often
erect. Leaf-blades flat, 3–25 cm long, 2.5–7.5(–12) mm wide, papillose-hispid
especially along the margins. Inflorescence composed of (1–)3–9 linear to narrowly
oblong spikes 1.2–6.5 cm long, ascending or often radiating horizontally from the
top of the culm. Spikelets 3–4-flowered, broadly ovate, 3.5–4.5 mm long; glumes
subequal, 1.5–2.2 mm long, the lower lanceolate in profile with a thick scabrid keel,
the upper elliptic to narrowly obovate in profile, the smooth keel extended into a
stout divergent scabrid awn 0.5–2 times as long as the glume; lemmas narrowly
ovate to ovate in profile, 2.5–4 mm long, the keel gibbous, concave and scabrid
above the middle and often extended into a stout cusp or mucro up to 1 mm long;
palea-keels winged or unwinged; anthers 0.25–0.8 mm long. Grain about 1 mm long,
broadly obovate to obtriangular in profile, transversely rugose. A widespread weed
of open situations in grassland and open woodland, common by roadsides and on
waste ground; sea-level to 2100 m.
Dactyloctenium giganteum B.S. Fischer & Schweick.: Robust tufted annual; culms
(30–)70–160 cm high, erect or geniculate ascending and occasionally rooting at the
lower nodes. Leaf-blades flat, 11–45 cm long, 5–12 mm wide, almost glabrous to
papillose-hispid especially along the margins. Inflorescence composed of (1–)3–9
linear spikes 3.5–11 cm long; spikes usually ascending to form a brush-like cluster at
the top of the culm, less often widely spreading. Spikelets 2–7-flowered, elliptic, 4–
6.2 mm long; glumes subequal, 1.5–2.2 mm long, the lower lanceolate to narrowly
elliptic in profile with a thick scabrid keel, the upper elliptic in profile, the keel
extended into a divergent scabrid awn (1.5)–1.75–4 times as long as the glume;
lemmas lanceolate to narrowly ovate in profile, 3–4 mm long, acuminate, the keel
scabrid, more or less straight or only slightly concave above the middle, extended
into an awn-point 0.7–2 mm long; palea-keels unwinged; anthers 1.3–2.1 mm long.
Grain 0.7–1.1 mm long, broadly obovate to obtriangular, transversely rugose. A
common grass of roadsides, old cultivations, and other disturbed sites, favoring light
sandy soils; 200–2000 m (Clayton 1970; Clayton et al. 1974).
Both species serve as fodder for goats. Seeds ground and used as fish poison
(Neuwinger 2004).
Dactyloctenium aegyticum (L.) Willd. . . . 405
References
Clayton WD. Flora of tropical East Africa: Gramineae (Part 1). London: Crown Agents for
Overseas Governments & Administrations; 1970.
Clayton WD, Renvoize SA, Phillips SM. Flora of tropical East Africa: Gramineae (Part 2). London:
Crown Agents for Overseas Governments & Administrations; 1974.
Dalbergia lactea Vatke
FABACEAE

Grace N. Njoroge
Local Names
Dalbergia lactea: Kikuyu: Mwaritha; Shambaa: Kowa (Beentje 1994; Gachati

1989; Kokwaro 2009).
Botany and Ecology
Dalbergia lactea Vatke: Small tree or shrub, generally scandent if support available
and sometimes with aid of coiled branchlets or peduncles, 3–9(–25) m tall, ever-
green, bark purplish-brown (at least when dried) and striate on branches, ultimately
brown and rather fibrous. Corolla 6.5–9(–10) mm long, white flushed mauve to
purplish (particularly on standard inside), standard broadly ovate to suborbicular,
cordate and very shortly clawed, wings as long as standard and much exceeding keel.
Stamens (9–)10, partially or wholly divided into 2 subequal phalanges. Fruit oblong,
rather abruptly narrowed or rounded at the ends, with the 0.5–1.5 cm long stipe (8–)
R. W. Bussmann (*)
G. N. Njoroge

10–15 cm long, 3–4.5 cm wide, scarcely thickened over the seed-cavity, chartaceous,
greenish or purplish turning dull pale brown or straw-colored, glabrous, with very
laxly reticulate slightly immersed venation. Leaves 13–30 cm long, stipules oblong,
pointed, 5–6 mm long, fugaceous, lateral leaflets 6–8(–10) on either side of rachis,
oblong or oblong-elliptic, rarely slightly obovate-oblong, 4–9 cm long, 2–5.5 cm
wide, rather abruptly rounded or emarginate at apex, very broadly cuneate, rounded
or slightly cordate at base, glabrous to pubescent, primary lateral nerves 14–20 on
either side of midrib. Panicles on leafy branches, 5–15 cm long, but flowering
branches often growing out with leaves all or mostly suppressed forming compound
inflorescences up to 40–50 cm long, flowers congested on ultimate axes, many, axes
with dark brown persistent felty tomentum, bracts ovate to oblong-elliptic,
1–3.5 mm long, caducous, bracteoles near top of short 0.5–2(–3) mm long pedicel,
oblong-elliptic to obovate, 1.5–2 mm long. Calyx 3.5–5 mm long, pubescent to
tomentose with short dark brown to almost black hairs, rarely glabrous. Branchlets
brownish tomentose, usually soon glabrescent (Gillett et al. 1971a, b).
Dalbergia lactea: Leaves and roots are used for abdominal pain. Roots boiled and
extract applied externally for rheumatism and as bath for skin diseases (Bekalo et al.
2009; Kokwaro 2009). Used as an antimalarial (Njoroge and Bussmann 2006a), and wet
leaves are applied externally for asthma treatment (Njoroge and Bussmann 2006b). The
roots are employed to treat various sexually transmitted diseases (Njoroge and
Bussmann 2009). Also used to treat back pains (Njoroge et al. 2004). In Madagascar,
Dalbergia monticola is used to treat bilharzia (Rakotoarivelo et al. 2015), and Dalbergia
purpurascens is used for cranial system problems (Randrianarivony et al. 2017).
The wood and leaves of Dalbergia sissoo are used as antiseptic, for dysentery, skin
eruptions, gonorrhea, headache, leprosy, and skin diseases (Verma et al. 2007), and as
expectorant (Muhammad et al. 2019). Also used for mastitis (Singh et al. 2019). It is
also used for bladder and kidney stones, as laxative, for piles, bronchial problems,
asthma, cough, rheumatism, sunburns, and as blood purifier (Umair et al. 2019).
Dalbergia melanoxylon is used for East Coast Fever in livestock (Kioko et al. 2015)
and is used for firewood (Tian 2017). Dalbergia sissoo serves to treat mastitis in
livestock (Singh et al. 2019).
References
Dalbergia lactea Vatke 409
nationalities and peoples regional state, Ethiopia. J Ethnobiol Ethnomed. 2009;5:26. https://doi.
org/10.1186/1746-4269-5-26.
Gillett JB, Polhill RM, Verdcourt B. Flora of tropical East Africa: Leguminosae, subfamily
Papilionoideae, Part 1. London: Crown Agents for Overseas Governments & Administrations;
1971a.
Papilionoideae, Part 2. London: Crown Agents for Overseas Governments & Administrations;
1971b. [E. Africa, Legum. (Part 4)].
borne diseases among Maasai people in Northern Tanzania. Vet World. 2015;8:755.
2009;8(2):262–9.
Rakotoarivelo NH, Kuhlman A, Rakotoarivony F, Ramarosandratana AV, Jeannoda V,
Randrianasolo A, Bussmann RW. Medicinal plants used to treat the most frequent diseases in
the Ambalabe rural commune. J Ethnobiol Ethnomed. 2015;11:68.
2017. https://doi.org/10.1186/s13002-017-0147-x.
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;18(14):1.
https://doi.org/10.32859/era.18.14.1-11.
10.1186/s13002-016-0131-x.
10.1186/s13002-019-0285-4.
Datura stramonium L.
SOLANACEAE

Grace N. Njoroge
Synonyms
Datura stramonium L.: Datura bertolonii Parl. ex Guss.; Datura inermis Juss. ex
Jacq.; Datura lurida Salisb.; Datura nigra Hassk; Datura parviflora Salisb.; Datura
pseudostramonium Sieber ex Bernh.; Datura stramonium var. tatula (L.) Torr.;
Datura tatula L.; Stramonium foetidum Scop.; Stramonium spinosum Lam.; Stra-
monium tatula (L.) Moench; Stramonium vulgatum Gaertn.
Local Names
Datura stramonium: Kikuyu: Magurukia, Ndatura; Kipsigis: Barutu,

Chemogong’; Luo: Koth Kiyombi; Taita: Mwalola; Luhya: Silulu (Gachati
1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Datura stramonium L. is native to the Americas and has been introduced in many
tropical, subtropical, and even temperate regions. It is a naturalized weed in many
African countries but is probably seriously underreported. Annual. Plant (12)20–100
(120) cm tall, fetid. Root fusiform, with numerous slender branches, white. Stem
simple or dichotomously branched above, green, glabrous, branches pubescent on
inner side, usually diverging at acute angle. Leaves with petioles equaling lamina or
about half as long, basal leaves up to 20 cm long and broad, ovate, acuminate,
cuneate at base, margin with large unequal acute or acuminate teeth, simple, rarely
incised, upper surface dark green, lighter beneath, both surfaces sparsely puberulent,
slightly more densely underneath (or pubescent only underneath). Rowers solitary in
bifurcations of stem and branches, on straight, erect, pubescent, 7–12 mm long
pedicels. Calyx 4–6 cm long, pale green, 5-angled, tubular, slightly inflated, with
5 acuminate teeth. Corolla 6–10(12) cm long, white, infundibuliform, with long
narrow tube and plicate broad limb, cleft into 5 (very rarely 6) short broadly
triangular lobes, sharply tapering above into slender 5–8 mm long cusp. Ovary
densely covered with soft bristles, style slender, long. Fruit ovoid or subglobose
capsule, surrounded below by recurved persistent, calyx base, spiny, often sparsely
so in lower part, spines distinctly longer and thicker at apex, dehiscing by 4 valves,
though not reaching base. Seeds reniform 3(3.5) nun long, black, with very finely
pitted reticulate surface and large shallow indentations, mainly near bulging external
margin. Ruderal plant, growing near habitations, in kitchen gardens, on garbage and
dunghills, along pasture edges, in mountains. The question of the origin of
D. stramonium has not yet been satisfactorily resolved, although several suggestions
have been offered on this subject. Most authors are inclined to look for the native
habitat of D. stramonium in the Eastern Hemisphere. Closely related species are also
found in the Old as well as the New World. In Eurasia, the following species (with
white flowers) are found to be closely related: D. bertolonii Pari, ex Guss. – Sicily,
D. inermis Jacq. – Ethiopia, D. wallichii Dun. – Nepal, the first two are distinguished
from D. stramonium by glabrous capsules, the last by leaves canescent on the upper
surface. Of the American species, the Mexican ones are closely related:
D. quercifolia H.B.K. and to a lesser extent D. discolor Bemh. Both species have
an anthocyanin-colored corolla as in D. tatula L., the species closest to
D. stramonium, regarded by several authors as a separate species assumed to be a
native of Central or South America (Shishkin and Boborov 1955) (Figs. 1, 2, 3, 4, 5,
6, and 7).
Phytochemistry
Datura stramonium: A highly poisonous and medicinal plant. All parts of Datura
contain alkaloids: hyoscyamine in very large quantities (about 0.28% in leaves on an
average and 0.33–0.48% in seeds) and a small quantity of scopolamine and atropine
(an isomer of hyoscyamine). The seeds are also found to contain fatty oil (Datura
Datura stramonium L. 413
Fig. 1 Datura meteloides

(Solanaceae), Bale Mountains
R.W. Bussmann)
oil) 16–25%, linool 15%, palmitic acid 10%, butyric acid 6.2%, glycerol 9.6%, a
small quantity of alcohols, aldehydes, ketones, etc. The peak quantity of alkaloids
accumulates at the end of the summer and remains constant until the end of the
vegetative period.
The leaves of Datura and, frequently, also the seeds are used in the pharmacopeia of
many countries. Extracts, tinctures, and candles are made from the seeds, while the
leaves are included in the composition of antiasthmatic powders and cigarettes.
Datura preparations are used for treating neuralgia resulting from rheumatism and
asthma as an antispasmodic and narcotic remedy.
Datura stramonium and Datura metel have largely similar medicinal uses through-
out the world. The most widely known use of Datura stramonium and of other
Datura species is for relieving asthma, cough, tuberculosis, and bronchitis by
smoking the dried leaves, roots, or flowers. “Asthma cigarettes” have been shown
to be very effective in some cases, but in other cases, they had little or no effect.
Cigarettes made with the leaves are also used to treat Parkinson’s disease. A
decoction or infusion of leaves is given as a sedative to mental and schizophrenic
patients. The leaves are applied as a dressing to cure rheumatic pain, swellings,
Fig. 2 Datura meteloides

R.W. Bussmann)
wounds, gout, burns, ingrown toenails, fungal infections, tumors, and ulcers. Dried
pulverized leaves are dusted on wounds or applied after mixing the powder with fat
or Vaseline. In East Africa, the fruit is heated, and the juice then applied for earache.
Leaf poultices are used for rheumatism and swellings. In Kenya, the seed oil is used
as massage oil (Kokwaro 2009). Leaves are crushed and applied to abscesses with
pus, deep thorns, infected swollen wounds, etc. This will extract the thorn/pus
(Bussmann et al. 2011). The smoke of both species is inhaled to treat respiratory
tract diseases, as toothache remedy, analgesic, and narcotic (Mohagheghzadeh and
Faridi 2006). The leaf powder is applied to external injuries (Teklehaymanot et al.
2007) and for other applications (Lulekal et al. 2008; Wondimu et al. 2007). Also
applied to tinea versicolor and for toothache (Yineger et al. 2008).
The leaves of Datura and, frequently, also the seeds are used in the pharmacopeia
of many countries. Extracts, tinctures, and candles are made from the seeds, while
the leaves are included in the composition of antiasthmatic powders and cigarettes.
Datura preparations are used for treating neuralgia resulting from rheumatism and
asthma as an antispasmodic and narcotic remedy.
In DR Congo, pounded fresh root and fresh leaves are soaked in water and the
liquid is given in enema as an abortifacient. In Zimbabwe, a hot poultice of leaves
and roots is applied to goiter. A leaf infusion is drunk to treat venereal diseases, and
to cure ulcers, the skin is washed with an infusion of roots and leaves. In Burundi,
leaf ash is eaten as a cure for whooping cough. In Rwanda, a leaf infusion is taken as
Fig. 3 Datura stramonium

R.W. Bussmann)

R.W. Bussmann)
an antispasmodic and to reduce stomach acidity. In Kenya, dried and ground leaves
and seeds are eaten mixed with fat to treat ringworm. Headache is relieved by
rubbing the scalp with leaves or leaf sap. Hair loss is countered by applying fruit
sap or leaf pulp, and these also serve to remedy dandruff. In Ethiopia, pieces of

R.W. Bussmann)
young fruit are sucked against tonsillitis and sore throat and applied to abscesses and
swollen glands. In Kenya and Lesotho, the fruit is heated in hot ash, and after
cooling, juice is squeezed and used as ear drops to treat earache. In Zimbabwe, an
infusion of fruit ash is drunk to treat stomachache. In Ethiopia, the smoke of burning
seeds is inhaled to relieve toothache, while in Kenya, fresh green fruit is applied for
this purpose. In Namibia, a leaf extract is administered to cows to ensure a rapid
expulsion of the afterbirth and pulped roots are mixed with water and given to cattle
to cure lung diseases. The dried leaves and seeds of Datura stramonium are included
in the pharmacopoeias of many Western countries as an antispasmodic and for
treatment of asthma, whooping cough, and Parkinson’s disease. In the Altai, Datura
is used for depression, convulsions, hiccups, stuttering, tetanus, whooping cough,
epilepsy, mental illnesses, and externally for ulcers, burns, tumors, mastitis, con-
junctivitis, and erysipelas. In the Northern Caucasus, Datura is used for pleurisy and
as anthelmintic. In the Ural, the extract serves to treat asthma, rheumatism, arthritis,
pleurisy, stomach ulcers, burns, toothache, and eczema (Sokolov 1990). The leaves
are crushed and applied to abscesses with pus, deep thorns, and infected swollen
wounds. This will extract the thorn/pus (Bussmann et al. 2011). The leaves can be
applied to boils and sores, and the flowers are used to remedy dandruff and hair loss
(Joshi et al. 2010; Kunwar et al. 2009, 2010, 2015). In Kenya, the extract is used for
tonsillitis (Njoroge et al. 2004; Njoroge and Bussmann 2006) and skin swellings
(Njoroge et al. 2004; Njoroge and Bussmann 2007). The leaves are used for

(Solanaceae), Cusco, Peru.

R.W. Bussmann)
hemorrhoids and to stop weight loss in children (Giday et al. 2007). It is also applied
for toothache (Giday et al. 2009; Mekonnen et al. 2015).
The narcotic use of Datura stramonium varies between cultures. In Central and
South America, hallucinogenic uses are common among native tribes. In Africa,
before they enter fighting contests, young men of the Fulfulde people of the border
area of Niger and Nigeria are served drinks containing Datura seeds. This increases
their courage and pain tolerance. The leaves are most commonly used as a narcotic,
either smoked or boiled and eaten, seeds are similarly used. Roots, seeds, or leaves
are added to alcoholic drinks to increase the intoxicating effect. Side effects include
dry mouth and throat, eye pain, blurred vision, restlessness, dizziness, arrythmia,
flushing, and faintness. An overdose will cause headache, nausea, vomiting and
affect the central nervous system causing disorientation, hallucinations, euphoria,
inappropriate affect, short-term memory loss, and coma. The seeds are also used for
criminal purposes. Hospital admissions and fatalities, most often of adolescents, are
not uncommon. It is for this reason that several countries including France removed
Datura cigarettes from the pharmacopoeia in 1992 (Bussmann 2016).
Datura stramonium: Reports on the use of the plant as an insecticide vary from
good control of aphids in crops in Namibia to no effects in Australia. In East Africa,
the leaves yield a green dye that is used to dye cloth; in Lesotho, the twigs yield a
blue-green dye that is used for house decoration. In Ethiopia, the plant has been used
to tattoo the gums, partly as a treatment of gingivitis or dental decay. The stems are
used as firewood. Leaf infusions are used in coccidiosis in chicken and for bloat in
cattle (Kokwaro 2009). The plant is used as insect repellent in livestock
(Karunamoorthi et al. 2009).
In veterinary medicine, the extract is used topically as antispastic and for wound
healing. The extract is used as insecticide for aphids, spider mites, bedbugs, and
caterpillars. It is highly toxic (Sokolov 1990).
References
Bussmann RW. Magic plants. In: Albuquerque U, Alves R, editors. Introduction to ethnobiology.
Heidelberg: Springer International Publishing; 2016. p. 163–9.
in Kumaun Himalaya. Med Aromat Plant Sci Biotechnol. 2010;4(Special issue 1):43–6.
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in Far-West Nepal: a
medicines in Far-West Nepal. J Ethnopharnacol. 2015;163:210–9.
10.1186/1746-4269-4-10.
Shishkin BK, Boborov EG. Flora of the USSR, Volume 22: Solanaceae and Scrophulariaceae.
Leningrad: Akademia Nauk; 1955 (English 1993). 745 pp.
Sokolov PD, editor. Plant resources of the USSR: flowering plants, their chemical composition, use.
Volume 5. Families of Caprifoliaceae – Plantaginaceae. Leningrad: Akademia Nauk; 1990.
ailments at Bale Mountains National Park, Southeastern Ethiopia. J Med Plants Res. 2008;2
(6):132–53.
Dioscorea dodecaneura Vell.
Dioscorea dumetorum (Kunth) Pax
Dioscorea minutiflora Engl.
DIOSCOREACEAE

Grace N. Njoroge
Synonyms
Dioscorea dodecaneura Vell.: Dioscorea bangii R. Knuth; Dioscorea bulbifera L.;

Dioscorea colocasiifolia Pax; Dioscorea discolor R. Knuth; Dioscorea dodecandra
Steud.; Dioscorea dodecaneura var. maronensis Uline ex R. Knuth; Dioscorea hebantha
Mart. ex Griseb.; Dioscorea huallagensis R. Knuth; Dioscorea illustrata W. Bull.;
Dioscorea pilosiuscula Bertero ex Spreng.; Dioscorea racemosa Rusby; Dioscorea
septemnervis Vell.; Dioscorea sororia Kunth; Dioscorea vittata W. Bull ex Baker
Dioscorea dumetorum (Kunth) Pax: Dioscorea buchholziana Engl.; Dioscorea
triphylla L.; Dioscorea triphylla var. dumetorum R. Knuth; Helmia dumetorum
Kunth
Dioscorea minutiflora Engl.: Dioscorea acarophyta De. Wild.; Dioscorea armata
De Wild.; Dioscorea brevispicata De Wild.; Dioscorea cayenensis Jum.; Dioscorea
demeusei De Wild. & T. Durand.; Dioscorea ealaensis De Wild.; Dioscorea ekolo
De Wild.; Dioscorea engbo De Wild.; Dioscorea grandebulbosa R. Knuth;
Dioscorea grandibulbosa R. Knuth; Dioscorea hystrix R. Knuth; Dioscorea lilea
R. W. Bussmann (*)
G. N. Njoroge

De Wild.; Dioscorea litoie De Wild.; Dioscorea multiflora Mart.; Dioscorea

praehensilis var. minutiflora Baker P.P.; Dioscorea pynaertioides De Wild.
Local Names
Dioscorea dodecaneura: Luhya: Litugu (Kokwaro 2009).

Dioscorea dumentorum: Giriama: Mariga; Sukuma: Maringa, Ndiga (Kokwaro
2009).
Dioscorea minutiflora: Kikuyu: Gikwa (Gachati 1989; Kokwaro 2009).
Botany and Ecology
Dioscorea dodecaneura Vell.: Inflorescence, especially the calyx tube, scurfy

tomentose, gray. Stems slender, strongly sulcate, petioles 2–5 cm long, ribbed,
leaves 7–15 cm long and broad, ovate-cordate, with broad, shallow sinus Abruptly
acute, thin, 9 ribbed, with ribs prominent underneath. Racemes short-peduncled,
10–20 cm long, slender, costate, loosely flowered. Pedicels short, ovary 5–7 mm
long, 2–3 mm wide, oblong, obtusely triangular, downy. Styles nearly equaling
perianth, shortly recurved, thick, bifid.
Dioscorea dumetorum (Kunth.) Pax: Tuber replaced annually, much divided with
short or cylindric round-ended root-bearing lobes up to 2.5 cm in diameter, spread-
ing or descending to 30 cm. Twining stems up to 10 m long, pubescent and beset
sparsely or rather densely with prickles. Leaves alternate, 3-foliolate, petiole up to
20 cm long, pubescent, usually with a few scattered prickles, leaflets with petiole up
to 10 mm long, adpressed pubescent, glabrescent above, discolorous and lanate,
rarely sparsely hirsute beneath, median leaflet obovoid, acutely acuminate, cuneate
or rounded at the base, conspicuously 3-nerved from just above the base. Male
inflorescence paniculate, the ultimate branches spreading in all directions forming
dense subsessile cylindric spikelets up to 15 mm long, subsessile or on peduncles up
to 5 mm long, bracts broadly ovate, adpressed to the perianth and partly concealing
it, densely pubescent, perianth subglobose, glabrous, with the 3 inner segments
1 mm long and the 3 outer considerably smaller and thinner. Stamens 6. Female
inflorescence pendulous, spicate, 5–10 cm long, with the flowers close together at
first, the internodes elongating greatly in age, flowers directed downwards, perianth
depressed subglobose, 2 mm in diameter, pubescent, lobes subequal. Ovary 7 mm
long, densely pubescent. Male inflorescence paniculate, the ultimate branches
spreading in all directions forming dense subsessile cylindric spikelets up to
15 mm long, subsessile or on peduncles up to 5 mm long, bracts broadly ovate,
adpressed to the perianth and partly concealing it, densely pubescent, perianth
subglobose, glabrous, with the 3 inner segments 1 mm long and the 3 outer consid-
erably smaller and thinner. Stamens 6. Female inflorescence pendulous, spicate,
5–10 cm long, with the flowers close together at first, the internodes elongating
greatly in age, flowers directed downwards, perianth depressed subglobose, 2 mm in
Dioscorea dodecaneura Vell. . . . 423
diameter, pubescent, lobes subequal. Ovary 7 mm long, densely pubescent. Capsule

(3–)4(–5) cm long, rather sparsely pubescent. Seeds 2 cm long, with wing on the
basal side only. Edges of lowland rainforest, dry evergreen forest, evergreen bush-
lands and on termite hills in Brachystegia woodland, persisting in plantations and in
secondary thickets and grasslands, 0–1650 m (Milne-Redhead 1975).
Dioscorea minutiflora Engl.: Rootstock perennial, in age consisting of several
horizontally radiating woody arms with a shoot at the outside end, eventually
dying back towards the center to form separate plants, each arm protecting several
vertical fleshy tubers about 25 cm long. Twining stems prickly, up to 10 m high,
glabrous. Leaves opposite or occasionally alternate, glabrous, petiole up to 10 cm
long, blade broadly ovate, orbicular or the uppermost reniform, cordate or rounded at
the base, shortly acuminate, the lower up to 12 cm long and 9 cm wide, the upper
5 cm long and 6 cm wide. Aerial tubers absent. Inflorescence glabrous. Rainforest,
1050–1750 m, in Senegal and Guinée east to Zaire and south to Gabon and Angola
(Milne-Redhead 1975) (Figs. 1, 2, 3, 4, 5, 6, and 7).
Dioscorea dodecaneura: Boiled tubers are given to children for measles (Kokwaro
2009).
Fig. 1 Dioscorea
sp. (Dioscoreaceae),
Marañon, Peru. (Photo
R.W. Bussmann and
Fig. 2 Dioscorea
R.W. Bussmann and
Dioscorea dumentorum: Roots are dried and ground, and the powder is mixed with
water to treat bilharzia. Tubers must be soaked at least overnight to remove toxins
(Kokwaro 2009). Dioscorea quartiniana is used medicinally in Ethiopia (Lulekal
et al. 2007, 2008).
Dioscorea belophylla species are used medicinally at a global scale, and new
species have been described recently (Romero-Hernández et al. 2019). In India,
Dioscorea bulbifera is used as an expectorant for asthma and bronchitis, to treat
diarrhea, urinary discharge, leukoderma, and as febrifuge (Joshi et al. 2010), as well
as to treat jaundice (Raj et al. 2018) and fever and boils (Singh et al. 2017).
Dioscorea deltoidea is used as anthelminthic (Kunwar et al. 2009, 2010, 2013),
for urogenital disorders, and cough (Malik et al. 2015). Dioscorea belophylla is used
for pain relief and against pneumonia (Raj et al. 2018).
Dioscorea trifida: Fresh tuber is used to treat inflammations, renal disease, uterus
disease and discharge, cysts, internal inflammation, cancer of the uterus, inflamma-
tion of the ovaries, vaginal discharge, and inflammation of the kidneys (Bussmann
and Sharon 2006, 2015a, b; Paniagua Zambrana et al. 2020). The species has
antibacterial properties (Bussmann et al. 2010). Dioscorea belophylla is a remedy
for body and stomach pain and pneumonia (Raj et al. 2018).
Fig. 3 Dioscorea
R.W. Bussmann and
Fig. 4 Dioscorea coriacea

(Dioscoreaceae), Marañon,
Fig. 5 Dioscorea
syringiflolia (Dioscoreaceae),
R.W. Bussmann and
Fig. 6 Dioscorea
R.W. Bussmann and
Local Food Uses
Dioscorea praehensilis and other species are eaten as famine food (Balemie and
Kebebew 2006; Giday et al. 2009). Dioscorea soso and Dioscorea ovinala are
frequently eaten in Madagascar (Randrianarivony et al. 2016a, b). The same holds
true for Dioscorea fandra and Dioscorea sansibarensis, after careful detoxification
(Randrianarivony et al. 2016a, b). Dioscorea belophylla is eaten as appetizer (Raj
et al. 2018).
Fig. 7 Dioscorea
R.W. Bussmann and
A variety of Dioscorea species are eaten by cattle (Bussmann et al. 2011). Dioscorea
sansibarensis is used as poison in Madagascar (Randrianarivony et al. 2016a, b),
and, like Dioscorea bulbifera, sometimes used as fish poison (Neuwinger 2004).
References
medicinal flora of Northern Peru. St. Louis: William L. Brown Center, MBG; 2015a. ISBN:
978-0-9960231-2-2.
medicinal del Norte de Peru. St. Louis: William L. Brown Center, MBG; 2015b. ISBN: 978-0-
9960231-3-9.
Bussmann RW, Malca G, Glenn A, Sharon D, Chait G, Díaz D, Pourmand K, Jonat B, Somogy S,
Guardado G, Aguirre C, Meyer K, Rothrock A, Townesmith A, Effio-Carbajal J, Frías-
Fernandez F, Benito M. Minimum inhibitory concentration of medicinal plants used in Northern

Peru as antibacterial remedies. J Ethnopharmacol. 2010;132:101–8.
region, Ethiopia. J Ethnobiol Ethnomedi. 2011;7:28.
doi.org/10.1186/1746-4269-5-34.
in Kumaun Himalaya. Med Aromat Plant Sci Biotechnol. 2010;4(Special issue 1):43–6.
Kunwar RM, Burlakoti C, Chowdhary CL, Bussmann RW. Medicinal plants in Far-West Nepal:
their Indigenous uses and pharmacological validity. Med Aromat Plant Sci Biotechnol. 2010;4
(Special issue 1):28–42.
and management in Far-West Nepal. J Ethnobiol Ethnomed. 2013;9:24.
10.1186/1746-4269-4-10.
Milne-Redhead E. Flora of tropical East Africa: Dioscoreaceae. London: Crown Agents for
Overseas Governments & Administrations; 1975. 25 pp. ISBN: 0-85592-031-9.
Paniagua Zambrana NY, Bussmann RW, Romero C. Dioscorea tambillensis R. Knuth, Dioscorea
trifida L. f. In: Paniagua Zambrana NY, Bussmann RW, editors. Ethnobotany of mountain
regions – ethnobotany of the Andes. Cham: Springer International Publishing; 2020. https://doi.
org/10.1007/978-3-319-77093-2_101-1.
RW. Indigenous uses of ethnomedicinal plants among forest dependent communities of North-
ern Bengal, India. J Ethnobiol Ethnomed. 2018;14(1):8. https://doi.org/10.1186/s13002-018-
0208-9.
services from Agnalavelo sacred forest and their relationship with forest conservation. Mada-
gascar Conserv Dev. 2016a;11(2):44–51.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016b. ISBN: 978-0-9960231-4-6.
Romero-Hernández C, Téllez-Valdés O, Bussmann RW. Dioscorea chusqueifolia (Dioscoreaceae),
a new species from northern Peru Brittonia. 2019. https://doi.org/10.1007/s12228-019-09587-8.
itants of Jakholi Block, Rudraprayag district, Western Himalaya, India. J Etnobiol Ethnomed.
2017;13(49). https://doi.org/10.1186/s13002-017-0178-3.
Diospyros abyssinica (Hiern.) F. White
Diospyros mespiliformis Hochst. ex A. DC.
EBENACEAE

Grace N. Njoroge
Synonyms
Diospyros abyssinica (Hiern.) F. White: Disopyros ferrea (Willd.) Bakh.;

Disopyros piscatoria Gürke; Disopyros ubanghensis A. Chev.; Disopyros
welwitschii Hiern; Ebenus abyssinica (Hiern) Kuntze; Maba abyssinica Hiern;
Maba ubanghensis A. Chev.; Maba warneckei Gürke
Diospyros mespiliformis Hochst. ex A. DC.: Diospuros holtzii Gürke; Diospyros
sabiensis Hiern
Local Names
Diospyros abyssinica: Kikuyu: Muiruthi; Swahili: Mdaa-Mwitu; Borana: Roho;

Gabbra: Lookko; Giriama: Mlala; Ilelewa: Motya-mowgi; Kamba: Mukololo;
Kipsigis: Cheptuiyet; Luhya: Lusui; Luo: Ochol; Maa: Ol-Chartoiyan; Meru:
Mutimuiro; Mutharagwe; Nandi: Cheptuiyet, Kenduiwet; Samburu: Ilchanai
R. W. Bussmann (*)
G. N. Njoroge

orok; Sebei: Cheptua; Taveta: Mesesevu (Beentje 1994; Gachati 1989; Kokwaro
2009).
Diospyros mespiliformis: Ilelewa: Mukowolo; Meru: Mukoro; Oromo: Kolati-
gurati; Somali: Kolati; Taveta: Muyongolo; Turkana: Egum; Acholi: Chumu;
Makua: Mdaka; Kamba: Mukoo; English: Ebony (Beentje 1994; Kokwaro 2009).
Botany and Ecology
Diospyros abyssinica (Hiern.) F. White: Small, medium-sized or large tree up to

36 m tall, but sometimes flowering as a shrub 2 m tall. Bole long, straight, slender.
Bark dark gray or blackish, rough, reticulate and exfoliating on old trees. Leaves
subcoriaceous, drying gray-green or blackish, lamina 3 1.2–12 4 cm, elliptic,
oblanceolate-elliptic or lanceolate-elliptic, apex obtuse to shortly and bluntly sub-
acuminate, rarely distinctly acuminate, lower surface glabrescent, lateral nerves in
5–12 pairs, venation prominent and closely reticulate on both surfaces. Male flowers
axillary and ramuligerous, in contracted cymes, peduncle 0.1 cm long, pedicels
0.1 cm long, fulvous-setulose. Female flowers axillary or ramuligerous, in (1–2)
3–5(8)-flowered fascicles, pedicels 0–2 cm long, fulvous-setulose. Calyx 0.2 cm
long, shallowly cyathiform, with 3–4 short, broadly deltate lobes, glabrous outside
except for a few minute marginal hairs, glabrous inside. Calyx 0.6 cm long,
cyathiform, glabrous outside, finely strigulose towards the base inside, divided
almost to the base, lobes 3–4 up to 0.6 0.6 cm, suborbicular, sometimes apiculate,
strongly imbricate. Corolla 0.5–0.6 cm long, sub-rotate, glabrous, tube 0.15 cm long,
lobes 3–4, 0.45 0.3 cm, broadly elliptic, apex obtuse. Corolla slightly shorter than
the calyx, otherwise as in male. Stamens 10–15, 0.2–0.4 cm long, anthers lanceolate-
apiculate, sparsely setulose towards the apex. Pistillode 0.1 cm long or absent,
glabrous. Pistillode 0.1 cm long or absent, glabrous. Corolla 0.5–0.6 cm long,
sub-rotate, glabrous, tube 0.15 cm long, lobes 3–4, 0.45 0.3 cm, broadly elliptic,
apex obtuse. Stamens 10–15, 0.2–0.4 cm long, anthers lanceolate-apiculate, sparsely
setulose towards the apex. Male floiaers axillary and ramuligerous, in contracted
cymes, peduncle 0.1 cm long, pedicels 0.1 cm long, fulvous-setulose. Calyx 0.2 cm
long, shallowly cyathiform, with 3–4 short, broadly deltate lobes, glabrous outside
except for a few minute marginal hairs, glabrous inside. Staminodes 3–4 glabrous,
0.2 cm long, filiform, exserted, attached to the throat of the corolla and alternating
with the lobes. Ovary 0.4 0.2 cm, conoidal, glabrous, gradually merging into the
short undivided style, locules 6, uniovulate, stigmatic lobes, 3, about 0.1 cm long,
ascending. Ovary 0.4 0.2 cm, conoidal, glabrous, gradually merging into the short
undivided style, locules 6, uniovulate, stigmatic lobes, 3, about 0.1 cm long,
ascending. Corolla slightly shorter than the calyx, otherwise as in male. Staminodes
3–4 glabrous, 0–2 cm long, filiform, exserted, attached to the throat of the corolla
and alternating with the lobes. Female flowers axillary or ramuligerous, in (1–2) 3–5
(8)-flowered fascicles, pedicels 0.2 cm long, fulvous-setulose. Calyx 0.6 cm long,
cyathiform, glabrous outside, finely strigulose towards the base inside, divided
almost to the base, lobes 3–4 up to 0.6 0.6 cm, suborbicular, sometimes apiculate,
Diospyros abyssinica (Hiern.) F. White. . . 431
strongly imbricate. Seed(s) 1 (very rarely 2), 0.9 0.6 cm, globose to sub-ellipsoid,
black, endosperm smooth. Fruiting calyx scarcely accrescent, 0.7 cm long, becom-
ing patelliform. Fruit up to 1.4 0.9 cm, glabrous, ellipsoid or sub-globose, style
persistent (White and Verdcourt 1996). Found in the undergrowth of humid
Afromontane forests (Bussmann 2002). Slow germination under light conditions
(Bussmann and Lange 2000).
Diospyros mespiliformis Hochst. ex A. DC.: Evergreen tree 5–27(–35) (excep-
tionally 45) m tall, with spreading densely rounded crown, but occasionally
flowering as a shrub only 1–3 m tall, has been described as mango-like with
leaves near the ground. Bole 1.5 m and crown 7.5 m wide, stem bark blackish or
dark brown, very rough, longitudinally channeled, flaking in irregular scales,
slash pinkish brown or black outside, pink inside, branchlets knobby, tomentose
with pinkish adpressed hairs when young, glabrescent. Leaves drying pale red-
dish brown when young, dull gray-green above and yellowish green beneath
when old, mostly narrowly elliptic to narrowly oblong-elliptic or oblong to
oblanceolate-elliptic, 3.5–19 cm long, 1.5–7.5 cm wide, mostly acute to
sub-acuminate, less often obtuse or rounded at the apex, cuneate to rounded at
the base, sub-coriaceous, minutely strigulose puberulous with adpressed and
spreading flexuose hairs beneath, lateral nerves 15–20 pairs, ascending at 45°,
indistinct, together with the tertiary nerves and venation forming a reticu-
lum prominent on both sides, petiole 7–10 mm long, the margins revolute.
Male flowers subsessile in 3’s on peduncles 4–6 mm long arising from the axils of
deciduous reduced leaves at the base of the current year’s growth or of the first-
formed normal leaves, flower-buds adpressed silvery-silky velvety, calyx 3 mm
long, calyx-lobes 4–5, triangular, 1.5 mm long, corolla white or greenish yellow,
sweetly scented, narrowly urceolate, 6 mm long, sericeous-tomentose outside,
corolla-tube 5 mm long, glabrous inside and thickened at the throat, corolla-lobes
4–5, triangular, 1 mm long, stamens 14, 4 mm long, included, filaments 1 mm
long, glabrous, inserted on the receptacle, anthers narrowly lanceolate, apiculate,
glabrous except for a few hairs on the connective, rudimentary ovary minute,
tomentose. Female flowers subsessile, solitary or rarely 2–3 together, in the axils
of reduced leaves at the base of the current year’s shoot, calyx and corolla similar
to the male but calyx 8 mm long and lobes triangular-cordate with undulate-
plicate reflexed margins, corolla 1–1.2 cm long, staminodes 6–12, filiform, 4 mm
long, glabrous, inserted at the base of the corolla, ovary 3 mm in diameter with
style scarcely differentiated, locules 4 or 6, stigma sessile. Male flowers sub-
sessile in 3’s on peduncles 4–6 mm long arising from the axils of deciduous
reduced leaves at the base of the current year’s growth or of the first-formed
normal leaves, flower-buds adpressed silvery-silky velvety, calyx 3 mm long,
calyx-lobes 4–5, triangular, 1.5 mm long, corolla white or greenish yellow,
sweetly scented, narrowly urceolate, 6 mm long, sericeous-tomentose outside,
corolla-tube 5 mm long, glabrous inside and thickened at the throat, corolla-lobes
4–5, triangular, 1 mm long, stamens 14, 4 mm long, included, filaments 1 mm
long, glabrous, inserted on the receptacle, anthers narrowly lanceolate, apiculate,
glabrous except for a few hairs on the connective, rudimentary ovary minute,
tomentose. Female flowers subsessile, solitary or rarely 2–3 together, in the axils
of reduced leaves at the base of the current year’s shoot, calyx and corolla similar
to the male but calyx 8 mm long and lobes triangular-cordate with undulate-
plicate reflexed margins, corolla 1–1.2 cm long, staminodes 6–12, filiform, 4 mm
long, glabrous, inserted at the base of the corolla, ovary 3 mm in diameter with
style scarcely differentiated, locules 4 or 6, stigma sessile. Fruit yellow, globose,
1.5–2.5 cm diameter, shiny, verruculose, at first adpressed ferruginous hairy, at
length glabrous save for a few hairs near the base of the persistent style, soft,
edible with dry sweet flesh, calyx patelliform or slightly cyathiform, the lobes
with recurved strongly undulate margins. Seeds 3–6, reddish brown or dark,
compressed oblong-ellipsoid, 10 6.5 4.5 mm, rugulose when dry, the
endosperm deeply ruminate. Moist semideciduous forest, riverine forest,
scattered tree grassland and interfaces with fringing forest or woodland also
dryer Combretum woodland, woodland in rocky places, semi-evergreen thicket
on lava and dry scrub, 0–1500 m (White and Verdcourt 1996). Found in the
undergrowth of humid Afromontane forests (Bussmann 2002).
Diospyros abyssinica: Various parts of the plants are used in traditional medi-
cine. In Ghana, bark and roots are used by the Krobo people against various
diseases. In Mali, leaf and roots decoctions are used to treat malaria and dysen-
tery, and to promote wound healing. In Tanzania, a root decoction is taken to treat
leprosy.
Diospyros mespiliformis: Roasted and pulverized roots are taken to treat jaun-
dice, and root decoctions as anthelmintic, to ease childbirth, and to treat malaria,
pneumonia, and syphilis. Bark preparations are administered to treat cough,
bronchial diseases, tuberculosis, syphilis, and leprosy, and applied externally to
wounds, ulcers, bruises, and furuncles. The bark is also used in veterinary
medicine as vermifuge. Leaf decoctions or infusions are taken to treat fever,
diarrhea, dysentery, trypanosomiasis, menorrhagia, whooping cough, hiccough,
and poisoning. Leaf preparations are externally applied to treat fever, pneumonia,
conjunctivitis, and otitis, and as hemostatic and antiseptic to wounds, yaws, and
furuncles. Fruit decoctions or infusions are taken to treat dysentery, diarrhea, and
menorrhagia. Fruit ash is applied to fungal skin infections and fruit powder to
ulcers, whereas seed decoctions are administered against headache. Twigs are
chewed to clean the teeth. Bark decoction is used for stomachache (Kokwaro
2009).
Diospyros has its distribution center in Madagascar, where a wide variety of species
are sold as medicines in local markets, specially to treat cysticercosis, intestinal
parasites, toxoplasmosis, emphysema, diabetes, albumin regulation, and allergies
(Randriamiharisoa et al. 2015). Some species are used to treat malaria (Razafindraibe
et al. 2013). Diospyros lotus and Diospyros kaki are used to treat dysmenorrhea and
diarrhea (Sher et al. 2016).
Diospyros abyssinica (Hiern.) F. White. . . 433
Local Food Uses
Diospyros mespiliformis: The fruit is sweet but acidulous with a slight lemon-like
taste. It is often eaten raw when fully ripe, particularly by children, but sometimes
also dried and kept for later use when food is scarce at the end of the dry season. The
fruits are also used in the production of fruit juice and alcoholic drinks. They can be
ground into flour, which is sometimes an ingredient of porridge. The seeds are also
eaten, and they have a nut-like flavor. The leaves are occasionally eaten as vegetable.
Diospyros lotus, Diospyros kaki, and other species are planted as fruit trees
(Bussmann 2017; Mehdiyeva et al. 2017).
Diospyros abyssinica: The wood is hard, tough, and moderately heavy. The rates of
shrinkage are moderately high. The wood has a low durability and is susceptible to
attacks by fungi and pinhole borers and moderately susceptible to termite attack. The
wood is used for heavy flooring, poles, interior trim, mine props, furniture, cabinet
making, masts of dhows, agricultural implements, musical instruments, tool handles,
ladders, toys, novelties, pestles, mortars, golf club heads, sticks, carving, and
turnery. It is also in demand for loom shuttles in weaving sisal cloth in Kenya
(Beentje 1994). The wood is commonly used as firewood and for charcoal produc-
tion (Kiefer and Bussmann 2004, 2008).
Diospyros mespiliformis: Various parts of the tree are used in ritual ceremonies. The
wood is used for posts in house construction, flooring, joinery, furniture, ship
building, vehicle bodies, musical instruments such as drums, household utensils
such as cups, spoons, pestles and mortars, tool handles, walking sticks, combs,
agricultural implements such as plows, boxes, carvings and turnery, and for canoes.
The bole is used traditionally for dug-out canoes. The wood is also used as firewood
and is valued for charcoal production (Beentje 1994). The foliage is browsed by
livestock. The gum from the bark is used to mend broken pottery, and the fruit pulp is
used to glaze and varnish pottery. The species are planted for reforestation, as
ornamental shade tree, and as windbreak. The flowers serve as source of nectar for
honeybees. Powdered roots are used against witchcraft (Kokwaro 2009).
Lots of species, including Diospyros sakalavarum and Diospyros capulifera, and
a wide variety of other species are used for construction in Madagascar
(Rakotoarivelo et al. 2013; Randrianarivony et al. 2016). Diospyros species are
widely cut illegally for the lucrative rosewood trade.
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002;91(1/2), 27–79,
Bussmann RW, editor. Ethnobotany of the Caucasus, vol. XXVII. Cham: Springer International
Publishing International Publishing; 2017. 746 p. ISBN: 978-3-319-49411-1.
Nat Hist. 2000;89(1–2):101–11.
2004;6(2):161–9.
Mehdiyeva N, Alizade V, Batsatsashvili K, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S,
Tchelidze D, Paniagua Zambrana NY, Bussmann RW. Diospyros lotus L. In: Bussmann RW,
editor. Ethnobotany of the Caucasus. Cham: Springer International Publishing; 2017.
org/10.1186/s13002-015-0046.
2013;9:73.
White F, Verdcourt B. Flora of tropical East Africa: Ebenaceae. Rotterdam: Balkema; 1996. 51 pp.
ISBN: 90-6191-371-3.
Dodonaea viscosa Jacq.
SAPINDACEAE

Grace N. Njoroge
Synonyms
Dodonaea viscosa Jacq.: Dodonaea angustfolia L. f.; Dodonaea bialata Kunth;

Dodonaea burmanniana DC.; Dodonaea dioica Roxb. ex DC.; Dodonaea
bombeyana Blume; Dodonaea eriocarpa fo. galapagensis Sherff; Dodonaea var.
vaccinioides Sherff; Dodonaea spatulata Sm.; Dodonaea viscosa subsp.
angustifolia (L. f.) J.G. West; Dodonaea viscosa subsp. burmanniana (DC.)
J.G. West; Dodonaea viscosa var. amgustifolia (L. f.) Benth.; Dodonaea viscosa
var. galapagensis (Sherff) Porter; Dodonaea viscosa var. spatulata (Sherff) Porter;
Dodonaea viscosa var. vulgaris Benth.; Ptelea viscosa L.
Local Names
Dodonaea viscosa: Kikuyu: Murema-Muthua; Swahili: Mkaa-Pwani, Mkengata;

Borana: Hidesa; Kamba: Kithongoi; Luhya: Muendu; Luo: Oking’; Maa:
Orgeturai, Ol-Tuyesi; Marakwet: Tabilikwa; Sebei: Tombolokwa; Tugen:
R. W. Bussmann (*)
G. N. Njoroge

Tupulukwo; Hehe: Luhahi; Pare: Mwanjio, Njitwe (Beentje 1994; Gachati 1989;
Kokwaro 2009); English: Hopbush, Switch sorrel, Sand olive.
Botany and Ecology
Dodonaea viscosa Jacq.: Monoecious or dioecious semiprostrate shrub or small

tree, 0.5–9 m tall, bark black or brown, rough, twigs black or reddish brown,
glandular, glabrescent, ridged and developing vertical fissures. Leaves simple,
variable, blades oblanceolate or broadly to narrowly elliptic, (1–)4–13 cm long,
(0.4–)1.3–4.2 cm wide, widest at or above the midpoint, obtuse and minutely
apiculate at apex, gradually narrowed below the middle to a narrowly cuneate
base, margin entire, recurved (in dried specimens), both surfaces glabrous and
glandular, coated especially when young with viscid glandular exudate, lateral
nerves 15–20(–30), often indistinct, petioles 0–2.5 mm long. Flowers greenish
yellow, unisexual or bisexual, in loose thyrsoid panicles at the ends of twigs, pedicels
0.8–1.5 cm long in flower and fruit, glabrous and glandular. Bisexual flowers
similar: sepals 3–4, greenish yellow, ovate, 2–2.5 mm long, spreading, anthers
1.6–1.9 mm long. Female flowers: calyx green, cup-like but sepals free, staminodes
lacking or with anthers 1.5 mm long, ovary green, oblong in outline, flattened,
bilobed, glandular, style 2–3-lobed, equalling the ovary, disk inconspicuous. Male
flowers:sepals 3–4, straw-colored or greenish yellow, 2–2.5 mm long, spreading,
stamens 7(–9), anthers 2–3 mm long, ovary rudiment minute. Male flowers: sepals
3–4, straw-colored or greenish yellow, 2–2.5 mm long, spreading, stamens 7(–9),
anthers 2–3 mm long, ovary rudiment minute. Female flowers: calyx green, cup-like
but sepals free, staminodes lacking or with anthers 1.5 mm long, ovary green,
oblong in outline, flattened, bilobed, glandular, style 2–3-lobed, equalling the ovary,
disk inconspicuous. Bisexual flowers similar: sepals 3–4, greenish yellow, ovate, 2–
2.5 mm long, spreading, anthers 1.6–1.9 mm long. Fruit disk-shaped, the body 1.5–
2.3 cm long, 8–11 mm wide, with an encircling papery venose wing 3–7 mm wide or
a few with 3(–4) locules and wings. Seeds 2 per locule, black, subspherical, or
somewhat compressed with a raised dorsal rim, 2–3 mm diameter, 1.5–3 mm thick.
The center of origin of Dodonaea viscosa is believed to be Australia, but it occurs
throughout the tropics and subtropics. Outside Australia, two varieties have been
recognized: var. viscosa, which in Africa occurs naturally along the coasts of West
Africa (from Senegal to Nigeria) and East Africa (from Kenya to Mozambique), and
in Madagascar, and var. angustifolia (L.f.) Benth., which occurs naturally from the
Democratic Republic of Congo in the west to Ethiopia and Somalia in the east, and
South Africa in the south, and also in Madagascar. Both varieties are now pantrop-
ical. Common undergrowth shrub in dry Afromontane forests (Bussmann 2002a, b,
2006) (Figs. 1, 2, 3, 4, 5, and 6).
Dodonaea viscosa Jacq. 437
Fig. 1 Dodonaea viscosa

(Sapindaceae). Chachapoyas,
Peru. (Photo R.W. Bussmann)



(Sapindaceae). Pusac, Peru.

Dodonaea viscosa Jacq. 439

Dodonaea viscosa: The plant is used in traditional medicine worldwide, adminis-

tered orally or as poultice to treat a great variety of ailments. Stem or leaf infusions
are used to treat sore throats, and root infusions are used to treat colds. The stems and
leaves are used to treat fever, and seeds (in combination with those of other plants
and coated in honey) are used to treat malaria. The stems are used as fumigants to
treat rheumatism. The leaves are used to relieve itching, and a lotion made from
unspecified plant parts is used to treat sprains, bruises, burns, and wounds. Digestive
system disorders, including indigestion, ulcers, diarrhea, and constipation are com-
monly treated in traditional medicine with an orally administered decoction of either
the leaves or roots. Trachoma is treated with applications of leaf juice, and powdered
leaves are given to expel roundworms. Pulverized roots are a component of anthel-
mintic preparations. The roots, either in decoction or fresh, are taken by women in
East Africa to stimulate milk production after giving birth and to treat dysmenor-
rhoea and irregular menstruation. In Ethiopia, women make an amulet of Dodonaea
viscosa and other plants wrapped in red silk thread and worn around the thigh to
ward off infertility. In some areas, twigs are used as toothbrushes. The roots are
boiled as galactagogue after childbirth. Leaf extract in cold water is used for diarrhea
and as styptic (Kokwaro 2009). Used to treat snakebites (Desta 1995). Seeds are
used to treat malaria (Giday et al. 2007).
Dodonaea viscosa: The wood is used for tool handles, tools, brooms, digging sticks,
spears, clubs, and poles, when available in large sizes, also for posts and frames for
permanent structures and in Kenya to make ribs of dhows (Beentje 1994). Through-
out its range, the species is an important source of fuelwood, and in East Africa, it is
favored for the production of charcoal. In tropical Africa, Dodonaea is browsed by

livestock and camels and used as an emergency fodder during the dry season. It is a
source of nectar and pollen for honey. The seeds yield a fish poison. Dodonaea
viscosa is used in reforestation, reclamation of marshes and degraded land, and as a
soil stabilizer. It makes a good live fence for dry areas and is planted as a windbreak.
It is commonly cultivated as an ornamental throughout its range on account of its
abundant fruiting and glossy leaves. Cultivars with leaves ranging from bluish-red,
pink, and red to yellowish-green have been developed. The roots are boiled as
galactagogue for cattle (Kokwaro 2009).
References
2006;11(1):41–66.
Dombeya torrida (J.F. Gmel.) Bamps
Dombeya sp.
MALVACEAE

Grace N. Njoroge
Synonyms
Dombeya torrida (J.F. Gmel.) Bamps: Dombeya goetzenii K. Schum.
Local names
Dombeya torrida: Kikuyu: Mueku; Maraket: Borawa; Maa: Ol-Subukiai,

Ol-subukiai-Orok; Elgeyo: Boloet, Silip; Kipsigis: Silibwet; Maraklwet: Borowa;
Meru: Mukeu; Nandi: Sibukuet, Silipchet; Samburu: Il-Porowai; Tugen: Boroa;
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Dombeya sp.: Trees or shrubs, deciduous when flowering, more rarely evergreen,
pubescent with stellate, simple, and glandular hairs. Leaves petiolate; blade 3–5( 7)
palmately lobed, or entire, base often cordate, margin serrate, dentate, or rarely
crenate; stipules narrowly triangular, caducous. Inflorescence axillary, pedunculate,
cymose, bracteate, subumbellate, or dichasial with 10–40 flowers, pedicellate; or
epicalycular bracts whorled, usually equal, caducous, rarely alternate on pedicel.
Sepals 5, valvate, united at base, triangular, often reflexed, nectariferous adaxially at
base, outer surface pubescent. Petals 5, contorted, spreading, white, sometimes
red-veined or based, or pink, asymmetrical, dimidiately ovate or elliptic, shortly
clawed, glabrous, brown-marcescent. Androgynophore absent; stamens in a single
whorl, filaments united in a short tube (but 1.5–1.6 cm long in Dombeya amaniensis)
at base, fertile stamens 10–15, in groups of 3 alternating with 5 slightly longer or
subequal ligulate-spatulate petaloid staminodes; anthers dehiscing by slits. Ovary
ovoid to oblate, tomentose or rarely glandular, with 3–5 locules each with 2–8
collateral ascending ovules, placentation axile, style as long as stamens, shortly
(2-)3–5-branched. Fruit spherical to ovoid, brown, dry, slightly woody, dehiscing
loculicidally by 3–5 valves. Seeds 1-several per locule, subreniform or angular,
brown or black, not fleshy (Figs. 1, 2, 3, 4, 5, 6, 7, 8, 9, 10 and 11).
Dombeya torrida (J.F. Gmel.) Bamps: Deciduous tree (7-)12–20 m tall; stems
terete, hispid to glabrous, hairs simple, stellate, tufted and subsessile- (rarely
stalked-) glandular. Leaves suborbicular, usually unlobed, 7–28 cm long, 5–20 cm
wide, apex acuminate, margin finely and closely serrate, base deeply cordate, upper
surface scabridulous, rarely glabrous, lower surface puberulous, or tomentellous,
frequently with tufts of hairs at base of main nerves; petiole 3.6–20 cm long; stipules
with the basal half transversely oblong or ellipsoid-oblong, the upper half narrowly
Fig. 1 Dombeya
sp. (Malvaceae) Farfangana,
Madagascar. (Photo
R.W. Bussmann)
Dombeya torrida (J.F. Gmel.) Bamps. . . 443
Fig. 2 Dombeya
Madagascar. (Photo
R.W. Bussmann)
Fig. 3 Dombeya
Madagascar. (Photo
R.W. Bussmann)
Fig. 4 Dombeya
Madagascar. (Photo
R.W. Bussmann)
triangular, 0.5–2.1 cm long, 0.1–0.8 cm wide, caducous to persistent. Inflorescence

axillary, 5.5–22 cm long, with two or more branches, each sub-umbellate, many-
flowered; peduncle 2.3–13.4 cm; bracts lanceolate, 0.7–1.7 cm long, 0.3–0.6 cm
wide, acuminate, glabrescent, caducous; branches usually once-forked near the apex,
bracteoles 1–1.2 cm long; epicalyx bracts inserted at calyx base, ovate, 0.4–1.1 cm
long, 0.2–0.5 cm wide; apex acute to acuminate, caducous. Calyx lobes erect or
reflexed, lanceolate-triangular, 0.5–1.3 0.2–0.4 cm. Petals pink or white, with or
without a red base or veins, 1.1–1.9 cm long. Androecium 0.8–1.8 cm long; staminal
tube dilating towards apex or sides convex; staminodes 0.4–1.3 cm long; longest
stamen 0.6–1.1 cm; anthers 0.2–0.3 cm long. Ovary globose or ovoid, tomentose;
style 0.6–1.1 cm long, 5 branched. Capsule globose, rarely depressed-ovoid, tomen-
tose; seeds ovoidoblong, 0.3 0.2 cm, dark brown. (Cheek and Dorr 2007).
Germinates mostly under light conditions, not easy to store (Bussmann and Lange
2000). Common understorey tree in disturbed Afromontane forests (Bussmann
2002a,b,c, 2004) (Figs. 12, 13, 14, and 15).
Fig. 5 Dombeya cf. lucida

(Malvaceae) Midugu-du Sud,
Madagascar. (Photo
R.W. Bussmann)

Madagascar. (Photo
R.W. Bussmann)

Madagascar. (Photo
R.W. Bussmann)

Madagascar. (Photo
R.W. Bussmann)
Dombeya torrida: In East Africa a decoction of the flowers and bark is taken against
indigestion, especially after using large quantities of meat (Bekalo et al. 2009;
Kokwaro 2009).
Donbeya burgessiae is used to treat ringworm (Njoroge and Buissmann 2007), and
cough (Njoroge and Bussmann 2006). Dombeya rotundifolia is used to treat
dysentery (Muithee et al. 2011).
Fig. 9 Dombeya sp. cf.

lucida (Malvaceae) Midugu-
du Sud, Madagascar. (Photo
R.W. Bussmann)
Fig. 10 Dombeya burgessiae

(Malvaceae) Sekenani,
R.W. Bussmann)
Fig. 11 Dombeya burgessiae

R.W. Bussmann)
Dombeya torrida: The wood is suitable for flooring, ship and boat building, vehicle
bodies, furniture, handles and ladders, sporting goods, agricultural implements,
yokes, poles, and piles. In tropical Africa it is mainly used for bows, construction,
poles, tool handles, and spoons. In Tanzania logs are carved into traditional stools.
Dombeya torrida is also used as firewood and for making charcoal. The fiber from
the bark is made into rope, string and cloth (Beentje 1994). It is eaten by animals but
rarely by livestock, used as firewood, to make spears and sticks, the bark serves for
strings, and gives lots of honey (Bussmann 2006). Fallen leaves improve the soil.
Dombeya burgessiae: Bark used for ropes, to join house posts, leaves as soap
replacement (Bussmann et al. 2006). Dombeya kirkii leaves are eaten by cattle,
and the species yields good honey. Also used to make ropes, and as firewood
(Bussmann et al. 2011). Dombeya rotundifolia: Eaten by animals but rarely by
livestock, used as firewood, to make spears and sticks, the bark serves for strings,
and gives lots of honey (Bussmann 2006).
Dombeya lucida is used in Madagascar for construction, and the bark to make ropes
(Rakotarivelo et al. 2013).
Fig. 12 Dombeya torrida

Chogoria, Kenya. (Photo
R.W. Bussmann)

R.W. Bussmann)

R.W. Bussmann)

R.W. Bussmann)
References
doi.org/10.1186/1746-4269-5-26.
appendices 1-7 @ www.naturekenya.org/JournalEANH.htm
Bussmann RW. Succession and regeneration patterns in East African Mountain forests – A Review.
Bussmann RW. Vegetation Ecology and Regeneration of Tropical Mountain Forests. In: Ambasht
RS, Ambasht NK, editors. Modern trends in applied terrestrial ecology. New York: Kluwer;
2002c. p. 195–223.
Mountain Forests: The role of natural and anthropogenic disturbance. Lyonia. 2004;61:98–111.
Nat Hist. 2000;891-2:101–11.
Cheek M, Dorr L. Flora of Tropical East Africa: Sterculiaceae. Kew: Royal Botanic Gardens; 2007.
L. Brown Center, MBG; 2013. ISBN 978-0-9848415
Dovyalis abyssinica (A. Rich.) Warb.
Dovyalis caffra (Hook. f. & Harv.) Warb.
Dovyalis macrocalyx (Oliv.) Warb.
SALICACEAE

Grace N. Njoroge
Synonyms
Dovyalis abyssinica (A. Rich.) Warb.: Aberia abyssinica Clos, Dovyalis engleri
Gilg, Flacourtia obtusa Hochst. ex Clos, Flacourtia obtusata Hochst. ex A. Rich.,
Hydnocarpus obtusa C. Presl, Rumea abyssinica A. Rich.
Dovyalis caffra (Hook. f. & Harv.) Warb.: Aberia caffra Hook. f. & Harv.
Dovyalis macrocalyx (Oliv.) Warb.: Aberia macrocalyx Oliv., Dovyalis adolfi-
friderici Mildbr. & Gilg, Dovyalis antunesii Gilg, Dovyalis chirindensis Engl.,
Dovyalis glandulosissima Gilg, Dovyalis luckii R.E. Fr., Dovyalis mildbraedii
Gilg, Dovyalis retusa Robyns & Lawalrée, Dovyalis salicifolia Gilg
Local Names
Dovyalis abyssinica: Kikuyu: Mukambura, Muirungi, Boni: Kurrawa, Chagga:

Maangwe, Elgeyo: Kibini, Kipchipinya, Ndorobo: Mukchat, Kisii:
Omokorogunyawa, Kipsigis: Nokiat, Luo: Sangla, Songla, Sangana, Maa:
R. W. Bussmann (*)
G. N. Njoroge

Ol-Morogi, Marakwet/Tugen: Mudariwa, Mintiriwa, Marakwet, Medililwa,

Mogonja, Meru: Muro, Nandi: Nukchat, Okiek: Tamugot, Pokot: Karatutwa,
Sebei: Mundariluluet, Samburu: Limoro, Lomoro, Imoro, Taita: Mubuchi,
Tugen: Kimkiet, Kinokuet, Turkana: Kudar, Ngomwo (Beentje 1994; Gachati
1989; Kokwaro 2009).
Dovyalis macrocalyx: Luhya: Likunga, Luo: Akudho
Botany and Ecology
Dovyalis abyssinica (A. Rich.) Warb.: Shrub or tree, much branched, up to 8 m tall,
trunk up to 20 cm. across, with pale greyish-brownish bark. Branchlets sometimes
with rather slender axillary spines 0.5–1.5(rarely 2.5) cm long. Leaf-blades ovate-
oblong or -elliptic, apex gradually obtusely attenuate, or sometimes rounded, base
broadly attenuate to obtuse, thinly to more firmly chartaceous, sparsely and very
shortly pubescent on the nerves beneath, glabrescent, entire to repand-crenate or
subserrate, (3-)4–9 cm long, 2–3.5( 4.5) cm broad, lateral nerves 4–5( 7) curved-
ascending pairs, slightly raised beneath, reddish in fresh specimens, no proper
reticulation, petiole 2–4( 5) mm long. Stamens 40–60. Stamens 40–60.
Interstaminal glands hairy apically. Male flowers 1–3 in a fascicle in generally
already defoliate axils, rusty-puberulous or tomentellous all over, pedicels 5–8
( 13) mm long. Calyx-lobes (4-)5(rarely 8), subovate-oblong, subacuminate,
5–6 mm long, 2–3 mm broad. Interstaminal glands hairy apically. Female flowers
solitary or rarely in pairs in defoliate axils, pedicels stoutish, 6–8 mm long. Male
flowers 1–3 in a fascicle in generally already defoliate axils, rusty-puberulous or
tomentellous all over, pedicels 5–8( 13) mm long Calyx-lobes similar to, though
slightly larger than those in the male flowers. Calyx-lobes (4-)5(rarely 8), subovate-
oblong, subacuminate, 5–6 mm long, 2–3 mm. broad. Disk annular, crenulate, finely
hairy. Disk annular, crenulate, finely hairy. Ovary rusty-puberulous to -velutinous,
styles 4–5(rarely 6). Female flowers solitary or rarely in pairs in defoliate axils,
pedicels stoutish, 6–8 mm long. Calyx-lobes similar to, though slightly larger than
those in the male flowers. Ovary rusty-puberulous to -velutinous, styles 4–5(rarely
6). Fruit globular, yellowish-reddish with minute paler dots, papillose-puberulous or
glabrescent towards maturity, 2 cm across. Seeds few, appressed hairy. Upland
rainforest to riparian and dry evergreen forest, sometimes in open wooded grassland,
1500–3000 m (Bussmann 2002; Sleumer 1975) (Figs. 1, 2, and 3).
Dovyalis caffra (Hook. f. & Harv.) Warb.: Tree or shrub up to 6 m tall, evergreen,
dioecious or rarely polygamous, armed with spines up to 6 cm long, bark ashen-
white to brown with longitudinal ridges and fissures, ultimate branchlets often much
abbreviated, up to about 2 cm long or forming cushion-like structures. Leaves
fascicled or, on young shoots, alternate, blade soft to coriaceous, obovate or some-
times elliptic-rhomboid, 2–5.5 cm long, 1–3 cm broad, glabrous or rarely
puberulous, with prominent venation on both sides, 3- to 5-veined from the base,
apex acute to obtuse or emarginate, base usually cuneate, margin entire, slightly
revolute, petiole 2–4 mm long, glabrous. Male flowers light green, fascicled, with
Dovyalis abyssinica (A. Rich.) Warb. . . . 455
Fig. 1 Dovyalis abyssinica

(Salicaceae), Bale Mountains
R.W. Bussmann)
5–10 flowers in groups of 3, calyx 2- to 5-lobed, lobes 3 mm long, elliptic,

pubescent, stamens numerous, filaments 5 mm long, nectaries hairy. Female flowers
light green, solitary or in fascicles of 2–3 on abbreviated shoots, pedicels 4–10 mm
long, calyx deeply 5- to 7-lobed, lobes 3 mm long, elliptic-lanceolate, puberulous,
acuminate, somewhat recurved, disc sparsely hairy, ovary 5- to 7-lobed, unilocular,
placentas 5–7, each placenta with 2 ovules, styles 5–7. Fruit subglobose, 3–6 cm in
diam., minutely velvety, bright yellow. Seeds many, woolly, 1 cm long (Sleumer
1975).
Dovyalis macrocalyx (Oliv.) Warb.: Bush or small tree up to 7 m tall, young
branches slender, pubescent at first, becoming glabrous, usually armed with slender,
straight spines up to 5 cm long. Leaf-lamina 2.5–9 1.4–4.3 cm, membranous or
thinly coriaceous, somewhat shining above, narrowly ovate, elliptic or ovate, apex
obtuse or subacute, broadly cuneate, obtuse or rarely slightly cordate at the base,
margin remotely crenulate or entire, glabrous on both sides, with 5–7 basal nerves,
venation laxly reticulate and subprominent on both sides, petiole 2 mm long,
puberulous. Male flowers in 1- to 4-flowered axillary fascicles, pedicels 2–3 mm
long, shortly and densely pubescent, calyx lobed almost to the base, lobes 5–6,

R.W. Bussmann)
3–4 1–2.5 mm, lanceolate to ovate-lanceolate, apex acute, margin entire or

occasionally with a few coarse teeth and sometimes with one or two glandular
hairs, stamens 20, filaments slender, 5 mm long, inter-staminal glands minute,
puberulous. Female flowers 1–2 per axil, calyx of 6–10 lobes divided almost to
the base, 6.5 2 mm, narrowly lanceolate, often incurved at the acuminate apex,
margin with dense stalked glands, both sides pubescent, annular disk pubescent,
undulate and segmented, ovary ovoid, pubescent with two pubescent divergent
styles 3 mm long, stigmas narrow, bilobed. Male flowers in 1- to 4-flowered axillary
fascicles, pedicels 2–3 mm long, shortly and densely pubescent, calyx lobed almost
to the base, lobes 5–6, 3–4 1–2.5 mm, lanceolate to ovate-lanceolate, apex acute,
margin entire or occasionally with a few coarse teeth and sometimes with one or two
glandular hairs, stamens 20, filaments slender, 5 mm long, inter-staminal glands
minute, puberulous. Female flowers 1–2 per axil, calyx of 6–10 lobes divided almost
to the base, 6.5 2 mm, narrowly lanceolate, often incurved at the acuminate apex,
margin with dense stalked glands, both sides pubescent, annular disk pubescent,
undulate and segmented, ovary ovoid, pubescent with two pubescent divergent
styles about 3 mm long, stigmas narrow, bilobed. Fruiting calyx accrescent and

R.W. Bussmann)
pinkish-green or red, the marginal glands are also much enlarged and become
fimbriate-ciliate, the calyx-lobes equal or slightly exceed the fruit. Fruit fleshy, red,
ellipsoid, puberulous, 2-seeded. Seeds 8 6 mm, obovoid, testa with a dense brown
wool (Sleumer 1975).
Dovyalis abyssinica: Root decoction used to treat venereal diseases, especially

gonorrjhoea, bilharzia, stomach-ache, and fever (Beentje 1994; Kokwaro 2009).
Leaf infusion used for indigestion (Kokwaro 2009). Used for chest pain, colds,
joint pain, stomachache, and as mouthwash (Delbanco et al. 2017; Gafna et al. 2017;
Nankaya et al. 2019), as well as indigestion (Muthee et al. 2011). Also employed as
steam bath for skin problems (Flatie et al. 2009; Yineger et al. 2008), and boiled
roots for postpartum weakness (Njoroge and Bussmann 2009). Used to treat malaria
(Njoroge and Bussmann 2006) and gonorrhea (Bussmann et al. 2006).
Dovyalis caffra: Used to treat malaria (Njoroge and Bussmann 2006).
Dovyalis macrocalyx: Root decoction drunk for headache and against nightmares.,
Bark decoction for venereal diseases (Kokwaro 2009).
Local Food Uses
Dovyalis abyssinica: Fruits edible (Beentje 1994; Bussmann 2006; Bussmann et al.
2011), but very sour.
Dovyalis abyssinica: Used for firewood, leaves browsed by cattle (Bussmann 2006;
Bussmann et al. 2011), and planted as live fence (Bussmann 2006).
Dovyalis caffra: Planted as live fence (Mekonnen et al. 2015).
Dovyalis macrocalyx: Used for firewood (Kiefer and Bussmann 2008).
References
importance, uses, and origin. Econ Bot. 2017;71(1):13–31.
Flatie T, Gedif T, Asres K, Gebre-Mariam T. Ethnomedical survey of Berta ethnic group Assosa
Zone, Benishangul-Gumuz regional state, mid-west Ethiopia. J Ethnobiol Ethnomed.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.

Ethnomed. 2006;2:8.
2009;8(2):262–9.
Sleumer HO. Flora of tropical East Africa: Flacourtiaceae. London: Crown Agents for Overseas
2008;2(6):132–53.
Dracaena afromontana Mildbr.
Dracaena steudneri Engl.
ASPARAGACEAE

Grace N. Njoroge
Local Names
Dracaena afromontana: Kikuyu: Mutharathare Muthari; Kipsigis: Labatiet; Maa:

Ol-ebenyan; Nandi: Lebekuet (Beentje 1994; Gachati 1989; Kokwaro 2009)
Dracaena steudneri: Kikuyu: Ithare; Swahili: Masanaka; Samburu: Sasaante; Sebei:
Mololosti; Taita: Issai; Tugen: Roke (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Dracaena afromontana Mildbr.: Shrub or tree, 1.5–7 m tall, palm-like, sometimes

scandent. Leaves 1.5–2.5 12–35 cm, lanceolate, narrow, base gradually
amplexicaul, apex acute. Flowers 13–15 mm long, whitish-cream, panicles 20–
60 cm long. Fruits orange, round, 15–18 cm. (Demissew 2006). A common com-
ponent of broad-leafed Afromontane forests (Bussmann 2002a, b) (Figs. 1 and 2).
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Dracaena afromontana (Asparagaceae), Mt. Kenya Nastional Park, Chogoria, Kenya.
Dracaena steudneri Engl.: Shrub or small tree, 2–12 m tall. Leaves 4–112 cm
wide 40–100 cm long, narrow, lanceolate, base slightly narrowed, widening to
amplexicaul base, apex acute. Flowers creamy-white to yellowish-green, in 30–
100 cm long panicles. Corolla 10–15 mm. Fruits orange, yellow or red, roundish.
10–15 mm wide (Demissew 2006). A common component of broad-leafed
Afromontane forests (Bussmann 2002a, b, c) (Figs. 3, 4, and 5).
Dracaena steudneri: The leaves are used to make a decoction for treating rabies
(Bekalo et al. 2009; Giday et al. 2010).
Dracaena gracilis and Dracaena reflexa are used medicinally in India (Reang
et al. 2016), and Dracaena reflexa is used in Madagascar for malaria, epilepsy, to
expel a retained placenta after birth and to treat placental apposition (Rakotoarivelo
et al. 2013). Also used to treat boils (Rabearivony et al. 2015). The stems and leaves
of Dracaena fragrans are employed to remedy cough, bronchitis and asthma
(Bussmann and Sharon 2006).
Local Food Uses
Dracaena reflexa: The flowers are eaten raw and used as condiment (Rakotoarivelo
et al. 2013).
Dracaena afromontana Mildbr. . . . 463
Fig. 2 Dracaena
afromontana (Asparagaceae),
Mt. Kenya Nastional Park,
R.W. Bussmann)
The leaves of both Dracaena afromontana and Dracaena ellenbeckiana are used to
weave small containers (Bussmann 2006).
Dracaena steudneri: The root is burnt and the smoke inhaled to treat evil eye
Dracaena xiphophylla is used in Madagascar ceremonially to attract a spouse
and to bless agricultural fields (Randrianarivony et al. 2016a, b). Dracaena mannii
is used as fishing poison (Neuwinger 2004).
Fig. 3 Dracaena steudneri

Nastional Park, Chogoria,
Kenya. (Photo
R.W. Bussmann)
Fig. 4 Dracaena ellenbeckiana (Asparagaceae), Laikipia Plateau, Kenya. (Photo R.W. Bussmann)
Dracaena afromontana Mildbr. . . . 465
Fig. 5 Dracaena ellenbeckiana (Asparagaceae), Laikipia Plateau, Kenya. (Photo R.W. Bussmann)
References
doi.org/10.1186/1746-4269-5-26.
p. 195–223.
Demissew S. Flora of tropical East Africa: Asparagaceae. Kew: Royal Botanic Gardens; 2006.

Randrianasolo A, Bussmann RW. Guide de plantes d’Ambalabe – Vol. 1. St. Louis: William
services from Agnalavelo sacred forest and their relationship with forest conservation. Madag
Conserv Dev. 2016a;11(2):44–51.
Randrianasolo A, Bussmann RW. Guide des plantes utiles d’Analavelona et de ses environs –
Vol. 1. St. Louis: MBG, William L. Brown Center; 2016b. ISBN 978-0-9960231-4-6.
Aromat Plants. 2016:5–2.
Drypetes gerrardii Hutch.
PUNTRANJIVACEAE

Grace N. Njoroge
Local Names
Drypetes gerrardii: Kikuyu: Munyenye, Luhya: Manamakulu, Sinamagulu, Maa:

Ol-Durdien, Samburu: Ikalai, Lecheniioibor (Beentje 1994; Gachati 1989).
Botany and Ecology
Drypetes gerrardii Hutch.: A much-branched evergreen shrub or tree up to 30 m,

with a densely leafy rounded crown and pendent branches. Seeds compressed-ovoid,
up to 1 cm long, mottled brownish. Female flowers: pedicels 1–2 mm long,
extending to 0.5–1(–2) cm in fruit, sepals as in the male disc shallowly cupular,
subentire, glabrous, ovary 2-locular, subglobose, 2 mm long, 2.5 mm wide, densely
fulvous-tomentose, styles 2, 1–1.5 mm long, united at the base, reflexed, persistent,
stigmas linear. Fruit obovoid-subglobose, usually shallowly bilobed, 1–1.5 cm long,
1–1.5 c. diameter, smooth, yellowish to golden-brown tomentose. Male flowers:
R. W. Bussmann (*)
G. N. Njoroge

pedicels 3–6(–8) mm long, evenly to densely yellowish-pubescent, sepals 4, sub-

orbicular-ovate, 2 mm long, 2 mm wide, evenly to densely yellowish-pubescent
without, sparingly to evenly pilose within, ciliate, yellowish green, yellow or creamy
white, fragrant, stamens 4, 2–3 mm long, anthers 1 mm long, disc crenulate to subentire,
plicate, the folds protruding a little between the stamens, smooth, with a small cylindric
central projection, glabrous. Young shoots and petioles yellowish-brown tomentose or
pubescent at first, later glabrescent. Petioles 3–7 mm long, leaf-blade ovate, ovate-
rhombic, elliptic-ovate or lanceolate, (2–)5–12(–17) cm long, (1–)2–6(–9) cm wide,
acute, subacute or obtuse, often slightly acuminate, somewhat asymmetrically
rounded-cuneate, shallowly and remotely crenate-dentate to subentire, thinly to mod-
erately coriaceous, shiny above, lateral nerves 6(–8) pairs, looped within the margin,
tertiary nerves reticulate, those nearest the midrib perpendicular to it and subparallel
for a short distance, slightly prominent above, more so beneath, sparingly yellowish-
pubescent above and more evenly so beneath, and sometimes more densely so on the
midrib beneath, or else quite glabrous above and beneath, dark green, drying brownish.
Stipules triangular-ovate, 1.3–1.5 mm long, sparingly to densely yellowish-pubescent,
soon deciduous. Flowers axillary, the males glomerulate-fasciculate with (2–)4–12(–
15) flowers per fascicle, produced after the females, the females usually solitary. Bark
fairly smooth, pale to dark greyish brown. Wood white, heavy. Twigs brownish, often
striate-sulcate (Smith 1987; Smith and Carter 1988). Important shrub in afro-tropical
mountain forests (Bussmann 2002, 2006).
Drypetes gerrardii: Good timber (Beentje 1994). Drypetes madagascariensis is

used for consttuctyion in Madagascar (Rakotoarivelo et al. 2013). The stem bark
of Drypetes arborescens, Drypetes dinklagei, Drypetes gossweileri, Drypetes
klainei, and Drypetes leonensis is used as fish poison (Neuwinger 2004).
References
2006;111:41–66.
1988.
Ehretia cymosa Thonn.
EHRETIACEAE

Grace N. Njoroge
Synonyms
Ehretia cymosa Thonn. Ehretia corymbosa Bojer ex A. DC.
Local Names
Ehretia cymosa: Kikuyu/Meru: Murembu, Mukui; Kipsigis: Mutereriet; Luhya:

Shekutu, Shikui; Maa: Endalai-ekolok; Marakwet: Kabinbonet, Morori; Nandi:
Boranet, Shamut; Okiek: Mundereriet; Chagga: Yambu; Samburu: Leachachur;
Taita: Mundana; Tugen: Makongiet, Buruburuti (Beentje 1994; Gachati 1989;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Ehretia cymosa Thonn.: Shrub or small tree 2–9( 20) m tall, with spreading crown
and often weak drooping branches. Young stems pubescent or glabrescent, soon
glabrous. Fruits orange, red or black, ovoid to subglobose, 2–6 mm in diameter,
apiculate, breaking up at maturity into 4 mericarps; pyrenes subovoid, 3.3 mm long,
2.8 mm wide; one inner face with an excavation towards the apex, the other not
excavated but with an indentation bordering the straight margin; outer face convex
with 4–5 ribs strongest towards the base and smooth towards the right-hand margin.
Leaf-blades elliptic or elliptic- to ovate-oblong, 7.5–20 cm long, 3.5–12 cm wide,
acuminate at the apex, cuneate, rounded or more or less subcordate at the base, glabrous
above save for the midrib and beneath save for pubescent nervation and hairy domatia,
more or less subcoriaceous; venation more or less impressed above, raised beneath;
petiole 1.2–3.5 cm long, glabrous, puberulous or hairy. Ovary ovoid, 1 mm long,
glabrous; heterostyly frequent; style white or pale purple, exserted 2–4 mm in long-
styled flowers, 0.7–0.8 mm long and included in short-styled flowers; stigmatic
branches 0.5–1 mm long. Flowers hermaphrodite, fragrant, in often copious panicles
of corymbose cymes up to 15 cm long and wide, the axes glabrous to pubescent or long
densely spreading hairy, sessile or subsessile or terminal flowers shortly pedicellate;
pedicels articulate at the base, 1–2( 3) mm long. Calyx campanulate, 1.5–2 mm long,
with 4–5 lanceolate lobes 0.7–1.2 mm long, acute, glabrous or puberulous, ciliate.
Corolla white, yellowish or pinkish white, campanulate, 4–8 mm long; tube 1.5–
3 mm long; lobes 4–5, oblong, 1.5–4 mm long, 1.5–1.8 mm wide, obtuse or subacute,
reflexed, glabrous or sparsely to densely ciliate at the apex. Stamens exserted; filaments
2.5–3.5 mm long; anthers brownish or blackish. (Verdcourt 1991). Important under-
growth tree in afro-tropical mountain forests (Bussmann 2002a, b, 2006). Slow germi-
nating under light conditions, not germinating in the dark (Bussmann and Lange 2000).
Ehretia cymosa: Roots and leaves toxic but used as aphrodisiac. Leaf juice applied
to wounds (Kokwaro 2009). The root is chewed for stomach ailments (Wondimu
et al. 2007).
Local Food Uses
Ehretia cymosa: The fruits are edible (Balemie and Kebebew 2006).
Ehretia cymosa: Used for utensils, tool handles, yokes, and firesticks
(Beentje 1994).
Ehretia cymosa Thonn. 471
References
Bussmann RW. Succession and regeneration patterns in east African Mountain forests - a review.
Bussmann RW, Lange SH. Germination of important east African mountain forest trees. J East Afr
Nat Hist. 2000;891-2:101–11.
Verdcourt B. Flora of tropical East Africa: Boraginaceae. Rotterdam: Balkema; 1991.
Ekebergia capensis Sparrm.
MELIACEAE

Grace N. Njoroge
Synonyms
Ekebergia capensis Sparrm.: Charia chevalieri C. DC., Charia indeniensis A. Chev.,

Ekebergia buchananii Harms, Ekebergia chevalieri (C. DC.) Harms, Ekebergia
complanata Baker f., Ekebergia holtzii Harms, Ekebergia indeniensis (A. Chev.)
Harms ex Engl., Ekebergia meyeri C. Presl ex C. DC., Ekebergia mildbraedii Harms,
Ekebergia petitiana A. Rich., Sorindeia doeringii Engl. & K. Krause, Trichilia capensis
(Sparrm.) Pers., Trichilia ekebergia E. Mey. ex Sond., Trichilia rueppelliana Fresen.
Local Names
Ekebergia capensis: Kikuyu: Mukuria-Hungu, Murera-Hungu, Mununga, Swahili:

Mpoto wa ndovu mkuu, Marakwet: Kerebut, Bajun: Mpotowanduvu-Mkuu,
Kamba: Kyuasi, Mukongui, Boni: Mrongoleh, Ndorobo: Orouet, Rararuet,
R. W. Bussmann (*)
G. N. Njoroge

Oroyuet, Kipsigis: Araruet, Ardwi, Kisii: Omonyamavi, Luo: Tido, Maa:

Ol-Subukiai, Meru: Muchogomo, Nandi: Teldet, Sebei: Bumet, Kipumetet,
Taita: Mbo, Manuka-Masi, Tugen: Temwaa Arariet, Kondilwa, Turkana:
Eng’amwo (Beentje 1994; Gachati 1989; Kokwaro 2009), English: Cape ash,
Dog plum, Mountain ash, Ekebergia,
Botany and Ecology
Ekebergia capensis Sparrm.: Medium-sized evergreen or semi-evergreen tree up to

30 m tall, but usually less, bole slightly buttressed or fluted at base, up to 1 m
diameter at breast height, second-year branchlets slender, usually less than 0.6 cm
diameter, smooth, with scattered leaf-scars, closely lenticellate with large whitish
lenticels. Leaves imparipinnate, usually entirely glabrous, sometimes pubescent,
rarely tomentose, petiole and rhachis up to 35 cm long, leaflets opposite or sub-
opposite, 7–15, subsessile or shortly petiolulate, lanceolate to oblong-lanceolate, up
to 14.5 6 cm, usually much smaller, tapering to an acuminate or subacuminate
apex, base asymmetric, lower surface rarely drying whitish. Flowers white or pinkish
white, sweet-scented, borne in many-flowered cymose panicles. Calyx 0.2 cm long,
sparsely to densely puberulous. Petals elliptic-oblong, 0.4–0.5 cm long, densely
puberulous on both surfaces. Staminal tube 0.2 cm long, puberulous outside, densely
bearded at throat inside. Ovary 0.15 0.2 cm, densely setulose, style 0.05–0.1 cm
long. Fruit a drupe 1.5 1.5 cm, deep red, with 2–4 pyrenes. Montane, mid-altitude
and riparian forest, often at edges, more rarely (Uganda) in woodland and wooded
grassland, 600–2650 m. Ekebergia capensis is widespread, from Senegal east to
Eritrea and Ethiopia, and south to Botswana, eastern South Africa and Swaziland.
(Styles and White 1991). Common tree in Cassipourea malosana and Juniperus
procera dominated Afromontane forests (Bussmann 2002a, b) (Figs. 1, 2, 3, 4, 5,
and 6).
Ekebergia capensis: Bark decoctions, infusions and macerations are taken to treat
gastritis, heartburn, dysentery, epilepsy, gonorrhea and as vermifuge, and are applied
externally to ulcers, abscesses, boils, scabies, acne, pimples and itching skin. A
powder prepared with the bark is sniffed against headache, colds, and sinusitis. A
root decoction is taken as a diuretic and to treat kidney problems, dysentery,
heartburn, headache, and respiratory complaints. The root is chewed as an expecto-
rant. Charred pulverized roots are sniffed for treatment of headache and blocked
nose. Leaf macerations are used internally or externally to treat headache, fever,
cough, and skin complaints, and they are taken as a vermifuge. The wood is used by
Zulu people to facilitate childbirth. Decoctions of various parts of Ekebergia
capensis are used traditionally in central Ethiopia as an anthelmintic for the treat-
ment of livestock. Bark and roots have been used as ordeal poisons. Root chewed as
Ekebergia capensis Sparrm. 475
Fig. 1 Ekebergia capensis

(Meliaceae), Bale Mountains
Ethiopia. (Photo R.W:
Bussmann)
Fig. 2 Ekebergia capensis (Meliaceae), Bale Mountains National Park, Demaro forest, Ethiopia.
(Photo R.W: Bussmann)

Bussmann)
expectorant. Bark decoction in steam bath to treat itchy rashes (Kokwaro 2009).
Used as antimalarial (Njoroge and Bussmann 2006). The galls on Ekeberigia leaves
are used for abortions (Njoroge and Bussmann 2009).
Local Food Uses
Ekebergia capensis: The fruit is edible.
Ekebergia capensis: The wood is locally valued for furniture, and it is also used for
light construction, poles and tool handles. It is suitable for light flooring, joinery,
interior trim, ship building, vehicle bodies, sporting goods, toys, novelties, vats, food
containers, boxes, crates, matches, turnery, veneer, and plywood. It is also used as
firewood and for charcoal production (Beentje 1994). In southern Africa the bark is

Bussmann)
Fig. 5 Ekebergia capensis (Meliaceae), Bale Mountains National Park, Demaro forest, Ethiopia.

Bussmann)
used for tanning. The foliage is browsed by livestock in the dry season. Ekebergia
capensis is planted as an ornamental, particularly as a roadside tree, but also as a
garden tree for its attractively colored fruits and for shade. It is occasionally planted
for soil conservation, as a windbreak and as a shade tree in coffee and banana
plantations. The flowers are a source of nectar and pollen for honeybees. Used as
firewood (Bussmann et al. 2011), and as fish poison (Neuwinger 2004).
References

Ethnomed. 2006;2:8.
2009;8(2):262–9.
Styles BT, White F. Flora of tropical East Africa: Meliaceae. Rotterdam: Balkema; 1991.
Englerina woodfordioides (Schweinf.) Balle
ex M.G. Gilbert
LORANTHACEAE

Grace N. Njoroge
Synonyms
Englerina woodfordioides (Schweinf.) Balle ex M.G. Gilbert: Englerina

woodfordioides var. adolfi-friderici (Engl. & Krause) Balle, Englerina woodfordioides
var. umbelliflora (De Wild.) Balle, Ischnanthus ehlersii (Schweinf.) Tiegh., Ischnanthus
woodfordioides (Schweinf.) Tiegh., Loranthus adolfi-friderici Engl. & Krause, Loranthus
bagshawei Rendle, Loranthus brachyphyllus Peter, Loranthus ehlersii Schweinf.,
Loranthus eucalyptoides Peter, Loranthus longifolius Peter, Loranthus rugegensis Engl.
& Krause, Loranthus tschertscherensis Pax, Loranthus umbelliflorus De Wild., Loranthus
viminalis Engl. & Krause, Loranthus woodfordioides Schweinf., Tapinanthus adolfi-
friderici (Engl. & Krause) Danser, Tapinanthus bagshawei (Rendle) Danser, Tapinanthus
brachyphyllus (Peter) Danser, Tapinanthus ehlersii (Schweinf.) Danser, Tapinanthus
eucalyptoides (Peter) Danser, Tapinanthus longifolius (Peter) Dansere, Tapinanthus
rugegensis (Engl. & Krause) Danser, Tapinanthus tschertscherensis (Pax) Danser,
Tapinanthus umbelliflorus (De Wild.) Danser, Tapinanthus viminalis (Engl. & Krause)
Danser, Tapinanthus woodfordioides (Schweinf.) Danser
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Englerina woodfordioides: Kikuyu: Kieha, Kipsigis: Ketarwet (Gachati 1989;

Kokwaro 2009).
Botany and Ecology
Englerina woodfordioides (Schweinf.) Balle ex M.G. Gilbert: Shrub with spread-

ing to drooping stems to 1 m. or so, twigs slightly compressed, glabrous. Calyx 0.5–
1 mm long, ciliolate. Receptacle 1–1.5 mm long. Flowers 6–16( 20) per umbel,
peduncle 1–5( 6) mm long, pedicels (1-)2–5 mm long, bract ovate-triangular from a
shallow cup, 1–2 mm long, slightly gibbous, ciliolate. Petiole 3–10( 15) mm long,
grooved, the sides often crispate, the channel sometimes scurfy puberulous, lamina
thin, coppery reddish when young, becoming fresh green, often with red veins, and a
little more coriaceous, linear-lanceolate to oblong-lanceolate or ovate-lanceolate, 3–
13( 15) cm long, 0.6–3( 5) cm wide, often attenuate, acuminate at the apex, base
cuneate, rounded or rarely slightly cordate, glabrous except sometimes at youngest
unfolded stage, with 4–8 pairs of lateral nerves, the lower 2–4 often (but not always)
strongly ascending. Berry bright red, obovoid, with prominent lobed disc, 6–9 mm
long, 3.5–5 mm in diameter, seed orange. Corolla (1.1 )1.5–2.5 cm long, blue-grey
with red lobes, inside tube blue-black, buds slightly inflated in lower part, narrow
above vents to oblong-ellipsoid apical swelling (2 )2.5–3.5( 4) mm long, tube
(4 )6–12 mm long, with V-slit to base, papillate inside especially along sutures,
lobes erect, linear with spathulate tips, (7 )8–15 mm long. Filaments blackish
below, green above, tightly coiled, tooth 0.5–1.2 mm long, anthers 1.5–3 mm
long. (Polhill and Wiens 1999). Common in Afromontane forests dominated by
Faurea saligna and Ilex mitis (Bussmann 2002). Mostly useless for the local
population (Bussmann 2006; Bussmann et al. 2011) (Figs. 1, 2, and 3).
Englerina woodfordioides: Leaf and bark decoction drunk for liver problems.
Heated twigs are applied as poultice for pneumonia (Kokwaro 2009). The whole
plant is used to treat common cold (Njoroge and Bussmann 2006). Fresh leaves used
to treat earache (Yineger and Yewhalaw 2007).
Englerina woodfordioides (Schweinf.) Balle ex M.G. Gilbert 483
Fig. 1 Englerina woodfordioides (Loranthaceae). Mt. Kenya National Park, Sirimon, Kenya.
References
Polhill RM, Wiens D. Loranthaceae. In: Flora of tropical East Africa. Rotterdam: Balkema; 1999.
121 pp. ISBN 90-6191-383-7. Reference page.
Ensete ventricosum (Welw.) Cheesm.
MUSACEAE
Rainer W. Bussmann, Narel Y. Paniagua-Zambrana,

Grace N. Njoroge, and John Grimshaw
Synonyms
Ensete ventricosum (Welw.) Cheesm.: Ensete edule Bruce ex Hiran., Musa arnoldiana
De Wild., Musa davyae Stapf, Musa ensete J.F. Gmel., Musa ventricosa Welw.
Local Names
Ensete ventricosum: Kikuyu: Ihindu, Thendu, Luo/Luganda: Kitembe, Meru:

Mukobo, Kipsigis/Nandi: Sasuriet, Kamba: Iklulutui, Maa: Ol-musalala,
Marakwet: Sosurwa, Kipsigis/Nandi: Sasuriet, Meru: Gikobo, Mukoko, Taita:
Isengeru (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge
J. Grimshaw
The Yorkshire Arboretum, Castle Howard, York, UK
e-mail: director@yorkshirearboretum.org

Botany and Ecology
Ensete ventricosum (Welw.) Cheesm.: Giant herb arising from a short upright
rhizome. Bracts of the male part of the inflorescence persistent or partially decidu-
ous, each subtending 30–40 flowers. Outer tepal of male flowers 3-lobed, the lobes
variable in length, 3.5–5.5 cm long, white with orange-yellow tips, inner tepal
serrate-apiculate, 1–1.5 1–1.7 cm, the apiculum 0.3–1.3 cm, or occasionally
absent, stamens 5, 3–5 cm long, anthers violet to purple, filaments white, staminode
present or not, acicular, 0.1–1 cm long, style acicular, 1–2 cm long. Bracts of the
female/hermaphrodite part of the inflorescence persistent, partially covering the
fruits. Is the basis of an agricultural system in southern Ethiopia. Seeds irregularly
subspherical, 1.2–2.3 1.2–1.8 0.9–1.6 cm, striate to smooth, hard, black,
embedded in orange pulp. Pseudostem formed of the overlapping leaf-bases,
1.5–5 m. tall. Leaf-blades erect or spreading, forming a large rosette, oblong-
lanceolate, to 5 1.5 m., glaucous or not, midrib red or green. Outer tepal
3-lobed, sometimes with 1–2 smaller extra acicular lobes attached to it internally,
inner tepals 1–3, variable in shape with 2 wings and an apiculum up to 1.5 cm long,
stamens 0–5, 3.5 cm long, colored as in the male flowers, staminodes variable
according to the number of stamens present, style 2.5–4 cm long, terete, with a
large capitate stigma. Fruits 5–20 in the axil of each bract, long-obovoid, 8–15 3–
4.5 cm, orange at maturity. Inflorescence appearing from the center of the rosette,
pendulous when mature. (Lock 1993). Naturally growing in wet, lower Afromontate
forests (Bussmann 2002) (Figs. 1 and 2).
Ensete ventricosum: Stems boiled for liver problems. Leaf decoction to prevent
miscarriages. The liquid accumulating at the leaf bases is drunk to avoid afterbirth
problems. Leaf and fruit juice taken for liver problems. The burned stalk ash is
employed in a similar way. Root decoctions are used for ear problems and stomach-
ache. The pseudo-stem sap serves top clean snakebites (Kokwaro 2009). The root
decoction is used to treat bone fractures (Giday et al. 2010; Mekonen et al. 2015).
Powdered seeds of Ensete perrieri is used in Madagascar to treat eye problems like
cataracts and conjunctivitis; A leaf decoction serves to relieve colics
Local Food Uses
Ensete ventricosum: One of the most important staples in the Afromontane forest
region of Ethiopia. Often grown together with Coffeea arabica and Colocasia
esculenta (Bekalo et al. 2009; Luizza et al. 2013; Mekonen et al. 2015).
Ensete ventricosum (Welw.) Cheesm. 487
Fig. 1 Ensete ventricosum (Musaceae), cultivated, Oromiya Province, Ethiopia. (Photo

J. Grimshaw)
Fig. 2 Ensete ventricosum (Musaceae), cultivated, Oromiya Province, Ethiopia. (Photo

J. Grimshaw)
Ensete ventricosum: Petioles yield strong, fine fibers. Leaves used as thatch
(Beentje 1994).
References
doi.org/10.1186/1746-4269-5-26.
Lock JM. Flora of tropical East Africa: Musaceae. Rotterdam: Balkema; 1993.
Res Appl. 2013;11:315–39.
Mekonen T, Giday M, Kelbessa K. Ethnobotanical study of homegarden plants in Sebeta-Awas 72
District of the Oromia region of Ethiopia to assess use, species diversity and management 73
practices. J Ethnobiol Ethnomed. 2015;11(64).
Randrianarivony TN, Andriamihajarivo TH, Rakotoarivony F, Rabarimanarivo M, Randrianasolo A,
Bussmann RW. Guide des plantes utiles d’Analavelona et de ses environs – Vol. 1. St. Louis:
William L. Brown Center, MBG; 2016b. ISBN 978-0-9960231-4-6.
Eragrostis tef (Zuccagni) Trotter
POACEAE

Grace N. Njoroge
Synonyms
Cynodon abyssinicus (Jacq.) Raspali; Eragrostis abessinica (Jacq.) Link; Eragrostis

abyssinica (Jacq.) Link; Eragrostis pilosa subsp. abyssinica (Jacq.) Asch. &
Graebn.; Eragrostis pilosa var. tef (Zuccagni) Fiori; Poa abyssinica Jacq.; Poa
cerealis Zuccagni; Poa tef Zuccagni
Local Names
Eragrostis tef: Tef
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Eragrostis tef (Zuccagni) Trotter: Annual; culms 20–90 cm high, erect. Leaf-
blades flat, up to 30 cm long and 4 mm wide. Panicle lanceolate to broadly ovate,
10–40 cm long, diffuse or contracted, with slender pedicels and long flexuous
branches of which the lowest form a whorl, the axils hairy or glabrous. Spikelets
4- to 12-flowered, narrowly oblong, 5.5–9 mm long, 1.5–2 mm wide, straw-colored
to dark green or reddish, the florets at first appressed to the rhachilla, but diverging as
the grain swells, remaining intact at maturity; glumes lanceolate, acuminate,
unequal, the lower 1.2–2.5 mm long and 0- to 1-nerved, the upper 1.7–3 mm.;
lemmas oblong-elliptic, 2–2.7 mm long, acute; palea scaberulous on the keels,
eventually falling very tardily with the lemma; anthers 3, 0.3–0.6 mm long. Cary-
opsis oblong, 1–1.2 mm long, turgid, white or reddish brown. Weedy places; 750–
2500 m. A staple cereal crop in Ethiopia, introduced experimentally to other tropical
countries, and sometimes found as an adventiv. Tef originated in northern Ethiopia,
where it is widely cultivated. Details of its domestication are unknown, but it may
predate the introduction of wheat and barley to the region (Fig. 1).
Fig. 1 Eragrostis tef (Poaceae) threshing in Bale region, Ethiopia. (Photo R.W: Bussmann)
Eragrostis tef (Zuccagni) Trotter 491
Local Food Uses
Eragrostis tef: Grain cultivation of Tef has been confined mainly to Ethiopia and to
some extent the highlands of Eritrea. It is also grown in northern Kenya. Small-scale
commercial tef production takes place in South Africa, the United States, Canada,
Australia, Europe (the Netherlands), and Yemen. Tef is grown as a forage grass, for
instance in South Africa, Morocco, Australia, India, and Pakistan. It has been
introduced experimentally into other tropical countries, either for its grain or for
hay, e.g. in other parts of East Africa and in southern Africa. It is commonly found as
an escape from cultivation.
In Ethiopia and Eritrea Tef flour is mainly used to prepare “injera,” a thin, flat,
pancake-like bread, made from dough fermented for 2–3 days. “Injera” is prepared
in a range of sizes and is consumed with various sauces (“wot”), based on meat or
pulses. Tef flour produces the best quality “injera”: pliable, soft with glossy appear-
ance, which does not fall apart under handling or stick to the fingers and has a
slightly sour taste. Fenugreek (Trigonella foenum-graecum L.) can be added to Tef
flour in a small proportion to improve the “injera” flavor. It also increases the lysine
content. Tef flour is also mixed with barley or sorghum flour to make “injera.” Other
traditional preparations from tef flour include “kitta” (unleavened bread), “atmit” or
“muk” (gruel), porridge and local alcoholic beverages. Several recipes that fit
Western tastes have been developed from tef flour particularly in the United States,
where it has found niches in the health food market and as a gourmet food. Tef flour
is used as a thickening agent in a range of products, including soups, stews, gravies,
and puddings (Bekalo et al. 2009; Giday et al. 2007, 2009; Kioko et al. 2015;
Teklehaymanot et al. 2007; Teklehaymanot 2009; Teklehaymanot and Giday 2010;
Yineger and Yewhalaw 2007; Yineger et al. 2008).
In Ethiopia tef straw is used as forage, especially during the dry season. Mixed with
clay it is used as plastering material for local houses and to make bricks, stoves,
granaries, beds, and pottery.
References
doi.org/10.1186/1746-4269-5-26.

borne diseases among Maasai people in northern Tanzania. Vet World. 2015;812.
District, Jimma Zone, southwestern Ethiopia. J Ethnobiol Ethnomed. 2007;3:24.
10.1186/1746-4269-4-11.
Erica arborea L.
ERICACEAE

Grace N. Njoroge
Local Names
Erica arborea: Kikuyu: Muthithinda; Kipsigis: Kaibeyuwunyot; Kwaipeyot; Maa:

Olikbejus; Marakwet: Kwaloliong; Nandi: Sisinuet (Beentje 1994; Gachati 1989).
Botany and Ecology
Erica arborea L.: Shrub or tree 0.3–7.5 m high, trunk diameter to at least 5 cm;
much-branched, with ascending branches, the branches red-brown; stems pubescent
with short smooth hairs and longer dendritic hairs to 1 mm. Leaves in whorls of 3–4,
appressedor ascending, needle-like, 2–6.5 mm long, 0.5–1 mm wide, margins
minutely denticulate, glabrous; petiole 0.4–0.7 mm long, glabrous. Flowers clus-
tered towards the end of short lateral branches, where their density may give
impression of continuous flowers along branches; pedicels 1.5–4 mm long, glabrous
R. W. Bussmann (*)
G. N. Njoroge

or rarely with a few hairs, with bract and 2 ciliolate bracteoles to 1.3 mm long below
the middle. Calyx 4-merous, 1.1–1.9 mm long, the lobes ovate, 0.6–1.2 mm long,
ciliolate (sometimes with dendritic hairs) to glabrous, saccate at base. Corolla white
or pink, campanulate, pendulous, 1.5–4 mm long, widest at the mouth or almost so;
stamens 8, included in corolla, anthers bifid and dehiscing by an oblique pore, with
two small dorsal/basal appendages; disc present; ovary glabrous, style 0.5–2 mm
long, stigma capitate, 0.2–0.5 mm in diameter. Fruit a glabrous capsule. (Beentje
2006). A common invader in burnt Afromontane timberline forest areas (Bussmann
1996; Bytebier and Bussmann 2000). In itself a very common component at the
upper reaches of Afromontane forests dominated by Hagenia abyssinica and Hyper-
icum revolutum and forming forests of its own together with Ginidia glauca
(Bussmann 2002a, b, 2006) (Figs. 1, 2, 3, 4, and 5).
Erica arborea: The smoke of the wood is inhaled to treat respiratory infections
(Luizza et al. 2013).
Fig. 1 Erica arborea (Ericaceae), background, Mt. Kenya National Park, Naro Moru Route
Moorlands, Kenya. (Photo R.W. Bussmann)
Erica arborea L. 495
Fig. 2 Erica arborea

(Ericaceae), Mt. Kenya
Route Moorlands, Kenya.
Erica arborea: Used as forage for livestock, firewood, and honey production


Fig. 5 Erica wytheana (Ericaceae), Mt. Kenya National Park, Naro Moru Route Moorlands,
References
Beentje H. Flora of tropical East Africa: Ericaceae. Kew: Royal Botanic Gardens; 2006.
Bussmann RW. Islands in the desert -forest vegetation of Kenya’s smaller mountains and highland
Bussmann RW. Succession and regeneration patterns in east African Mountain forests -a review.
2006;11(1):41–66.
Res Appl. 2013;11:315–39.
Erythrina abyssinica DC
FABACEAE

Grace N. Njoroge
Synonyms
Erythrina abyssinica DC: Chirocalyx abyssinicus (Lam.) Hochst.; Chirocalyx

tomentosus Hochst.; Corallodendron suberifra (Welw. ex Baker) Kuntze; Erythrina
abyssinica subsp. suberifera (Welw. ex Baker) Kuntze; Erythrina bequartei De
Wild.; Erythrina comosa Hua; Erythrina eggelingii Baker f.; Erythrina huillensis
Welw. ex Baker; Erythrina kassneri Baker f.; Erythrina mossambicensis Sim;
Erythrina pelligera Fenzl; Erythrina platyphylla Baker f.; Erythrina suberifera
Welw. ex Baker; Erythrina tomentosa R. Br.; Erythrina tomentosa R. Br. ex Richard;
Erythrina tomentosa var. longicauda Baker f.; Erythrina warneckei Baker f.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Erythrina abyssinica: Kikuyu: Muhuti; Maa: Ekirikiti, Ol-Goroshe; Kamba: Kivuti,

Muvuti; Acholi: Lucoro; Swahili: Mjafari, Mbamba Ngoma; Luo: Murembe, Orembe;
Luhya: Omurembe, Kumurembe, Umtembe; Meru: Muuti; Nyankore/Rutoro:
Omuko; Iraqwi: Qanqari; Duruma: Mgalla; Kipsigis: Kipisorwet, Kogoruet;
Marakwet: Gorgorwa; Maa: Ol-Opongi; Nandi: Kakaruet; Pokot: Korkorwo;
Samburu: Garacha; Sebei: Kaborte (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Erythrina abyssinica DC.: Tree, rarely a shrub, 3–10 m tall, with a rounded crown,
bark brown, thick, corky, deeply longitudinally fissured, young branches usually
spiny. Terminal leaflet 6–14 6–16 cm, broadly elliptic to sub-rotund, often broader
than long, lateral leaflets 4.5–14 3–11 cm, broadly ovate-rhombic, asymmetric at
the base, petiole 8–17 cm long, armed with scattered prickles, rhachis 3–8 cm long,
petiolules 3–12 mm long, stipules 3–7 mm long, linear-triangular. Inflorescence a
terminal pseudoraceme, peduncle 4–16 cm long, densely tomentose, rhachis
5–20 cm long, bracts 6–8 2–3 mm, oblong, caducous, pedicels 2–5 mm long,
bracteoles 4–8 1 mm, filiform, caducous. Calyx spathaceous, split fully by the
emergent corolla along its abaxial edge, densely tomentose, tube 0.6–1.8 cm long,
lobes 0.6–4.5 cm long, shape very variable, filiform, linear, elliptic, obovate or
turbinate. Corolla red, glabrous, standard 2.8–4 1.1–1.5 cm, elliptic-oblong,
wings 7–11 4–6 mm, keel petals free, 5–6 3–4 mm. Vexillary stamen joined
only at the base, others fused along half their length into a tube. Ovary 1.5–2.5 cm,
narrowly cylindrical-oblong. Pod brown to black 4.8–10.2 cm, curved, moniliform,
woody, pubescent, 1–10-seeded. Seeds red, 6–12 5–7 3–5 mm, hilum black,
5–7 2–3 mm, narrowly elliptic. A scattered tree in wooded grassland, in open
deciduous woodland and on rocky hillsides, can be grown as a “living fence” from
truncheons 200–1400 m (Gillett et al. 1971a, b) (Figs. 1, 2, 3, 4, 5, and 6).
Fig. 1 Erythrina edulis
(Fabaceae), Leymebamba,
Pweu. (Photo R. W.
Bussmann & N. Y. Paniagua-
Zambrana)
Erythrina abyssinica DC 503

Pweu. (Photo R. W.
Zambrana)

Pweu. (Photo R. W.
Zambrana)
Erythrina abyssinica: Bark of young stems used for eye inflammation, also to treat
gonirrhea. Root decoctions are used for malaria, syphilis and snakebite and abdominal
pain Roasted bark is applied to swellings. Bark decoctions remedy gall- and kidney

Pweu. (Photo R. W.
Zambrana)
Fig. 5 Erythrina arborescens

(Fabaceae), Ilam, Nepal, at an
altitude of 2300 m at
Maipokhari. (Photo K. R.
Bhattarai)
problems and stones, as well as duodenal ulcers (Bekalo et al. 2009; Kokwaro 2009).
Also used to treat infertility and urinary tract problems (Muthee et al. 2011). The bark is
boiled to relieve postpartum pain (Njoroge and Bussmann 2009). The extract is also
used for wound treatment and has shown antibacterial activity (Njoroge and Bussmann
2007). The boiled bark extract is also used as antimalarial (Njoroge and Bussmann
2006a), and for cough and nosebleeds (Njoroge and Bussmann 2006b). In Ethiopia
used for gastrointestinal problems (Giday et al. 2007). Extracts of Erythrina
arborescens serve to treat stomach disorders (Kunwar et al. 2012).
Local Food Uses
Some species, e.g., Erythrina edulis are eaten (Bussmann and Sharon 2006).
Erythrina abyssinica DC 505
Fig. 6 Erythrina arborescens

(Fabaceae), Ilam, Nepal, at an
altitude of 2300 m at
Maipokhari. (Photo K. R.
Bhattarai)
Erythrina abyssinica: Wood used for stools, beehives, and doors (Beentje 1994). A
leaf decoction is used to treat blackleg in livestock (Yineger et al. 2007). Sometimes
the bark smoke is used ceremonially (Mohageghazdeh and Faridi 2006). The Bark
decoction used to remedy East Coast Fever in cattle (Kokwaro 2009). Used also as
firewood (Kiefer and Bussmann 2008). Erythrina brucei is planted as shade species
(Mekonnen et al. 2015).
References
doi.org/10.1186/1746-4269-5-26.
Papilionoideae, part 1. London: Crown Agents for Overseas Governments & Administrations;
1971a.
Papilionoideae, part 2. London: Crown Agents for Overseas Governments & Administrations;
1971b.
species in far-West Nepal – a valuable resource being wasted. J Mt Sci. 2012;9:589–600.
2009;8(2):262–9.
Erythrococca bongensis Pax
EUPHORBIACEAE

Grace N. Njoroge
Local Names
Erytrococca bongensis: Kikuyu: Muhare-Ngware, Muharangare; Kipsigis:

Kusisitiet; Luhya: Shiriedzo; Maa: Enduleleindu; Somali: Nitatumbe; Samburu:
Itulelei, Lechapirik; Tugen: Kipkaris; Turkana: Ekoromwai; Chagga: Kimbumbu;
Padhola: Ngesa; Luo: Adieremo (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Erythrococca bongensis Pax: Fruit tricoccous, or by abortion dicoccous, the cocci

3.5–4 mm diameter, sparingly adpressed pubescent, greenish or purplish. Female
flowers: pedicels 2–4 mm long, stouter than in ♂, calyx-lobes smaller than in ♂,
minutely ciliolate, disc-glands 3, ovate, flattened, scale-like, ovary trilobate, 1 mm
diameter, evenly adpressed pubescent, styles free, 1 mm long, reflexed, stigmas
R. W. Bussmann (*)
G. N. Njoroge

fimbriate-laciniate throughout, white. An erect or semi-scandent shrub or small tree

up to 6 m tall, but more commonly to 3.5 m. Female inflorescences fewer-flowered
than male, but otherwise similar. Male flowers: pedicels slender, flexuous, extending
to 1.2 cm at anthesis, glabrous, calyx-lobes 3(–4), triangular-ovate, 1.5 mm long,
1 mm wide, acute, glabrous, greenish cream, extrastaminal disc-glands 7, small,
rounded, distinct, glabrous, interstaminal glands numerous, erect, angular, truncate,
glabrous, dark green, stamens 9–15, 5–7 peripheral, 4–8 central, 0.5 mm long,
filaments broadened in the middle, purple-grey, anthers white or cream. Male
inflorescences densely glomerulate, sessile or shortly pedunculate, 0.5–1.5 cm
long, peduncles pubescent or glabrous, bracts minute, ciliate. Stipules aculeolate,
2 mm long, straight or hooked, pungent, slightly accrescent, yellowish. Petioles (2–)
3–9 mm long, leaf-blades ovate, elliptic or elliptic-lanceolate, (1–)3–7.5(–12) cm
long, (0.5–)1–3(–5) cm wide, obtuse or sometimes subacute at the apex, cuneate at
the base, crenate-serrate, often irregularly so, chartaceous, brittle, lateral nerves 5–7
pairs, commonly looped well within the margin, sometimes not looped, not prom-
inent above, slightly so below, sparingly pubescent or puberulous along the midrib
and main nerves above and beneath at first, later becoming quite glabrous, dark
green, paler beneath, often purplish tinged at first. Young shoots and petioles evenly
pubescent, soon glabrescent. Seeds 3–3.5 mm diameter, shallowly foveolate-
reticulate, aril orange-yellow or red. Twigs pale grey-green, not prominently spar-
ingly lenticellate. Bark light grey or brown, flaking. (Smith 1987; Smith and Carter
1988). Common in the undergrowth of semi-wet Afromontane forests, especially in
natural Coffea forests (Bussmann 2002a, b) (Fig. 1).
Fig. 1 Erythrococca
bnongensis (Euphorbiaceae),
Park, Odo-Bulu forest,
Ethiopia. (Photo R. W.
Bussmann)
Erythrococca bongensis Pax 509
Erythrococca bongensis: Leaf juice is used as cure for cough, stomach pain, and
AIDS (Kokwaro 2009).
Local Food Uses
Erytrococca bongensis: The fruits are commonly eaten, especially by children

Erytrococca bongensis: Wood used to make arrows. Leaf infusion to treat swellings
in cattle (Maasai) (Beentje 1994). Leaf extract used to treat black quarter in cattle
(Kokwaro 2009).
References
appendices 1–7 http://www.naturekenya.org/JournalEANH.htm.
1988.
Eucalyptus globulus Labill.
MYRTACEAE

Grace N. Njoroge
Local Names
Eucalyptus globulus: Kikuyu: Mubau, Muringamu (Gachati 1989; Kokwaro

2009).
Botany and Ecology
Eucalyptus globulus Labill.: A large or medium tree, up to 50 m tall, bark smooth,

bluish, deciduous except in lower part of trunk, juvenile leaves opposite, a consid-
erable number of pairs, sessile to amplexicaul, glaucous, cordate or ovate or broadly
lanceolate, 7–16 cm long and 1–9 cm broad, mature leaves alternate, petioled, dark
green, lustrous, lanceolate or cordate-lanceolate, acuminate, 10–30 cm long and
3–4 cm broad, flower buds usually solitary, sessile or very short stalked, tubular,
4-angled, warty or tuberculate, 30 mm long and 20 mm broad, operculum
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Eucalyptus
sp. (Myrtaceae), Batumi,
Georgia. (Photos R. W.
Zambrana)
cup-shaped, with dome-shaped appendage, warty, sometimes much shorter than

tube, anther obovate, dehiscing by broad parallel slits, visible on anterior side,
fruit sessile, depressed-spherical or broadly tubular, warty, with 4 main ribs and
1 to several short subsidiary ribs, 10–15 mm long and 15–30 mm broad, disk large,
convex, very thick. The main component is cineolic acid (58–70%). Eucalyptus
globulus is native to Tasmania and south-eastern Australia but is now widely planted
and naturalized in subtropical regions around the world (Macbride and Weberbauer
1936–1995) (Figs. 1, 2, 3, 4, 5, 6, and 7).
Eucalyptus globulus: Leaves boiled, and steam inhaled to treat flu, fever, and colds
(Kokwaro 2009). Used to treat malaria (Njoroge and Bussmann 2006a), pimples and
ringworm (Njoroge and Bussmann 2007), sexually transmitted diseases (Njoroge
and Bussmann 2009), and respiratory disorders (Njoroge and Bussmann 2006b). as
well as colds, flu, pimples, and pneumonia (Njoroge et al. 2004). In Madagascar it is
used to treat stomachache (Razafindraibe et al. 2013; Rabearivony et al. 2015) and
malaria (Rakotoarivelo et al. 2015), and is often found in local markets
(Randriamiharisoa et al. 2015). The bark and leaf decoction used as medicine in
Pakistan to treat diabetes (Ullah et al. 2019). Sold in local markets in Madagascar to
treat severe headache and colds, and abdominal pain (Rabearivony et al. 2015;
Eucalyptus globulus Labill. 513
Fig. 2 Eucalyptus
Zambrana)
Fig. 3 Eucalyptus
Zambrana)
Randriamiharisoa et al. 2015). The extract is used as antibacterial for wounds

(Muhammad et al. 2019). Sometimes used to treat hypertension (Malik et al.
2018). In Ethiopia used for rheumatism, stomach problems, constipation, fever, and
flu (Bekalo et al. 2009; Teklehaymanot et al. 2007; Wondimu et al. 2007; Yineger
et al. 2008). The smoke is used as expectorant and to relieve colds (Mohagheghzadeh
and Faridi 2006). For cold and cough often used as steam bath (Gedif and Hahn 2003).
Eucalyptus camadulensis leaves are used to treat malaria (Mekonnen et al. 2015;
Rakotoarivelo et al. 2015).
Fig. 4 Eucalyptus
Zambrana)
Fig. 5 Eucalyptus
Zambrana)
Fig. 6 Eucalyptus
Zambrana)
Eucalyptus globulus: Used in Kenya to treat respiratory tract problems in cattle

(Njoroge and Bussmann 2006c).Very frequently planted, and most widely used as
firewood. Used against ectoparasites in livestock (Wanzala 2017). The smoke is used
as insect repellent (Karunamoorthi et al. 2009a, b).
Fig. 7 Eucalyptus
Zambrana)
References
doi.org/10.1186/1746-4269-5-26.
Karunamoorthi K, Ilango K, Endale A. Ethnobotanical survey of knowledge and usage custom of
traditional insect/mosquito repellent plants among the Ethiopian Oromo ethnic group.
J Ethnopharmacol. 2009a;125:224–9.
J Ethnopharmacol. 2009b;121:49–53.

Malik K, Ahmad M, Bussmann RW, Ahmad M, Zafar M, Tariq A, Alqahtani A, Shahat A, Ullah R,
Rashid N, Zafar M, Sultana S, Nasar Shah S. Ethnobotanty of hypertensive plants used in
Northern areas of Pakistan. Front Pharmacol. 2018;9(789) https://doi.org/10.3389/
fphar.2018.00789.
of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006c;108:332–9.
2009;82:262–9.
org/10.1186/s13002-015-0046.
2013;9:73.
Appl. 2019;18 https://doi.org/10.32859/era.18.8.1-20.
2017:8647919, 33 pages. https://doi.org/10.1155/2017/8647919.
2008;26:132–53.
Euclea divinorum Hiern.
EBENACEAE

Grace N. Njoroge
Synonyms
Euclea divinorum Hiern.: Euclea balfourii Hiern ex Balf. f.; Euclea huillensis
Gürke; Euclea katangensis De Wild.; Euclea keniensis R.E. Fr.; Euclea kiwuensis
Gürke; Euclea laurina Hiern ex Balf. f.; Euclea stuhlmannii Gürke
Local Names
Euclea divinorum: Kikuyu/Kamba: Mukuthi, Kikuthi, Mukunyai; Swahili: Mdaa,

Mdala, Msirisha; Kipsigis/Tugen/Sebei: Uswa, Uswet; Luhya: Kumuchanjasi;
Luo: Ochol; Maa: Ol-Kinyei; Marakwet: Jeptuiya; Samburu: Lechingei, Lchinge;
Sebei: Shiendet, Cheptuishak, Wuswet; Tugen: Kanarape; Chagga: Mkenye
(Beentje 1994; Gachati 1989; Kokwaro 2009); English: Magic gwarri, Diamond-
leaved Euclea.
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Euclea divinorum Hiern.: Evergreen shrub or small tree 1.8–9(–15) m tall, bark grey-
brown, red-brown, pale mauve-grey or cinnamon or much darker and almost black,
mostly rough with shallow irregular longitudinal fissures, flaking longitudinally, slash
pink to crimson or pale yellow to orange, sapwood ochraceous or cream, turning apricot
on exposure to air, young shoots with rusty peltate scales but otherwise glabrous.
Leaves drying grey-green to chestnut, opposite or subopposite, mostly diamond-shaped
(sub-rhombic) or narrowly rhombic-lanceolate, widest near the middle, 1.6–12 cm
long, 0.6–4.5 cm wide, narrowed to a rounded or emarginate apex, attenuate to
narrowly or fairly broadly rounded at the base, the margin often crinkly, glabrous
save for rusty peltate scales beneath, venation sometimes slightly raised above but
inconspicuous, petiole 4–6 mm long. Inflorescences paired in at least some axils,
unbranched, dense and contracted, up to 1.5 cm long. Male flowers 3.5 mm long,
calyx yellow, patelliform with short broadly triangular teeth, glabrous save for rusty
scales, corolla creamy white, pale yellowish or salmon buff, the lobes sometimes with a
pale brown spot, deeply lobed, widely open at the throat, each lobe with more than
25 striga-like hairs along the mid-petaline line, stamens 16, strigulose, rudimentary
ovary with 2 simple or bilobed styles. Female flowers without staminodes, ovary
yellow, densely strigulose. Male flowers 3.5 mm. long, calyx yellow, patelliform with
short broadly triangular teeth, glabrous save for rusty scales, corolla creamy white, pale
yellowish or salmon buff, the lobes sometimes with a pale brown spot, deeply lobed,
widely open at the throat, each lobe with more than 25 striga-like hairs along the
mid-petaline line, stamens 16, strigulose, rudimentary ovary with 2 simple or bilobed
styles. Female flowers without staminodes, ovary yellow, densely strigulose. Fruit
brown to black, globose, 5–7 mm diameter, with short white hairs. Grassland, mostly
with scattered trees, open bushland, thicket on anthills, secondary forest, margins of
evergreen forest, often on stony slopes, 0–2700 m. Euclea divinorum is widespread
from Sudan and Ethiopia to Namibia, Botswana, and South Africa (White and
Verdcourt 1996). Common in the undergrowth of dry, Juniperus procera dominated
Afromontane forests (Bussmann 2002a, b) (Figs. 1 and 2).
Euclea divinorum: Fruits are taken as a mild laxative but can have strong purgative
action. They are applied in the fermentation process to make beer. The roots and twigs
are popular as toothbrushes and roots are chewed as a disinfectant and to color the lips
and mouth red. In traditional medicine root extracts and dried powdered roots are
applied for the treatment of gastro-intestinal disturbances, cancer, ulcers, wounds,
arthritis, miscarriage, jaundice, snakebites, and gonorrhea. The Shangaan people apply
ground roots on the skin against leprosy and to relieve headache and toothache. Root
decoction used as purgative (Luo) and bark infusion to increase appetite (Kipsigis)
(Beentje 1994). Root boiled with Croton megalocarpus to treat chest pain, pneumonia,
abscesses, and as purgative and anthelminthic, also as tonic. Used in purification
Euclea divinorum Hiern. 521
Fig. 1 Euclea divinorum

(Ebenaceae). Mature leaves,
Naro Moru, kenya. (Photo
R. W. Bussmann)
Fig. 2 Euclea divinorum

(Ebenaceae). Sapling,
Naro Moru, kenya. (Photo
R. W. Bussmann)
rituals (Sebei). Root also chewed for toothache, and bark pounded and applied to
snakebites (Kokwaro 2009). The root is boiled against malaria and as anthelminthic.
The branches serve as toothbrush (Bussmann et al. 2006). Both Euclea divinorum and
Euclea schimperi show antiviral properties (Gebre-Mariam et al. 2006). Euclea
divinorum serves to treat malaria, leprosy, gonorrhea, syphilis, amoebas, rabies, and
scabies (Geyid et al. 2005), as well as constipation (Muthee et al. 2011). The bark
decoction is used to treat excessive menstrual flow and diarrhea (Njoroge and
Bussmann 2009; Njoroge et al. 2004). Decoctions of the whole plant serve as
anthelminthic and for stomach probmels (Nankaya et al. 2019). Euclea schimperi is
used for stomach problems (Giday et al. 2003; Lulekal et al. 2008). The smoke of
Euclea natalita is used to relieve headache (Mohagheghzadeh and Faridi 2006).
Euclea racemosa bark serves to treat kidney problems (Wondimu et al. 2007).
Local Food Uses
Euclea divinorum: In East Africa, an infusion of the bark is used as a condiment in

the preparation of a fatty meat and milk soup. The thin fleshy part of the fruits is
edible (Bussmann et al. 2006). In western Kenya, Euclea divinorum is one of the
species used in the treatment of milk to increase its digestibility, palatability, and
preservation, sometimes treated milk can be stored for over a year without turning
bad, which is vital to the household food security.
Euclea divinorum: The bark is collected from the wild and used to produce fast
reddish-brown dyes, and for tanning hides and skins. Depending on how long the
fiber to be dyed is left in the boiling dye extract, the color obtained can range from
pale to dark brown or black. In Tanzania and southern Africa, the roots are pounded
and boiled to dye the various vegetable fibers used in the weaving of mats into
various shades of brown and even black when a mordant of iron-rich mud is applied
after dyeing. When used to dye wool, the roots give a wide range of brown colors.
Leather produced by tanning with Euclea divinorum bark has a deep red color due to
the numerous naphthoquinones present in the plant. In south-western Ethiopia,
Euclea divinorum is used to purify drinking water by adding branches to the gourds
or pots and leaving them to soak in the water for several hours. The fruits are used as
a purple dye for basket fibers by the Kwanyama Ovambos of northern Namibia and
also boiled to produce purple ink. In East Africa, the wood is used for tool handles
and carving, and as firewood. Occasionally, Euclea divinorum serves as an orna-
mental and shade tree. Leaves are used to wrap meat to keep it fresh. Also used for
firewood and to tan leather (Bussmann et al. 2006). The boiled fruits are used for
anaplasmosis in cattle (Njoroge and Bussmann 2006).
References
Euclea divinorum Hiern. 523
Gebre-Mariam T, Neubert RS, Schmidt PC, Wutzler P, Schmidtke M. Antiviral activities of some
Ethiopian medicinal plants used for the treatment of dermatological disorders. J Ethnopharmacol.
2006;104:182–7.
Geyid A, Abebe D, Debella A, Makonnen Z, Aberra F, Teka F, Kebede T, Urga K, Yersaw K,
Biza T, Haile Mariam B, Guta M. Screening of some medicinal plants of Ethiopia for their anti-
10.1186/1746-4269-4-10.
2009;8(2):262–9.
White F, Verdcourt B. Flora of tropical East Africa: Ebenaceae. Rotterdam: Balkema; 1996.
Euphorbia candelabrum Welw.
Euphorbia tirucalli L.
EUPHORBIACEAE

Grace N. Njoroge
Synonyms
Euphorbia tirucalli L.: Arthrothamnus tiruncalli (L.) Klotzsch & Barcke; Euphor-
bia geayi Constantin & Gallaud; Euphorbia laro Drake; Euphorbia media N.E. Br.;
Euphorbia rhipaloides Lem.; Euphorbia rhipsaloides Willd.; Euphorbia scoparia
N.E. Br.; Euphorbia suareziana Croizat; Euphorbia tiruncalli var. rhipsaloides
(Willd.) A. Chev.
Local Names
Euphorbia candelabrum: Kikuyu: Kithuri, Mubububngu, Githuri, Muthuri,

Kibubung’i; Luo: Bondo; Kipsigis: Kuresyet; Maa: Ol-Pongoni, Ol-Bobongo;
Swahili: Mtupa, Mtungutungii; Boni: Baraidi; Borana: Adama; Digo: Ganga;
Kamba: Kyaa; Luhya: Ludua; Somali: Darkhen; Tugen: Kuress; Turkana:
Yoopong (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Euphorbia tiruncallii: Kikuyu: Kariaria, Ndaru, Nyanjoe; Marakwet: Asubgwa;

Shambaa: Ngesa; Luo: Ojuok; Swahili: Mtupa mwitu; Digo: Utudi; Ilwana: Dana;
Kamba: Ndau; Maa: Ol-Oile; Oromo: Wadida; Pokot: Tumwen; Somali: Dana
Botany and Ecology
Euphorbia candelabrum Welw.: Tree to 12( 20) m high, with a simple trunk to
90 cm in diameter and rough fissured grey bark; branches persistent from 3 m
upwards, suberect, densely rebranching to form eventually a large broadly rounded
crown; terminal branchlets fleshy, (3–)4( 5)-angled, 5–10 cm wide, square in cross-
section to distinctly winged, with wings to 2.5 cm wide, usually constricted at
irregular intervals into oblong segments 15–25 cm long; angles straight to shallowly
sinuately toothed, with teeth 1–1.5 cm apart. Seeds subglobose, slightly compressed
laterally, 3 mm in diameter, greyish brown speckled with paler brown, smooth.
Capsule shortly exserted on a stout pedicel 5 mm long, 2–3-locular, subglobose,
8 12 mm, fleshy, green becoming red, hardening immediately before dehiscence to
6 9 mm and very obtusely 2–3-lobed. Spine-shields very obtusely triangular, to
8 7 mm, extending 5 mm above to include the flowering eye; spines stout, to 5 mm
long; prickles flexible, triangular, 1.5 mm long, soon deciduous; spines and spine-
shields soon becoming corky, rusty-brown and disintegrating. Leaves oblanceolate
on seedlings and young growth, to 7 1.5 cm, deltoid on older growth and soon
deciduous, 5 5 mm Cymes 1–6 crowded together, 1-forked, with stout peduncles
5–20 mm long and cyme-branches 5 m. long; bracts rounded, 5 6 mm Cyathia
4 9 mm, with broadly cup-shaped involucres; glands transversely elliptic,
2 4 mm, almost touching, golden yellow; lobes transversely elliptic,
2.2 3 mm Male flowers many: bracteoles spathulate, plumose; stamens 5.8 mm
long. Female flower: perianth irregularly divided into 3 or more filiform lobes 2–4 mm
long, sometimes with 1 or 2 teeth; styles 2 or 3, 3 mm long, joined at the base, apices
thickened, rugulose, bifid. (Smith 1987; Smith and Carter 1988). In both riverine, and
dry parts of Afromontane forests (Bussmann 2002a, b, 2004, 2006) (Figs. 1, 2, 3, and 4).
Euphorbia tirucalli L.: Trees or shrubs, 2–6 m tall, dioecious, eventually forming
trunk, 10–25 cm d.b.h.; bark rugose, gray or light. Stems green, succulent, very
finely longitudinally striate. Leaves alternate, present only on new growth; stipules
very small, caducous; petiole absent; leaf blade oblong-linear, 7–15 0.7–1.5 mm,
base attenuate, margin entire, apex obtuse. Cyathia clustered at apex of branches,
pedunculate, unisexual; involucral leaves minute, membranous, caducous; involucre
turbinate, ca. 2 1.5 mm, shortly pubescent inside; glands 5, peltate-ovate or
subrounded. Male flowers many, exserted from involucre. Female flower: ovary
glabrous, exserted from involucre; styles connate below middle; stigma 2-lobed.
Capsule 3-lobed, ca. 8 8 mm, smooth, sparsely pilose or glabrous. Seeds ovoid-
globose, ca. 4 4 mm, smooth; caruncle small. Flowering and fruiting July–October
(Wu et al. 1994–2013).
Euphorbia candelabrum Welw. . . . 527
Fig. 1 Euphorbia
candelabrum
(Euphorbiaceae), Laikipia
plain, Samburu, Kenya.
(Photo R. W. Bussmann)
Fig. 2 Euphorbia
candelabrum
Euphorbia candelabrum: A stem decoction is used after childbirth to expel the

placenta. The bark is used for skin diseases (Kokwaro 2009). A decoction of the pith
is used after childbirth (Maasai) (Beentje 1994). The latex is used for wounds and
Fig. 3 Euphorbia
candelabrum
Fig. 4 Euphorbia
candelabrum
plain, Samburu, Kenya (Photo
R. W. Bussmann)
warts (Njoroge and Bussmann 2007). The species serves also to treat joint pain,
general venereal diseases and infertility (Muthee et al. 2011). Used to treat gonorrhea
(Wondimu et al. 2007).
Euphorbia tiruncallii: The juice is used for sore throat and stomach problems. The
boiled root is eaten as emetic to treat snakebite, and also to treat sterility in women.
The plant is highly toxic, and any use requires high care (Kokwaro 2009). Used for
female health (Delbanco et al. 2017). Also used for malaria (Njoroge and Bussmann
2006a). The latex is used for warts (Njoroge and Bussmann 2007). Also used to treat
gonorrhea, hemorrhages, and help to ease delivery (Wondimu et al. 2007;
Teklehaymanot et al. 2007).
Many other species of Euphorbia are used medicinally. Euphorbia schimperiana
is used to treat syphilis and Euphorbia dumalis is used for the same purpose
(Bussmann et al. 2011). Euphorbia chamaesyce is used to treat constipation and
dysentery in children (Bhat et al. 2013). Euphorbia hypericifolia is used to treat
menorrhagia and diarrhea (Bhat et al. 2013). Euphorbia pilosa is used to treat
constipation and is used as an emetic in cases of food poisoning (Bhat et al. 2013).
Euphorbia macroceras serves for tooth pain (Bussmann et al. 2018). Euphorbia
hirta is used for bronchial infections, asthma, and warts (Kumar et al. 2011; Kunwar
et al. 2010a, b, 2015). In Madagascar, the species serves to treaty urine retention and
gonorrhea (Razafindraibe et al. 2013; Rabearivony et al. 2015), and for bronchitis in
infants, as well as premature closure of the fontanelle (Randrianarivony et al. 2016).
In Pakistan used to treat diabetes (Ullah et al. 2019). Euphorbia royleana latex is
used to treat arthritis (Kunwar et al. 2009, 2010a, b, 2013, 2016, 2019; Kunwar and
Bussmann 2009), for fractures (Singh et al. 2019), and as antiseptic (Malik et al.
2015). Euphorbia thomsoniana is employed in Pakistan for wet and dry eczema
(Sher et al. 2016). Euphorbia dracunculoides is used against skin parasites, acne,
snakebites, and epilepsy (Umair et al. 2019). Euphorbia prostrata has traditional
uses for diarrhea, hepatic ulcers, eczema, bladder stones, and is used as blood purifier
(Umair et al. 2019) and to treat diabetes (Ullah et al. 2019). Euphorbia pilulifera
serves for cough, bronchial asthma, diarrhea, indigestion, pain, burns, cuts, and
wounds (Umair et al. 2019). Euphorbia helioscopia serves to treat fungal infections
like athlete’s foot, eye sores, asthma, cholera, and constipation and serves as
anthelminthic (Muhammad et al. 2019; Umair et al. 2019). Euphorbia hetero-
chroma serves to treat fever and glandular problems (Delbanco et al. 2017).
Euphorbia amlpiphylla latex is used for warts (Giday et al. 2010; Mekonnen et al.
2015) and to treat rabies (Giday et al. 2007). Euphorbia crotonoides serves for
stomach problems (Giday et al. 2003). Euphorbia joyae is used for tonsillitis
(Njoroge and Bussmann 2006b) as well as postpartum hemorrhages (Njoroge and
Bussmann 2007). Eupohorbia cuneata is used for gonorrhea (Muthee et al. 2011).
Euphorbia alluaudii is used in Madagascar for dental treatments (Randrianarivony
et al. 2016). Euphorbia abyssinica serves to remedy venereal diseases and rabies
(Teklehaymanot et al. 2007), and ringworm (Teklehaymanot 2009). Euphorbia
breviarticulata is used for trachoma and other eye problems (Teklehaymanot
et al. 2010).
Local Food Uses
Euphorbia tiruncallii: Young branches may be roasted and chewed (Kokwaro

2009).
Euphorbia candelabrum: Roasted bark is applied to infected wounds and for

brucellosis in livestock, sap is applied to calves’ cheeks to treat East Coast Fever
(Kokwaro 2009). Euphorbia tiruncallii: Used as fish poison, and often planted as
hedge (Beentje 1994; Neuwinger 2004).
Euophorbia helioscopia and Euphorbia schimperiana used for skin problems in
livestock (Ali et al. 2019; Bekalo et al. 2009) and to treat anthrax (Yineger et al. 2007).
In Pakistan, Euphorbia helioscopia is used to repel the devil and diseases (Ahmad Jan
et al. 2019; Sher et al. 2016). Many species of Euphorbia are used to kill ectoparasites
in livestock (Kioko et al. 2015, Wanzala 2017). Euphorbia mandravioky is used to
better handle cattle, and as glue to catch birds (Randrianarivony et al. 2016).
References
2019. https://doi.org/10.1016/j.chnaes.2019.12.005.
Sher H, Bussmann RW, Shao J-W. Traditional uses of plants by Indigenous Communities for
veterinary practices at Kurram District, Pakistan. Ethnobot Res Appl. 2019;18:24. https://doi.
org/10.32859/era.18.24.1-19.
doi.org/10.1186/1746-4269-5-26.
in Kedarnath Wildlife Sanctuary of Garhwal Himalaya, India. J Ethnobiol Ethnomed. 2013;9(1).
p. 195–223.
2006;11(1):41–66.
Bussmann RW, Paniagua-Zambrana NY,Wood N, Ole Njapit S, Ole Njapit JN, Ene Osoi GE, Kasoe
Delbanco AS, Burgess ND, Cuni_Sanchez A. Medicinal plant trade in Northern Kenya: economic
borne diseases among Maasai people in Northern Tanzania. Vet World. 2015;8(6):755–62. 12.
Kunwar RM, Bussmann RW. Medicinal, Aromatic and Dye Plants of Baitadi and Darchula
Districts, Nepal Himalaya: status, uses and management. In: Hartmann M, Weipert J, editors.
Biodiversität, Naturausstattung im Himalaya, Bd. III. Erfurt: Verein der Freunde & Förderer des
Naturkundemuseums; 2009. p. 475–89.
macology of medicinal plants in far-west Nepal. Ethnobot Res Appl. 2009;7:5–28.
Kunwar RM, Burlakoti C, Chowdhary CL, Bussmann RW. Medicinal plants in Farwest Nepal: their
indigenous uses and pharmacological validity. Med Aromat Plant Sci Biotechnol. 2010a;4
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in far-west Nepal: a
pharmacological appraisal. J Ethnobiol Ethnomed. 2010b;6:35.
western Nepal. PlosONE. 2016. https://doi.org/10.1371/journal.pone.0167812.
Kunwar RM, Shrestha K, Malla S, Acharya T, Sementelli A, Kutal D, Bussmann RW. Relation of
medicinal plants, their use patterns and availability in the lower Kailash Sacred Landscape,
Nepal. Ethnobot Res Appl. 2019;18(7) https://doi.org/10.32859/era.18.6.1-14.

2009;82:262–9.
Madagascar. Indian J Tradit Knowl. 2016;151:68–78.
2013;9:73.
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;18(14) https://
doi.org/10.32859/era.18.14.1-11.
1988.
Appl. 2019;18(8) https://doi.org/10.32859/era.18.8.1-20.
10.1186/s13002-019-0285-4.
2017:8647919, 33 pages. https://doi.org/10.1155/2017/8647919.
Wu Z, Raven PH, Hong D, editors. Flora of China. Beijing/St. Louis: Science Press/Missouri
Fagaropsis angolensis (Engl.) Dale
RUTACEAE

Grace N. Njoroge
Local Names
Fagaropsis angolensis: Kikuyu: Mukaragati; Meru: Murumu; Luhya:

Shingulotso; Marakwet: Kwiril; Meru: Mukuriampungu; Nandi: Noiywet
Botany and Ecology
Fagaropsis angolensis (Engl.) Dale: Deciduous tree to 15 m. Leaves 12–30 cm,

leaflets opposite, in 2–4 pairs, lamina ovate or oblong-ovate to elliptic, 4–9 2–
4 cm, softly hairy on both surfaces, margin entire, with a row of closely spaced
pellucid gland dots, apex acute or acuminate, base asymmetric. Inflorescence of
panicles arising at the end of the previous year’s branches. Flowers 4-merous. Petals
4, 5–6 mm long, oblanceolate. Male flowers: stamens 8. Female flowers: staminodes
present, ovary 4-locular. Fruit 6–7 mm in diameter (Kokwaro 1982). Characteristic
R. W. Bussmann (*)
G. N. Njoroge

for the ground-layer of Brachylaena huilklensis – Elaeodendron buchananii domi-

nates deciduous Afromontane forests (Bussmann 2002).
Fagaropsis angolensis: The bark decoction is used by the Kikuyu to treat malaria
(Njoroge and Bussmann 2006).
Fagaropsis angolensis: Wood used for furniture (Beentje 1994).
References
appendices 1–7 http://www.naturekenya.org/JournalEANH.htm
Ethnomed. 2006;2:8.
Faurea saligna Harv.
PROTEACEAE

Grace N. Njoroge
Local Names
Faurea saligna: Kikuyu: Muthorothua; Sebei: Maiyokoro; Marakwet: Sirite,

Markwa, Maiyokwa; Maasai: Ol-Gerian, Ol-Orten; Bemba: Saninga; Kamba:
Kikaati, Mukaati; Luhya: Mugonju, Kukuyekwe; Nandi: Mosomboriet; Sebei:
Bwonget, Maakwet; Tugen: Musomboriet; Cherangani: Markwa; Elgeyo: Siriki;
Samburu: Ol-Bugui, Ilgilma; Ndorobo: Nkoma; Pokot: Muyokwa (Beentje 1994;
Botany and Ecology
Faurea saligna Harv.: Small to medium-sized tree up to 17 m high, usually with a

slender straight trunk to first branch. Spikes up to 12 cm long, terminal, peduncle
pubescent with greyish hairs, bracts deltoid, puberulous to pubescent Perianth pink
in bud, pale green when expanded but pinkish and brownish at base and apex of tube
respectively, 1.2 cm long, tube with a slightly bulbous tip in bud 3–4 mm long,
R. W. Bussmann (*)
G. N. Njoroge

pubescent without with greyish appressed hairs, densely so when young. Anthers
2.5–3 mm long. Young branchlets light brown with a grey appressed pubescence,
glabrescent. Leaves glaucous green, upper surface sometimes slightly shining,
lanceolate-elliptic or narrowly lanceolate-elliptic, sometimes slightly falcate, up to
16 cm long and rarely more than 3–3.5 cm broad, usually 5–6 times as long as broad,
acute at apex with a mucronate tip, tapering towards the base with a distinct petiole
up to 1.5 cm long, midrib distinctly visible above but not prominent, on lower
surface yellow and prominent, but secondary venation not obviously visible,
surfaces entirely glabrous above except when very young, entirely glabrous beneath
or sometimes with a few minute appressed hairs towards the base of the lamina,
petioles pink or red, mostly glabrous or with minute appressed hairs. Hypogynous
scales variable, ovate-triangular, 1–1.5 mm long (Brummitt 1993). Forming dense
Afromontane forests especially on very steep slopes, together with Ilex mitis
(Bussmann 2002a, b; Bytebier and Bussmann 2000).
Faurea saligna: Burnt bark used as tonic. Roots are boiled and the decoction drunk
for diarrhea and indigestion, also as tonic (Beentje 1994; Kokwaro 2009). Also used
for stomach problems (Muthee et al. 2011).
Faurea saligna: Timber used for furniture (Beentje 1994). Timber used as firewood
(Bussmann 2006).
References
Brummitt RK. Flora of tropical East Africa: Proteaceae. Rotterdam: Balkema; 1993.
syntaxonomical survey. J East Afr Nat Hist. 2000;89(1-2):45–71.
Ficus sycomorus L.
Ficus thonningii Blume
MORACEAE

Grace N. Njoroge
Synonyms
Ficus sycomorus L.: Ficus cocculifolia Baker; Ficus cocculifolia subsp.

sakalavarum (Baker) H. Perrier & Lenadri; Ficus cocculifolia var. sakalavarum
(Baker) H. Perrier; Ficus comorensis Warb.; Ficus damarensis Engl.; Ficus
exasperata Vahl; Ficus integrifolia Sim; Ficus sakalavarum Baker; Ficus scabra
Sim; Ficus sycomorus subsp. gnaphalocarpa (Miq.) C.C. Berg; Ficus trachyphylla
Fenzl; Sycomorus antiquorum Gasp.; Sycomorus gnaphalocarpa Miq.; Sycomorus
rigida Miq.; Sycomorus trachyphylla Miq.
Ficus thonningii Blume: Ficus burkei Miq.; Ficus petersii Warb.
Local Names
Ficus sycomorus: Kikuyu/Swahili/Kamba/Meru/Taita: Mukuyu; Swahili:

Mkuyu, Chivuzi; Borana: Od; Ilelewa: Makoyo; Kipsigis: Mogoiwet; Maa:
Orng’aboli; Nandi: Sebetwet; Oromo: Odha; Pokot: Mokongwa; Rendile: Santau,
R. W. Bussmann (*)
G. N. Njoroge

Gauden; Samburu: Lngaboli; Somali: Bardah; Tugen: Lokoiwo; Turkana:

Echoke; Ndorobo: Ingalobi; Luganda: Mukunyu (Beentje 1994; Gachati 1989;
Kokwaro 2009); Amharic: Shola, Bamba; Tigrigna: Saghla, Sagla, Shegla;
English: Wild fig, Stranglerfig, Sycamore, Bush fig, Common cluster fig.
Ficus thonningii: Kikuyu/Embu/Meru: Mugumo; Swahili: Mtschamwa,
Mrumbapori; Borana: Dambi; Kamba: Kiumo, Muumo; Kipsigis: Simotwet;
Luo: Pocho; Maa: Oreteti; Okiek: Sapoitit; Lang’o: Ananga; Lugbara: Laro;
Nyankore: Omutoma; Chagga: Mkuu (Beentje 1994; Gachati 1989; Kokwaro
2009); English: Strangler Fig, Wild Fig; Tigrigna: Shibaka.
Botany and Ecology
Ficus sycomorus L.: Tree up to 20(–30) m tall, trunk short, main branches
spreading. Leafy twigs (1–)2–6 mm thick, densely minutely puberulous and with
much longer white to yellowish hairs especially on the nodes, periderm flaking off
when dry. Lamina chartaceous to coriaceous, ovate to elliptic, obovate or sub-
circular, (1–)2.5–12(–21) (0.5–)2–11(–16) cm, apex rounded to obtuse, base
cordate to sometimes obtuse, margin subentire, slightly repand or denticulate,
upper surface scabrous to scabridulous, sometimes almost smooth, hispidulous to
strigillose, on the main veins whitish hirtellous to hirsute, lower surface puberulous
to hispidulous, on the main veins partly whitish hirtellous or hirsute, lateral veins
5–10 pairs, petiole (0.5–)1–4(–6) cm long, 1–3 mm thick, densely minutely white
puberulous and with much longer white to yellowish hairs, with the periderm
flaking off when dry, stipules 0.5–2.5 cm long, white puberulous to tomentose or
partly hirtellous to hirsute, caducous. Figs solitary or sometimes in pairs in the leaf-
axils or just below the leaves, on up to 10 cm long unbranched leafless branchlets or
on up to 20(–35) cm long branched leafless branchlets on the older branches down
to the trunk, peduncle 0.3–2.5 cm long, 1–3 mm thick, basal bracts 2–3 mm long.
Receptacle obovoid to pyriform or subglobose, often stipitate at least when dry,
1.5–5 cm in diameter when fresh, (1–)1.5–3 cm when dry, white to yellowish or
brownish velutinous or densely tomentose to sparsely puberulous or pubescent,
sometimes almost glabrous, yellowish to reddish at maturity. Forest edges, lake-
sides, riverine, extending into drier country especially where seasonal water collects
at foot of hills and scarps, rock outcrops, 0–2200 m (Berg and Hijman 1989)
(Figs. 1, 2, 3, 4, 5, and 6).
Ficus thonningii Blume: Tree up to 15(–30) m tall or a shrub, terrestrial or hemi-
epiphytic. Leafy twigs 1.5–7 mm thick, minutely puberulous to hirtellous or white to
brown pubescent, at least on the scars of the stipules or sometimes entirely glabrous,
periderm usually not flaking off. Leaves in spirals, occasionally subopposite, lamina
coriaceous, elliptic to lanceolate, oblanceolate, obovate or subovate, (1.5–)3–12
(–18) (1–)1.5–6(–7) cm, apex acuminate to obtuse or rounded, base subacute to
rounded or subcordate, margin entire, upper surface glabrous or sparsely (on the
midrib to rather densely) puberulous to pubescent, lower surface glabrous or
sparsely to densely white (to brownish) puberulous to pubescent on the whole
Ficus sycomorus L. . . . 539
Fig. 1 Ficus sp. (Moraceae),

Park, Demaro forest. (Photo
R. W. Bussmann)

R. W. Bussmann)
surface, the main veins or only the midrib, lateral veins (5–)7–12(–16) pairs, midrib
often reaching the apex of the lamina (even in leaves with a rounded apex), tertiary
venation reticulate or parallel to the lateral veins, petiole (0.5–)1–4(–6) cm long,

R. W. Bussmann)

R. W. Bussmann)
1–2 mm thick, often (not depending on the size of the lamina or the position of the
leaf on the twig) variable in length on the same twig, glabrous or puberulous,
hirtellous or pubescent, stipules 0.3–1(–1.5) cm long, white to brown pubescent,
puberulous or only ciliolate, caducous or subpersistent. Figs in pairs in the leaf-axils
or sometimes also below the leaves, sessile or on peduncles up to 1 cm long, basal
bracts 2–4 mm long, persistent. Receptacle globose to ellipsoid, 0.5–1.5(–2) cm in
diameter when fresh, 0.4–1.2(–1.7) cm when dry, glabrous or sparsely to densely
white to brown puberulous or pubescent, reddish, yellowish or brownish at maturity,
wall thin, mostly smooth or slightly wrinkled when dry, apex plane to strongly
protruding when dry. Forest, woodland, bushland and wooded grassland, sometimes
along rivers and lakes or among rocks, planted for ornament and barkcloth,
350–2500 m (Berg and Hijman 1989) (Figs. 6, 7, 8, 9, and 10).

R. W. Bussmann)

R. W. Bussmann)

R. W. Bussmann)
Ficus sycomorus: The bark is used for the treatment of scrofula, coughs, and throat
and chest diseases. The milky latex is used for the treatment of dysentery and chest
diseases or is applied to inflamed areas, while ringworm is treated with the bark and
milky latex. Leaves are said to be effective against jaundice and as an antidote for
snakebite, while the roots have laxative and anthelmintic properties. Bark also used
for general abdominal pain, stomachache, dysentery, diarrhea, sore throat, chest
pain, and glandular pain. Latex is used for toothache (Bekalo et al. 2009; Kokwaro
2009). Employed for uterine problems (Muthee et al. 2011). Used to restore the
uterus after childbirth, to extract the rest of the placenta in the uterus and to promote
the milk rise, to treat cough and colic, for childhood asthma, flu of children
(Randrianarivony et al. 2016a, b), as well as intestinal parasites (Rabearivony
et al. 2015). It is used to improve menstrual flow (Njoroge and Bussmann 2009)
and also used to treat boils and scabies (Njoroge and Bussmann 2007).
Ficus thonningii: The bark is important in local medicine, and it is used in treating
colds, sore throat, dysentery, wounds, constipation, nosebleed, and to stimulate
lactation. Latex is used for wound fever, while an infusion of the root and fiber is

R. W. Bussmann)

R. W. Bussmann)
taken orally to help prevent abortion. Powdered root is taken in porridge to stop
nosebleed, and the milky latex is dropped into the eye to treat cataracts. The bark
serves to treat flu and venereal diseases. The root bark decoction serves as
galactagogue (Wondimu et al. 2007). The latex helps to treat mouth sores (Kokwaro
Fig. 10 Ficus
sp. (Moraceae), Bale
Demaro forest. (Photo R. W.
Bussmann)
2009). Used to treat respiratory disorders (Muthee et al. 2011). Also used to treat
appetite loss and diarrhea (Teklehaymanot et al. 2007) and to improve menstrual
flow (Njoroge and Bussmann 2009).
Ficus botryoides is used for breast infections, diarrhea, and against fatigue in
Madagascar (Randrianarivony et al. 2016a, b). Ficus vasta latex is applied to
external injuries (Giday et al. 2003, 2007, 2009; Teklehaymanot and Giday 2010).
Ficus polita serves for stomachache and urine retention; Ficus polyphlebia for
gonorrhea and wounds; Ficus pyrifolia for cough and wounds; and Ficus politoria
for stomach pain and urinary disorders (Rabearivony et al. 2015).
Local Food Uses
Ficus sycomorus: Mature fruits are eaten fresh, stewed, or dried and stored for later
use (Beentje 1994). Fruit can also be used for the preparation of an alcoholic
beverage. Leaves are used in soups and groundnut dishes. The bark is chewed
together with kola nut. In Ghana, the wood ash is commonly used as a salt substitute.
Fruits are eaten (Bussmann et al. 2011).
Ficus thonningii: Jam can be made from the ripe fruits. Fruits are eaten (Bussmann
et al. 2011).
The fruits of Ficus pachyclada are eaten in Madagascar (Randrianarivony et al.

2016a, b).
Ficus sycomorus: A ceremonial tree for many African cultures (Beentje 1994).
Leaves are a much-sought fodder with fairly high nutritive value in overstocked
semiarid areas where the trees occur naturally. Fruits are eaten by livestock, wild
animals, and birds. It can be used as firewood and for making charcoal, and various
peoples throughout Africa use a piece of dry wood from this tree as the base block
when starting a fire by the friction method. The inner part of the root is used as
weaving fiber, and a strong rope can be made from the inner bark. The wood is
creamy brown, has a fairly uniform structure, is very light, soft to moderately hard,
tough, strong, easy to work, finishes smoothly, and holds nails firmly. It is not very
durable and is easily attacked by termites. Mainly used for making mortars and
pestles, drums, stools, doors, beehives, dugout canoes, carvings, and for house
building (Beentje 1994). Wild fig can be used for sand-dune fixation and riverbank
stabilization. The tree gives useful shade and is common at marketplaces, where
people gather under it for many social functions. Shed leaves form a valuable litter
improving the nutrient status, infiltration rate, and water-holding capacity of the soil.
An important tree planted for ornamental purposes near temples, roadsides, wells,
and community places such as market centers in rural areas. Used as firewood, and to
make beehives (Bussmann et al. 2011). Leaves are used to make bowls (Balemie and
Kebebew 2006). The fruits are used ceremonially to defend against evil spells
Figs are widely seen as sacred trees among many African Mountain communities.
Ficus thonningii: It is a ceremonial tree for many African cultures (Kokwaro 2009).
Livestock eat the dry leaves on the ground and to a lesser degree fresh leaves. Leaves
and twigs are eaten by bushbuck, dikdik, elephant, giraffe, impala, kudu, and nyala.
Dropped fruits are eaten by baboon, bushbuck, bushpig, civet, dikdik, gray duiker,
rock and tree hyrax, impala, kudu, slender mongoose, samango and vervet monkeys,
nyala, porcupine, and warthog. The ripe fruits are eaten by bats, barbets, bulbuls,
louries (turacos), parrots, pigeons, and starlings. Branches are used for firewood.
Bark cloth is obtained by cutting out a strip or cylinder of bark, which causes the tree
to produce a fine, matted covering of red, slender roots over the wound. Bark fiber is
used for making mats, and the twined bark produces a strong rope, which is mostly
used for fastening bundles of firewood before they are carried to the homestead for
fastening slates onto a roof. Branches are used as fire sticks (Beentje 1994). The
wood is creamy brown, has a fairly uniform structure, is light, soft to moderately
hard, with a rough texture, tough, strong, easy to work, finishes smoothly, and holds
nails firmly. Its durability is low, and it is easily attacked by termites. A considerable
amount of useful latex is produced by the tree. The sticky juice from pounded roots is
used to trap small animals like hares and birds. Truncheons can be planted close to
each other to help control erosion. The species is often planted to offer cover from
the sun. It can also be planted to provide shelter during the cold winter months. Leaf
litter helps in the improvement of the nutrient status and water-holding capacity of
the soil. Intercropping: In Uganda, the tree is intercropped with coffee and bananas.
This tree has an aggressive root system and should not be planted in a small garden
or near buildings, swimming pools, or paths. It makes an ideal shade tree in a large
garden or park, and it makes a successful container plant for the patio. It is also ideal
for use as a bonsai specimen. Used as firewood and to make beehives (Bussmann
et al. 2011). It is used for diarrhea and anaplasmosis in cattle (Njoroge and Bussmann
2006) and used also as shade tree (Mekonnen et al. 2015).
Figs are widely seen as sacred trees among many African Mountain communities.
Ficus palmata is used in ethnoveterinary medicine in Ethiopia (Yineger et al.
2007). Ficus botryoides is used ceremonially in Madagascar (Randrianarivony et al.
2016a, b).
Branches of Ficus exasperata are used as insect brush (Tabuti 2008). Ficus has a
particularly wide use in Nepal (Kunwar and Bussmann 2006).
References
doi.org/10.1186/1746-4269-5-26.
Berg CC, Hijman MEE. Flora of tropical East Africa: Moraceae. Rotterdam: Balkema; 1989.
Kunwar RM, Bussmann RW. Ficus species in Nepal: a review of diversity and indigenous uses.
Lyonia. 2006;11(1):85–97.

2009;8(2):262–9.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016a. ISBN 978-0-9960231-4-6
Madagascar. Indian J Tradit Knowl. 2016b;151:68–78.
Flacourtia indica (Burm. f.) Merr.
SALICACEAE

Grace N. Njoroge
Synonyms
Flacourtia indica (Burm. f.) Merr.: Flacourtia parvifolia Merr.; Flacourtia

ramontchi L’Hér.; Gmelina indica Burm. f.
Local Names
Flacourtia indica: Kikuyu: Muroro, Mroro, Mutahachu; Swahili: Mugovigovi;

Digo: Munyandoiya; Durma: Madangatunda; Giriama: Mugereaka; Kamba:
Kiathani; Kipsigit: Tungoroloet; Luhya: Bukarambi; Maa: Ol-leleroi,
Oldongururwo; Marakwet: Tungururwa; Meru: Muraga; Nandi: Lichet Nandi /
Sebei: Tugururiet; Sanya: Mogodonya; Samburu: Loloroi; Turkana: Echoke;
Sukuma: Muluka Mhuli; Pokot: Tingoswa (Beentje 1994; Gachati 1989; Kokwaro
2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Flacourtia indica (Burm. f.) Merr.: Shrubs or small trees, 2–4 m tall, deciduous,
bark gray-yellow, fissured, flaky, old branches usually not spiny, young branches
with axillary, simple spines, branchlets puberulous or subglabrous. Petiole red, short,
3–5 mm, puberulous, leaf blade greenish abaxially, deep green adaxially, rose red
when young, obovate to oblong-obovate, 2–4 1.5–3 cm, thickly papery, abaxially
glabrous or sparsely pubescent, hairs spreading and short, adaxially glabrous,
midvein raised abaxially, flat adaxially, lateral veins 5–7 pairs, reticulate veins
conspicuous, base mostly acute to obtuse, margin serrulate above middle, apex
rounded, sometimes retuse. Inflorescences axillary or terminating short lateral
twigs, racemose, short, rachis 0.5–2 cm, puberulous. Pedicels 3–5 mm, puberulous,
hairs spreading. Sepals 5 or 6, ovate, ca. 1.5 mm, outside glabrous or with a few
scattered short hairs, inside sparsely to densely pubescent, margin white ciliate in
dried material, apex obtuse. Staminate flowers: stamen filaments 2–2.5 mm, pubes-
cent or less often glabrous. Pistillate flowers: ovary globose, placentas 5 or 6, styles
5 or 6, united only at base, radiating, 1–2 mm, slender. Fruit dull to blackish red,
globose, 8–10 mm in diameter, longitudinally 5- or 6-angled, styles persistent. Seeds
5 or 6 (Sleumer 1975). Especially common in Afromontane forests dominated by
Coffea arabica (Bussmann 2002a, b; Bytebier and Bussmann 2000) (Figs. 1 and 2).
Flacourtia indica: Root decoction used as diuretic, for diarrhea, indigestion, gon-
orrhea, snakebite, infertility, and stomach pain. The leaf decoction serves to treat
asthma and anemia. Burnt roots are licked for cough and sore throat (Kokwaro
2009). Used to treat malaria (Flatie et al. 2009; Njoroge and Bussmann 2006). Also
used to treat constipation (Singh et al. 2019). To cure vertigo during pregnancy, to
treat the early closing of the fontanelle, for toothache, and to treat malaria and
Fig. 1 Flacourtiaceae cf.,

Amazonas, Peru. (Photo R. W.
Bussmann)
Flacourtia indica (Burm. f.) Merr. 551
Fig. 2 Flacourtiaceae cf.,

Bussmann)
children’s fever: decoction of the leaves (Randrianarivony et al. 2016a, b).

Flacourtia ramontchi is used for the same purpose (Randrianarivony et al. 2017).
Flacourtia jangomas is used to treat pain (Debbarma et al. 2017).
Local Food Uses
Flacourtia indica: Fruit edible (Beentje 1994; Randrianarivony et al. 2016a, b).
Flacourtia indica: The leaf decoction serves to treat screw worm in cattle (Kokwaro
2009). Wood used to build huts (Maasai) (Beentje 1994).
References
Ambasht NK, editors. Modern trends in applied terrestrial ecology: Kluwer, New York; 2002b.
p. 195–223.
syntaxonomical survey. J East Afr Nat Hist. 2000;89(1&2):45–71.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.

Njoroge GN, Bussmann RW. Phytotherapeutic management of diversity and utilization of antimalar-
ial ethnophytotherapeutic remedies among the Kikuyus Central Kenya. J Ethnobiol Ethnomed.
2006;2:8.
gascar Conservation and Development. 2016a; 11(2): 44–51.
Madagascar. Indian J Tradit Knowl. 2016b;15(1):68–78.
2017. https://doi.org/10.1186/s13002-017-0147-x.
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;18:14.
https://doi.org/10.32859/era.18.14.1-11.
Sleumer HO. Flora of tropical East Africa: Flacourtiaceae. London: Crown Agents for Overseas
Galinsoga parviflora Cav.
ASTERACEAE

Grace N. Njoroge
Synonyms
Galinsoga parviflora Cav.: Adventina parviflora Raf.; Galinsoga parviflora var.

semicalva A. Gray.; Galinsoga quinqueradiata Ruiz & Pav.; Galinsoga semicalva
(A. Gray) H. St. John & D. White; Sabezia microglossa DC.; Stemmatella sodiroi
Hieron; Wiborgia acmella Roth; Wiborgia parviflora (Cav.) Kunth
Local Names
Galinsoga parviflora: Kikuyu: Kang’ei, Mung’ei; Meru: Mnyenye (Gachati 1989;

Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Galinsoga parviflora Cav.: Plants 4–60 cm. Leaf blade 7–110 3–70(–80)
mm. Peduncles 1–40 mm, involucres campanulate, 2.5–5 mm in diameter, phyllaries
persistent, outer paleae persistent with distal inner phyllaries or deciduous, elliptic to
obovate, inner usually persistent, lanceolate to ovate or obovate, 2–3.5 mm, 3-lobed,
lobes to 1/3+ total lengths, acute. Ray florets (3–)5(–8), corollas usually dull white or
pink, lamina 0.5–1.8 0.7–1.5 mm. Disk florets 15–50. Ray achenes 1.5–2.5 mm,
pappus absent or of 5–10 laciniate scales 0.5–1 mm, disk achenes 1.3–2.5 mm,
glabrous or strigose, pappus absent or of 15–20 gray, sometimes white, linear,
fimbriate, obtuse or acute scales 0.5–2 mm. Flowering July–October (Wu et al.
1994–2013) (Figs. 1 and 2).
Fig. 1 Galinsoga parviflora

(Asteraceae). Pusac,
Bussmann and N. Y.
Paniagua-Zambrana)
Fig. 2 Galinsoga parviflora

(Asteraceae). Flowers, Pusac,
Bussmann and N. Y.
Paniagua-Zambrana)
Galinsoga parviflora Cav. 555
Galinsoga parviflora: Stem and leaves are chewed for cold sores (Kokwaro 2009).
The extract of the whole plant is used to treat nettle stings (Bhat et al. 2013). Crushed
leaves are applied to snakebites (Giday et al. 2009). The fruits are crushed and taken
with honey to treat common cold (Yineger et al. 2008).
Galinsoga parviflora: Browsed by wildlife, otherwise no use (Bussmann et al.

2011).
References
in Kedarnath Wildlife Sanctuary of Garhwal Himalaya, India. J Ethnobiol Ethnomed. 2013;9(1).
2008;26:132–53.
Garcinia volkensii Engl.
CLUSIACEAE

Grace N. Njoroge
Local Names
Garcinia volkensii: Kikuyu: Munyawa; Ilelewa: Mchichozi; Kamba: Mulaliondo,

Muthithu; Meru: Mchathi; Oromo: Dariss; Somali: Chan-farod; Taita: Munyanga,
Mungecha; Swahili/Giriama: Mkimwemwe; Marakwet: Magilion (Beentje 1994;
Botany and Ecology
Garcinia volkensii Engl.: Much-branched glabrous evergreen tree or shrub (2–)4–

20 m tall, branches stiff, grooved or winged, flattened, 3–4-angled or rounded, bark
gray-brown, smooth, yielding a milky juice or yellow latex when cut. Sepals
5, reddish, unequal, rounded to triangular-ovate, 1–2.5 mm long and wide, glabrous.
Flowers dioecious, in terminal lax condensed 1-many-flowered cymose inflores-
cences, peduncle 0–4 cm long, bracts scale-like, keeled, pedicels 0–2 mm long. Male
flowers with 5 yellow spongy honeycombed fasciclodes uniting to form a star in the
R. W. Bussmann (*)
G. N. Njoroge

center of the flower, alternating with 5 greenish cream staminal-bundles, the fila-
ments joined for most of their length, each bundle 3.5 mm long, 0.7 mm wide, with
5–9 red, brown or mustard-colored anthers. Petals 5, cream to greenish white,
sometimes tinged with pink, with yellowish linear glands radiating from the base,
fleshy, round to broadly obovate, 4–9 mm long, 3.5–7 mm wide, glabrous. Leaves
opposite or rarely in whorls of 3–4, blades very variable, lanceolate or oblanceolate
to broadly ovate or obovate, (1.2–)4–20 cm long, (0.9–)1.5–8 cm wide, rarely
exceeding 11 4.5 cm, acute to rounded at the apex but apiculate, cuneate to
rounded at the base, mostly dark green above and yellowish or pale green beneath,
mostly distinctly coriaceous, the margin often thickened, venation mostly prominent
on both faces when dry, the surface often rugulose, with translucent and opaque
canals usually visible, petiole 0.3–1.8 cm long, with prominent ligule 2–4 mm long.
Fruit green turning yellow, reddish brown or orange, globose, ovoid or 2–4-lobed,
1–3(–5) cm in diameter, smooth and glabrous. Female flowers with small fasciclodes
alternating with sterile staminal-bundles, ovary green or yellowish, 2–4-locular, stigma
white, peltate, 5-lobed. Seeds red, 1–4, ovoid, l–2.2(–4.5) cm long, 0.7–2(–3) cm wide
and 0.6–1.4 cm thick, often much compressed (Bamps et al. 1978). Common in
lower, Ocotea dominated, Afromontane forests (Bussmann 2002). Other species
found in dry forests (Bytebier and Bussmann 2000).
Garcinia volkensii: Fruit infusion is drunk as emetic (Kokwaro 2009). The fruits of
Garcinia buchananii are eaten to expel ascaris (Giday et al. 2010). A leaf decoction
of Garcinia chapelieri is used in Madagascar to treat cough (Rakotoarivelo et al.
2015). Garcinia pervillei is used for fever, malaria, yellow fever, and bilharzia
(Randrianarivony et al. 2016). Garcinia pauciflora for malaria, wound treatments,
and diarrhea (Randrianarivony et al. 2017).
Local Food Uses
In China, a great variety of Garcinia fruits are eaten, e.g., Garcinia bracteata,
Garcinia cowa, Garcinia esculenta, Garcinia multiflora, Garcinia oblongifolia,
and Garcinia xanthochymus. Many of these widespread species can give local
people substantial productivity, as each adult tree can produce about 60 kg fruits
per year. Garcinia pedunculata and Garcinia yunnanensis are the most uncommon
species and serve for certain small population ethnic groups (formerly for Jingpo
people and now for Wa people) as important wild edible fruits and other uses (Liu
et al. 2016).
Garcinia volkensii Engl. 559
Garcinia volkensii: Hardwood but susceptible to borer attack (Beentje 1994).

Garcinia livingstonii is used as firewood (Bussmann 2006). Garcinia chapelieri is
used in Madagascar as construction timber (Rakotoarivelo et al. 2013). Garcinia
paucinervis is found only in rather dry sparse or dense forests in limestone moun-
tains in China, and its only use is for its prestigious timber (Liu et al. 2016).
References
Bamps P, Robson N, Verdcourt B. Flora of tropical East Africa: Guttiferae. London: Crown Agents
for Overseas Governments & Administrations; 1978.
Syst Geogr Plants. 2002;71(2):959–74.
Liu B, Zhang X, Bussmann RW, Hart RE, Li P, Bai Y, Long C. Garcinia in Southern China:
Ethnobotany, management, and niche modeling. Econ Bot. 2016;70(34):416–30.
L. Brown Center, MBG; 2013. ISBN 978-0-9848415-7-8
vol. 1. St. Louis: William L. Brown Center, MBG; 2016. ISBN 978-0-9960231-4-6
2017; https://doi.org/10.1186/s13002-017-0147-x.
Gnidia glauca (Fresen.) Gilg.
THYMELAEACEAE

Grace N. Njoroge
Synonyms
Gnidia glauca (Fresen.) Gilg.: Gnidia eriocephala Meisn.; Gnidia rivae Gilg;
Gnidia volkensii Gilg; Lasiosiphon eriocephalus (Wall. ex J. Graham) Decne.;
Lasiosiphon eriocephalus var. zeylanicus Meisn.; Lasiosiphon glaucus Fresen.;
Lasiosiphon rivae (Gilg) H. Pearson
Local Names
Gnidia glauca: Kikuyu: Muchingiri, Mirrisaa: Ol-keleliet, Mirrisaa: Kiris, Mirrisaa,

Mirrisaa: Kapkarasiet, Mirrisaa: Sengewa, Mirrisaa: Miondurmi (Beentje 1994;
Gachati 1989).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Gnidia glauca (Fresen.) Gilg.: Large much branched shrub 1–6 m high or small tree
up to 9 m or higher. Branches densely leafy in the upper part, branchlets finely
pubescent when young, later glabrescent, bark gray, brown or blackish, smooth to
rugose. Leaves alternate, subsessile or with petiole 1–3 mm long, lamina 32–65
(84) 10–26 mm, narrowly elliptic to narrowly or broadly obovate, acuminate to
acute, rarely obtuse at the apex, gradually narrowed to the base, rigid, glaucous,
glabrous or finely pubescent, glabrescent, midrib prominent on the lower surface.
Inflorescences dense terminal heads 20–50 mm in diameter, 20–50-flowered, pedun-
cle pubescent. Bracts 6–12, cream-colored or salmon-pink, 10–15 4–10 mm,
elliptic to ovate, slightly coriaceous, softly tomentose on both sides, persistent.
Flowers 5(rarely 4)-merous, orange or golden-yellow, fading to brown, pedicels
1–2.5 mm long, tomentose. Calyx tube 9–14 mm long with no articulation, the lower
part with dense tufts of 2–4 mm long silky hairs, upper part softly tomentose, lobes
2.5–4 1.5–2 mm, ovate, rounded, entire or lobed, tomentose outside. Petals
1–2 mm long, elliptic or spathulate, entire, emarginate or lobed, membranous or
fleshy. Stamens in upper row slightly exserted, anthers 1–2 mm long. Ovary pubes-
cent especially at the apex, style 6–8 mm long, stigma globose. Seed
3–4 1.5–2 mm. Growing in edges of mountain forests, in wooded grassland,
also around stream beds, 1000–2350 m (Peterson 1978). Sometimes dominant in the
understory of dry Afromontane forests (Bytebier and Bussmann 2000; Bussmann
2002a, b, c, 2004, 2006a, b) (Figs. 1, 2, and 3).
Fig. 1 Gnidia glauca (Thymelaeaceae), young plants, Bale Mountains National Park, Odo Bulu
area, Ethiopia. (Photo R. W. Bussmann)
Gnidia glauca (Fresen.) Gilg. 563
Fig. 2 Gnidia glauca

(Thymelaeaceae), flowers,
Park, Odo Bulu area, Ethiopia.
Fig. 3 Gnidia glauca

(Thymelaeaceae), Bark cut,
Park, Odo Bulu area, Ethiopia.
Gnidia glauca: The root powder is ingested to treat rabies (Bekalo et al. 2009;
Teklehaymanot and Giday 2007). The bark extract is used as arrow poison
Gnidia stenophylla is used medicinally in Ethiopia (Lulekal et al. 2008). Leaves

of Gnidia danguyana serve against parasites and to stop bleeding in Madagascar
(Razafindraibe et al. 2013).
Gnidia glauca: Bark fibers are used to make ropes (Beentje 1994). Used as fire-
wood. The bark can be turned into ropes (Bussmann et al. 2011). The root extract of
Gnidia kraussiana is used as fish poison (Neuwinger 2004).
References
doi.org/10.1186/1746-4269-5-26.
Syst Geogr Plants. 2002a;71(2):959–74.
Ambasht NK, editors. Modern trends in applied terrestrial ecology. New York: Kluwer; 2002b.
p. 195–223.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist. 2002c;91(1/2):27–79,
Bussmann RW. Regeneration and succession patterns in African, Andean and Pacific tro-
pical mountain forests: the role of natural and anthropogenic disturbance. Lyonia.
2004;61:98–111.
2006b;11(1):41–66.
10.1186/1746-4269-4-10.
Peterson B. Flora of tropical East Africa: Thymelaeaceae. London: Crown Agents for Overseas
Gnidia glauca (Fresen.) Gilg. 565

2013;9:73.
Teklehaymanot T, Giday M. Ethnobotanical study of medicinal plants used by people in Zegie
Peninsula, Northwestern Ethiopia. J Ethnobiol Ethnomed. 2007;3:12. https://doi.org/10.1186/
1746-4269-3-12.
Grewia bicolor Juss.
Grewia tembensis Fresen.
Grewia tenax (Forssk.) Fiori
Grewia villosa Willd.
MALVACEAE

Grace N. Njoroge
Synonyms
Grewia bicolor Juss.: Brewia baillonii R. Vig.; Grewia damine Gaertn.; Grewia
distica Dinter & Burret; Grewia kwebensis N.E. Br.; Grewia miniata Mast. ex Hiern
Grewia tembensis Fresen.: Grewia membranacea A. Rich.; Grewia parviflora
(Hoschst.) A. Rich.
Grewia tenax (Forssk.) Fiori: Chadara texax Forssk.; Grewia picta Baill. var.
picta; Grewia populifera Vahl
Grewia villosa Willd.: Grewia corylifolia A. Rich.; Grewia echinulata Delile;
Grewia orbiculata Rottler
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Grewia bicolor: Swahili: Mkone, Mfukufuku; Sukuma: Mkoma; Borana: Arobera,

Harowessa; Kamba: Mulawa; Kipsigis: Setetit; Luo: Powo; Maa/Samburu:
Olsitete, Siteti; Oromo: Haroru; Pokot: Sitet; Somali: Tebi, Debhi; Luo: Powo;
Turkana: Ekali (Beentje 1994; Kokwaro 2009).
Grewia tembensis: Borana: Deka; Kamba: Mutuva; Kipsigis: Chesarebut; Maa:
Oyiri; Oromo: Deka-dubrta; Pokot: Toronwo; Somali: Dumeg; Samburu: Irii;
Somali: Murie-bonati; Taita: Mwebma; Turkana: Emalaker (Beentje 1994).
Grewia tenax: Bolana/Ilelewa: Deka; Gabbra: Deekaa; Njemps: Ilkongoni;
Oromo: Deka; Pokot: Toronwo; Rendile: Muluhanyo; Samburu: Iri, Laripai;
Somali: Damak, Dekha; Turkana: Eng’omo (Beentje 1994).
Grewia villosa: Bajun: Mukorobasha; Borana: Morudo, Ogumdi; Gabbra/
Oromo: Ogomdi; Kamba: Mulawa; Luo: Nre-Powo; Karamoja: Eponge;
Sukuma: Msarasi; Maa: Olmankulai; Marakwet: Mongurwa; Pokot/Tugen:
Toronwo, Mukowo; Rendile: Obepo; Samburu/Njemps: Lpupoi, Lpusan; Somali:
Kamasha, Kommasha; Taita: Mshoshote; Turkana: Engomo, Epongai (Beentje
1994; Kokwaro 2009).
Botany and Ecology
Grewia bicolor Juss.: Shrub or small tree to 6 m tall, young branches grayish to
brownish tomentose, rarely with slightly coarser stellate pubescence. Leaves usually
elliptic, rarely oblong or ovate, 0.9–8.2 cm long, 0.4–3.5 cm wide, rounded to bluntly
acute at the apex, rounded at the base, margins minutely serrate to entire, green and
glabrous to densely minutely pubescent above, evenly whitish tomentose beneath,
rarely slightly brownish on the veins, petiole 2–4 mm long, grayish to brownish
tomentose, stipules linear-lanceolate, 4–12 mm long. Inflorescence a (1–)3-flowered
cyme, 1–3 in a leaf-axil, the axes grayish to brownish tomentose, peduncle 4–13 mm
long, pedicels 5–14 mm long, bracts ovate to lanceolate, 2–5 mm long. Flowers yellow
to orange, sepals 6–11 mm long, petals oblong, 3–8 mm long, acute to emarginate at
the apex. Androgynophore absent or up to 1 mm long, glabrous. Stamens 6–7 mm
long. Ovary 1.5 mm long, densely hairy, style 5–6 mm long. Fruit usually 2-lobed,
or unlobed by abortion, the lobes 4–7 mm long, 4–8 mm wide, glabrous or with a
few scattered minute stellate hairs, green turning orange when ripe. Woodland, thicket,
Commiphora-Acacia bushland, scattered bush grassland, on dry sandy soil and along
streams, 650–1650 m (Whitehouse et al. 2001). Found in drier Afromonatne forests,
especially in the forest – savannah transition zone and in disturbed areas (Bussmann
2002a, b) (Figs. 1 and 2).
Grewia tembensis Fresen.: Shrub, 0.5–4 m tall. Leaves obovate-elliptic, sometimes
almost round, 0.8–4.5( 8) 0.5–2( 4) cm. Leaf base obtuse or rounded, some-
times cuneate, apex rounded or obtuse, margins serrate. Upper side sometimes
sandpaper like, sparesly hairy. Underside tomentose. Flowers white, sometimes
pink or pale lilac, sepals brown purple outside, in 1-many flowered, axillary
Grewia bicolor Juss. . . . 569
Fig. 1 Grewia bicolor

(Malvaceae), Mt. Kenya
National Park, Naro Moru,
Kenya. (Photo
R.W. Bussmann)
Fig. 2 Grewia bicolor

Kenya. (Photo
R.W. Bussmann)
cymes, sepals 7–12 mm long, petals 4–8 mm long. Fruits orange, 1–4 lobed, with
lobes 4–7 mm long (Whitehouse et al. 2001). Found in drier Afromonatne forests,
especially in the forest – savannah transition zone and in disturbed areas (Bussmann
2002a, b) (Figs. 3 and 4).
Grewia tenax (Forssk.) Fiori: Small shrub up to 2 m tall, branches glabrescent.

Leaf-lamina up to 3 2 cm, orbicular to obovate, rounded at the apex, margin rather
coarsely dentate, rounded or abruptly cuneate at the base, slightly scabrous-
pubescent especially below, or glabrescent, grayish-green, coriaceous, petiole up
to 1 cm long, but usually much less, pubescent, stipules up to 4 mm long, filiform,
pubescent. Flowers always borne singly, leaf-opposed, peduncles 10 mm long,
almost glabrous, slender, pedicels similar, up to 10 mm long, the whole giving the
appearance of an articulated pedicel, bracts minute, glabrescent, 1.5 mm long. Sepals
10–18 mm long, greenish and shortly pubescent outside, white and glabrous inside,
linear-oblong. Petals white, with a linear and often 2-dentate lamina almost as long
as the sepals and narrower than the basal nectariferous claw which is circumvillous
within ledged above and up to 1.5 mm long. Androgynophore with a basal glabrous
portion up to 1.5 mm long and a densely pubescent upper portion up to 1.5 mm long.
Ovary 4-lobed, glabrous or glabrescent, style 10 mm long, glabrous, stigma-lobes
broad. Fruit 10 mm in diameter, 4-lobed, shining, glabrous (Whitehouse et al. 2001).
Found in drier Afromonatne forests, especially in the forest – savannah transition
zone and in disturbed areas (Bussmann 2002a, b) (Figs. 5 and 6).
Fig. 3 Grewia cf. similis

Kenya. (Photo
R.W. Bussmann)
Fig. 4 Grewia cf. tembensis

Kenya. (Photo
R.W. Bussmann)
Grewia villosa Willd.: Shrub 1.2–4.5 m high, plants apparently either bisexual or
unisexual, stems flattened, young shoots densely covered with villous hairs. Leaves
broadly ovate to orbicular, 2–15 cm long, 1.6–18.5 cm wide, rounded to obtuse or
slightly apiculate at the apex, rounded to cordate at the base, margin serrate,
pubescent above, more so on the veins, densely stellate-pubescent beneath with
long simple hairs on the veins, veins prominent and finely reticulate, leaves
appearing bullate when young, petiole 0.6–5.5 cm long, densely villous, stipules
broadly ovate to ovate-lanceolate, 5–10 mm long, caducous. Inflorescence a 2–10-
flowered cyme. Flowers dull red, yellow or brownish, sepals 5–10 mm long, densely
hairy on the outside, usually reddish on the inside, petals much smaller than sepals,
1–3 mm long, usually pinkish. Androgynophore glabrous. Stamens up to 3 mm long,
filaments whitish, anthers yellow. Ovary densely villous, style 1.5–3 mm long,
stigma much divided. Fruit unlobed, subglobose, flattened vertically, 1–1.6 cm in
diameter, orange to rusty-brown, covered with small warts tipped by long hairs,
outer wall crustaceous and well-separated from the pulp. Dry Acacia-Commiphora
bushland, rocky hillsides and beside dry watercourses, 0–1550 m (Whitehouse et al.
2001). Found in drier Afromonatne forests, especially in the forest – savannah
transition zone and in disturbed areas (Bussmann 2002a, b).
Fig. 5 Grewia cf. tenax

Kenya. (Photo
R.W. Bussmann)
Fig. 6 Grewia
sp. (Malvaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
Grewia bicolor: The species has a wide range of applications in African traditional
medicine. The bark is used as a vermifuge, diuretic, and laxative, and to treat boils
and sores, intestinal inflammation, and syphilis. In Senegal, a macerate or decoction
of the bark is credited with both inebriating and tranquillizing effects and is also
taken to counter fatigue. In Kenya, the bark is applied in case of itching, while in
Tanzania, it is chewed and put on wounds as a bandage. The wood is credited with
anthelmintic activity. In Côte d’Ivoire, a decoction of the leaves is drunk and put into
baths against pain in the chest, after which sap from pounded roots is rubbed on the
patient. In East Africa, a cold infusion of the root is drunk to treat anemia, chest
complaints, cold, diarrhea, snakebites, mental illness, hernia, and female infertility.
The bark paste is applied to skin itches (Kokwaro 2009). In Sudan, a poultice of the
root is applied on pustulent skin lesions, and the root is taken as a tranquillizer. A
decoction of the root is given in case of a delayed afterbirth. In Niger, the powdered
root bark is applied on burns, and in Mali, the juice or a decoction of the inner bark of
the roots is applied on wounds. In Namibia, a syrup prepared from the roots is rubbed
onto swollen legs. The plant is also used in veterinary medicine, e.g., to treat
stomach problems. Used as toothbrush (Bussmann et al. 2006). In Ethiopia, it is
used for venereal diseases including syphilis (Teklehaymanot and Giday 2010).
Grewia similis: Used as toothbrush (Bussmann et al. 2006).
Grewia tembensis: A root decoction is used to cure cough (Turkana) (Beentje 1994).
Used as toothbrush (Bussmann et al. 2006).
Grewia tenax: The bark gum is used as insect repellent. Roots are mixed with roots
of Diospyros scabra and the decoction drunk for tuberculosis (Kokwaro 2009).
Grewia villosa: A leaf and twig decoction is used for rheumatism (Borana), and the
bark infusion is used against diarrhea (Samburu) (Beentje 1994). The roots are used
for stomachache and lower abdominal pain. Leaves are applied to earache and spleen
pain. The roots are boiled to treat diarrhea and aching bones (Kokwaro 2009). Used
for boils and fever (Teklehaymanot and Giday 2010).
Grewia ferruginea is used as anthelmintic (Assefa et al. 2010; Desta 1995) and to
treat gonorrhea (Wondimu et al. 2007). Grewia mollis serves to treat constipation
(Bekalo et al. 2009) and for wounds (Flatie et al. 2009). Grewia disperma serves to
stop bleeding and to treat bronchitis (Kunwar et al. 2010). Grewia kakothamnos is
used for chest pain, tuberculosis, coughing with blood, chest pain, and rheumatism
(Teklehaymanot and Giday 2010). Grewia lavanalensis is used in Madagascar
during pregnancy (Randrianarivony et al. 2016a).
Local Food Uses
Grewia bicolor: The sweet, mealy fruit pulp is eaten fresh, or dried as candy
(Beentje 1994). Juice from the fruit is drunk fresh, added to porridge, fermented
into beer, or distilled into liquor. The mucilaginous leaves and fibers from the leaf are
used as binding agent in sauces. Fresh leaves are made into a kind of tea. In Burkina
Faso, the bark or leaf fibers are used in the preparation of sorghum beer to make it
clean and to remove bitterness. The fruit can be eaten (Bussmann et al. 2006).
Grewia similis: The fruit can be eaten (Bussmann et al. 2006).
Grewia tembensis: The fruit is edible (Beentje 1994; Bussmann et al. 2006).
Grewia tenax: The fruit is edible (Beentje 1994).
Grewia villosa: The fruit is edible (Beentje 1994).
Grewia bicolor: The bark is used to clarify muddy water. The bark fiber is made into
cordage and used for weaving. The fresh and dry leaves, young stems, and fruits
serve as forage for domestic animals. The leaves and ash from burnt leaves are
sometimes used as soap and for cleaning garments. The tree is also used as an
ornamental, as a shade tree, and as bee forage. The wood is used in house construc-
tion (poles, beams) and made into a range of articles including tool handles, herding
staffs and walking sticks, bows, arrows, spear shafts, knobkerries and clubs, pegs,
rakes, and saddle frames. It is used for carving, and in Kordofan (Sudan), the wood is
hollowed out to make bowls and boards for the “kalah” game, and the stems are
made into picture frames. In Burkina Faso, sticks are woven into baskets. The wood
is also used for fire sticks, as fuel wood, and made into charcoal. Also excellent for
spears, bows, arrows, and rungus (clubs) (Beentje 1994). The root decoction is used
to expel the placenta in cattle (Kokwaro 2009). Eaten by goats and cows but must not
be cut as fodder. Bark used to join sticks (Bussmann et al. 2006). Serves as firewood
Grewia similis: Makes good sticks (Bussmann et al. 2006). Serves as firewood
Grewia tembensis: Eaten by goats and cows but must not be cut as fodder. Bark used
to join sticks (Bussmann et al. 2006). Serves as firewood (Bussmann et al. 2011).
Grewia tenax: The wood is also used for fire sticks, as fuel wood, and made into
charcoal. Also excellent for spears, bows, arrows, and rungus (clubs) (Beentje 1994).
Serves as firewood (Bussmann et al. 2011).
Grewia villosa: Wood used for construction (Beentje 1994). Serves as firewood
(Bussmann et al. 2011; Tian 2017).
Grewia glandulosa is used in Madagascar for construction and tools
(Randrianarivony et al. 2016b).
References
Assefa B, Glatzel G, Buchmann C. Ethnomedicinal uses of Hagenia abyssinica (Bruce) J.F.Gmel.
among rural communities of Ethiopia. J Ethnobiol Ethnomed. 2010;6:20. https://doi.org/
10.1186/1746-4269-6-20.
org/10.1186/1746-4269-5-26.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in far-west Nepal: a
Madagascar. Indian J Tradit Knowl. 2016a;151:68–78.
10.1186/s13002-016-0131-x.
Whitehouse C, Cheek M, Andrews S, Verdcourt B. Flora of tropical East Africa: Tiliaceae &
Muntingiaceae. Rotterdam: Balkema; 2001.
Hagenia abyssinica J.F. Gmel.
ROSACEAE

Grace N. Njoroge
Synonyms
Hagenia abyssinica J.F. Gmel.: Banksia abyssinica Bruce; Brayera anthelmintica

Kunth ex A. Rich.; Hagenia anthelmintica (Kunth ex A. Rich.) J.F. Gmel. ex
Eggeling
Local Names
Hagenia abyssinica: Kikuyu: Munyieri, Kamonde, Muhooru, Mumondo,

Muthithiku; Swahili: Mturunga, Mdobore; Chagga: Mwaanga; Meru: Mujagajoga;
Kipsigis: Bondet; Marakwet: Seweruwa; Kisii: Omukunakuna; Maa: Olboldo;
Meru: Mujogajoga; Nandi: Mjojoruet; Pokot: Sorwa; Sebei: Sokoruet (Beentje
1994; Gachati 1989; Kokwaro 2009); English: Kosso
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Hagenia abyssinica J.F. Gmel.: Tree up to 18 m high, with globose or umbrella-

shaped open crown, bark ridged and flaky, red-brown to brown. Branchlets densely
sericeous-villous with golden antrorse hairs 3–4 mm long, ringed by the scars of the
fallen sheathing leaf-bases, scars at first villous, later glabrescent. Leaves
imparipinnate, up to 40 cm long, viscid, leaflets 5–7(8) on each side, usually
12–15 3.5–5.5 cm, subopposite, sessile or almost so, narrowly oblong, acuminate
at apex, obliquely rounded to subcordate at the base, glabrescent above, densely
silvery sericeous-villous to glabrescent below, margins finely dentate, main leaflet
pairs alternating with 1(3) reduced subcircular ones, the latter with up to 5 teeth or
minute and entire, rachis and petiole with soft patent hairs up to 4 mm long, petiole
up to 15 cm long, stipules (3)6–9 cm long, membranous, reddish at first but
becoming brown, adnate to petiole throughout almost their whole length. Panicles
many-flowered, much-branched, terminal, drooping, up to 60 cm long, female
panicles bulkier than the male ones, flowers subtended by 2(3) broadly rounded
bracts. Male flowers orange-buff to white, 8 mm in diameter, calyx-tube obconical,
1.5 mm long, densely hairy outside, epicalyx-lobes 1.5 0.5 mm, oblong, calyx-
lobes 4–5 2·5–3 mm, oblong to obovate, recurved and with the abaxial surface
concave, clearly veined, petals 1.5 mm long, linear, fugacious or absent, stamens (8)
10–15(20), anthers sparsely hairy, carpels vestigial, enclosed by the receptacle.
Female and hermaphrodite flowers red, viscid, up to 1.5 cm in diameter, calyx-
tube 1 mm long, obconical, densely hairy outside, epicalyx-lobes 1 0.4 cm,
unequal, oblong-elliptic to obovate with apex obtuse, clearly veined, accrescent in
fruit, calyx-lobes 3 2 mm, broadly ovate-acute with the apex acute, clearly veined,
stamens 0, or if present then fewer and smaller than in male flowers, carpels (1)2,
apically villous, with styles 1.5 mm long, stigmas 0.75 mm wide, hairy. Male flowers
orange-buff to white, 8 mm in diameter, calyx-tube obconical, 4–5 2.5–3 mm,
oblong to obovate, recurved and with the abaxial surface concave, clearly veined,
petals 1·5 mm long, linear, fugacious, stamens (8)10–15(20), anthers sparsely hairy,
carpels vestigial, enclosed by the receptacle. Female and hermaphrodite flowers red,
viscid, up to 1.5 cm in diameter, calyx-tube c. 1 mm long, obconical, densely hairy
outside, epicalyx-lobes c. 1 0.4 cm, unequal, oblong-elliptic to obovate with apex
obtuse, clearly veined, accrescent in fruit, calyx-lobes c. 3 2 mm, broadly ovate-
acute with the apex acute, clearly veined, stamens 0, or if present then fewer and
smaller than in male flowers, carpels (1)2, apically villous, with styles 1.5 mm long,
stigmas 0.75 mm wide, hairy. Achenes enclosed within the calyx-tube with its
accrescent epicalyx and persistent calyx-lobes, often only one of the achenes devel-
oping, pericarp thin, fragile, brown, reticulately rugose. Hagenia abyssinica is
indigenous to montane regions of eastern, central, and southern Africa, mostly
above 2000 m altitude. It is found in the Democratic Republic of Congo, Sudan,
and Ethiopia, and south to Malawi, Zambia, and Zimbabwe (Graham 1960) (Figs. 1,
2, 3, 4, 5, 6, 7, and 8).
Hagenia abyssinica dominated forests form the timberline in many African
mountain systems (Bussmann 2002a, b, c, 2004; Lange et al. 1997), and in plateau
Hagenia abyssinica J.F. Gmel. 579
Fig. 1 Hagenia abyssinica (Rosaceae), large mature Hagenia abyssinica tree of about 1200 years,
Mt. Kenya National Park, Naro Moru, Kenya. (Photo R.W. Bussmann)
and drier areas are heavily affected by human activities (Bussmann 1996). In
Ethiopia, for example, a large part of the areas above 2800 m was originally covered
by this forest type and is nowadays mostly converted into agricultural land (Figs. 1,
2, 3, 4, 5,6, 7, 8, 9, 10, 11, 12, 13, and 14).
The species is relatively fast growing (Bussmann 1999) but only germinates well
after the seeds get a heat surplus, e.g., by fire. The seeds cannot be stored well
(Bussmann and Lange 2000).
At first sight, it seems fairly incomprehensible that fire should be the decisive
factor for the regeneration of the Hagenia forests, which are predominately located
in the misty regions of the East African mountains, as high precipitation, as well as
frequent cloud formations, might prevent the vegetation and dead plant material
from becoming sufficiently dry to burn. Martin (1982), however, mentioned that
fires tend to be most frequent in areas with a combination of comparatively high rates
of primary production, slow decomposition rates, and occasional drying of the fuels.
Even Houston (1995) stated that the alternation of wet and dry seasons produces
ideal conditions for fires in wet regions. On Mt. Kenya, for example, there are
rainfall anomalies in the form of severe droughts, as, for example, the one that
occurred in 1890. In recent times, the rainfall of the year 1990 was likewise
extremely low, leading to the assumption that extraordinary dry periods, favoring
the outbreak of fires, occur in approximately 100 year interval. Thus, in sufficiently
wet years, a relatively high plant productivity, together with the large litter produc-
tion of an adult Hagenia, which is shedding the older leaves continually (Miehe and
Miehe 1994) and the low decomposition rates in the cold climate of the subalpine
Fig. 2 Hagenia abyssinica

(Rosaceae), large mature
Hagenia abyssinica tree of
about 1200 years with burn
scars, Mt. Kenya National
Park, Sirimon, Kenya. (Photo
R.W. Bussmann)
forests, result in the accumulation of fuel, which seem to encourage the spreading of
fires in extremely dry years. The existence of charcoal layers in the soil profiles, dug
within the Hagenia forests, indicates that fires are really a characteristic feature of
these misty forests. Field observations yielded, that regeneration stages of Hagenia
abyssinica are always growing together with either young seedlings of Juniperus
procera and/or fire indicating vegetation. Further, the equal ages of the Hagenia
forest stands suggest that the Kosso is not regenerating successively but all of a
sudden after a disturbing event like fire. Germination tests yielded that the Hagenia
seeds were not germinating unless the dense undergrowth was removed. Fire is the
only natural factor that is able to destroy the undergrowth of larger areas and thus to
provide an environment with reduced competition, suitable for germination of the
seeds. Germination therefore depends on a fire event, which is removing the
competing undergrowth and further stimulates the germination of the Kosso with a
sudden heat surplus (Bussmann and Lange 2000).
The regeneration cycle of the Hagenia forests of Mt. Kenya (Lange et al. 1997;
Bussmann and Lange 1999) and other areas is shown in Fig. 15.
Stage 1 of the cycle describes the climax forests with adult Kosso trees of uniform
size and age, and Juniperus procera as codominant tree species. The canopy of these
forests is often closed to nearly 100%, and the undergrowth mainly consists of a
dense herb and grass layer, whereas higher bushes are rarely found. The conditions
of high competition and intensive shading of the ground are not suitable for the
Hagenia abyssinica J.F. Gmel.
Fig. 3 Hagenia abyssinica (Rosaceae), large male tree flowering, Mt. Kenya National Park, Naro Moru, Kenya. (Photo R.W. Bussmann)
581

(Rosaceae), male tree
flowering, Bale Mountains
R.W. Bussmann)

(Rosaceae), young female
flowers, Bale Mountains
R.W. Bussmann)
germination of Hagenia abyssinica. A fire event disturbs the climax community by

destroying herbs and leaf litter in the undergrowth, as well as all roots in the upper
soil layers, whereas the adult Kosso trees remain more or less severely damaged but
still alive in most cases. Miehe and Miehe (1994) supposed adult Hagenia trees to be
almost fire resistant, as their bark is peeling in large pieces, preventing a fire from

(Rosaceae), mature male
flowers, Bale Mountains
R.W. Bussmann)
reaching the essential vascular strands of the tree. Supporting this assumption, many
Kosso trees in the forests show sometimes heavy signs of former burning, but they
are still green and even flowering. After a fire, the heated ground, now bare of
vegetation and only covered with black ash, provides ideal conditions for the
germination of the Kosso seeds. It depends on the frequency of fires, whether
stage 3 or 4 of the regeneration cycle is developing in a certain area. In case the
time intervals between two fire events are sufficiently high (approximately
100 years), the young Hagenia trees are found scattered between a dense pioneer
vegetation of about 5–6 m height. Clutia abyssinica (Euphorbiaceae), Dombeya
torrida (Malvaceae), Helichrysum schimperi (Asteraceae), and Leonotis mollissima
(Lamiaceae) are characteristic species in the impenetrable shrub thicket of this
succession stage after fire. In contrast, more frequent burning results in a shift to a
less high and dense vegetation dominated by Gnidia glauca (Thymelaeaceae).
Hagenia abyssinica: An infusion made using dried and pounded female flowers or
bark is widely used against tapeworms and as general anthelminthic, and has been
reported very consistently in literature (Assefa et al. 2010; Beentje 1994; Bussmann
et al. 2011; Desta 1995; Fassil 2005; Gedif and Hahn 2003; Giday et al. 2003; Luizza

(Rosaceae), male flowers,
Park, Odo Bulu forest,
Ethiopia. (Photo
R.W. Bussmann)
et al. 2013; Mekonnen et al. 2015; Teklehaymanot and Giday 2007; Wondimu et al.
2007). This treatment has been used for centuries especially in Ethiopia, but its use is
now in decline due to availability of reliable alternatives. Health organizations
discourage the use of this infusion as the dosage cannot be controlled, and serious
side effects of overdosage have been reported. Honey, obtained from beehives
located near Hagenia abyssinica trees and collected immediately after their
flowering, is also said to be effective in expelling tapeworms. Roots of Hagenia
are cooked with meat to produce a soup that is consumed as a treatment for general
illness and malaria. The bark is used in treatments for diarrhea and stomachache
(Kokwaro 2009). It is also used, often in a mixture with parts of other plants, as a
medicine to treat syphilis, scrofula, malaria, fever, and cough.

(Rosaceae), male flowers,
Ethiopia. (Photo
R.W. Bussmann)
Local Food Uses
Hagenia abyssinica: The seeds are used as a condiment or (Bussmann 2006).
Hagenia abyssinica: The dark red wood is used for furniture, flooring and carving,
and carpentry (Beentje 1994) but is not durable. An attractive tree, Hagenia is
sometimes planted as an ornamental. The wood is also used as fuel and the leaves
as fodder and green manure. In the Kilimanjaro area, the bark is used for dyeing

(Rosaceae), regenerating
Hagenia under burnt old trees,
Marania, Kenya. (Photo
R.W. Bussmann)

(Rosaceae), young specimens
of Hagenia abyssinica in
burnt area, Mt. Kenya
National Park, Marania,
Kenya. (Photo
R.W. Bussmann)
textiles yellowish red. Used to prevent evil eye affecting livestock (Assefa et al.
2010) and have other veterinary applications (Yineger and Yewhalaw 2007). The
leaves are eaten by cattle (Bussmann et al. 2011).
Fig. 11 Hagenia abyssinica (Rosaceae), male tree as remnant in burnt area, Mt. Kenya National
Park, Marania, Kenya. (Photo R.W. Bussmann)
Fig. 12 Hagenia abyssinica (Rosaceae), Hagenia destroyed by agriculture, Mt. Kenya National
Park, Marania, Kenya. (Photo R.W. Bussmann)
Fig. 13 Hagenia abyssinica (Rosaceae), overaged forest, Mt. Kenya National Park, Naro Moru
route, Kenya. (Photo R.W. Bussmann)

(Rosaceae), Hagenia
abyssinica with beehives,
Ethiopia. (Photo
R.W. Bussmann)
Fig. 15 Hagenia abyssinica (Rosaceae), regeneration cycle (Lange et al. 1997; Bussmann 2002b)
References
10.1186/1746-4269-6-20.
Bussmann RW. Growth rates of important east African montane forest trees, with particular
reference to those of Mount Kenya. J East Afr Nat Hist. 1999;881-2:69–78.
p. 195–223.
BITÖK, editor. Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie; 1999;64. p. 1–16.
Nat Hist. 2000;89(1–2):101–11.
Fassil H. Beyond plants, professionals & parchments: the role of home-based medicinal plant use
and traditional health knowledge in primary health care in Ethiopia. Ethnobot Res Appl.
2005;3:037–49.
Gedif T, Hahn H-J. The use of medicinal plants in self-care in rural central Ethiopia.
Graham RA. Flora of tropical East Africa: Rosaceae. London: Crown Agents for Overseas
Houston MA. Biological diversity – the coexistance of species on changing landscapes. Cambridge:
Cambridge University Press; 1995.
Lange SH, Bussmann RW, Beck E. Ecology and regeneration of the subalpine Hagenia abyssinica
forests of Mt. Kenya. Botanica Acta. 1997;1106:473–80.
Res Appl. 2013;11:315–39.
Martin RE. Fire history and its role in succession. In: Means JE, editor. Forest succession and stand
development in the northwest. Corvalis: Forest Res. Lab., Oregon State University; 1982.
Miehe S, Miehe G. Ericaceous forests and heathlands in the Bale Mountains of South Ethiopia. In:
Ecology and man’s impact. Hamburg: Stiftung Walderhaltung in Afrika und Bundesforschung-
sanstalt für Forst- und Holzwirtschaft; 1994.
1746-4269-3-12.
Harungana madagascariensis Poir.
HYPERICACEAE

Grace N. Njoroge
Synonyms
Harungana madagascariensis Poir.: Haronga madagascariensis (Lam. ex Poir.)

Choisy; Haronga paniculata fo. oblongifolia Engl.; Haronga paniculata
fo. oblongifolia Engl.; Harongana paniculata (Pers.) Lodd. ex Steud.; Harungana
paniculata Pers.; Harungana robynsii Spirlet
Local Names
Harungana madagascariensis: Kikuyu: Muithatua; Luo: Aremo; Shambaa:

Mkuntu, Mnyampome; Meru: Munyamwe; Nyankore: Muitaha; Luhya: Musila;
Swahili: Mbura; Digo: Mbonobono, Mukokotsaka; Meru/Embu: Munyanwe;
Nandi: Chepsebil (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Harungana madagascariensis Poir.: Small to medium-sized shrubby tree with

distinctly russet young leaves and branchlets. Bark brown, rough and scaly, with
orange sap when damaged. Leaves opposite, elliptic, up to 20 cm long, dark shiny
green above, densely covered in rusty hairs below; margin entire. Young leaves at
stem apex flattened together; hence the common name “praying hands.” Flowers in
dense, branched, terminal heads, creamy-white, hairy inside, with glandular dots
near the apex, sweetly scented. Fruit in clusters, 2–4 mm in diameter, greenish-
orange to red when ripe (Milne-Redhead 1953).
Harungana madagascariensis: The leaves are crushed and rubbed on the head for
headache. A bark decoction is used for malaria. A root decoction is applied to hasten
breast development in young women. Roots and bark boiled in water are used for
colds, to interrupt menstruation, and as eyedrops. The leaf and bark decoction is
drunk for diarrhea, sore throat, and venereal diseases. A leaf paste is applied to burns
Harungana madagascariensis: The light wood is durable (Beentje 1994).
References
Milne-Redhead E. Flora of tropical East Africa: Hypericaceae. London: Crown Agents for Overseas
Hypericum revolutum Vahl
HYPERICACEAE

Local Names
Hypericum revolutum: Kikuyu: Mwania-Thenge, Susimua; Kipsigis:

Biriwaokiret; Elgeyo: Chepboriokororiet, Chepkoberet; Marakwet: Kwalolia;
Maa: Osasimwa; Pokot: Chepsigaga (Beentje 1994; Gachati 1989; Kokwaro 2009)
Botany and Ecology
Hypericum revolutum Vahl.: Much-branched evergreen shrub or tree (1)2–6(12) m

high, with scaly bark. Young sterns 4-angled, soon becoming woody and terete.
Leaves sessile, lamina (11)20–30(37) (2)5–6(10) mm, lanceolate or oblong-
lanceolate, acute at the apex, narrowing to a clasping base, with pinnate or reticu-
late venation, translucent glands (mostly longitudinal streaks or interrupted lines)
R. W. Bussmann (*)
G. N. Njoroge
J. Grimshaw

and small marginal glandular dots (translucent or sometimes dark). Flowers solitary,
terminal. Bracts (when present) leaf-like, with dark marginal dots. Sepals ovate,
somewhat concave, with dark sub-marginal dots, and sometimes also dark fringing
glands. Petals yellow, (1.5) 2.5-3 cm long, about 3 times as long as the sepals, often
tinged with orange, with a few dark marginal dots or glandular. Androecium of
5 stamen-fascicles, each with about 30 stamens. Ovary 5-locular, styles 5 with only
the lower parts united or almost free, 4–7 mm long. Fruit capsular, 5-valved. (Milne-
Redhead 1953). Characteristic timberline species in Afromontane forests (Bussmann
2002a, b, 2006a, b) (Figs. 1, 2, 3, 4, and 5).
Fig. 1 Hypericum revolutum

(Hypericaceae), Hypericum
densely covered in Usnea,
Naro Moru route, timberline,
Kenya. (Photo
R.W. Bussmann)
Hypericum revolutum Vahl 595

(Hypericaceae), flowering
branch, Mt. Kenya National
Park, Naro Moru route,
timberline, Kenya. (Photo
R.W. Bussmann)

(Hypericaceae), flower,
Naro Moru route, timberline,
Kenya. (Photo
R.W. Bussmann)

(Hypericaceae), flowering
branch, 3500 m, Bale
Mountains above Goba,
Oromiya Province. (Photo
J. Grimshaw)
Hypericum revolutum: Dried leaves and stems are soaked in water for rheumatism
and diarrhea. Pounded bark is applied to wounds and burns (Kokwaro 2009). The
leaves are boiled and given to babies with stomach problems, and to make them stop
crying (Bussmann et al. 2011).
Hypericum revolutum: Used as construction timber, especially for house posts.

Leaves sometimes browsed by cattle (Bussmann 2006a, b; Bussmann et al. 2011).
Hypericum revolutum Vahl 597

(Hypericaceae), flowers,
3500 m, Bale Mountains
above Goba, Oromiya
Province. (Photo
J. Grimshaw)
References
2006b;111:41–66.
Milne-Redhead E. Flora of tropical East Africa: Hypericaceae. London: Crown Agents for Overseas
Hyphaene compressa H. Wendl.
Hyphaene coriacea Gaertn.
Hyphaene thebaica (L.) Mart.
ARECACEAE

Grace N. Njoroge
Synonyms
Hyphaene compressa H. Wendl.: Chamaeriphes compressa (H. Wendl.) Kuntze;

Hyphaene benadirensis Becc.; Hyphaene incoje Furtado; Hyphaene kilvaensis
(Becc.) Furtado; Hyphaene mangoides Becc.; Hyphaene megacarpa Furtado;
Hyphaene multiformis Becc.; Hyphaene semiplaena (Becc.) Furtado
Hyphaene coriacea Gaertn.: Chamaeriphes coriacea (Gaertn.) Kuntze;
Chamaeriphes shatan (Bojer ex Dammer) Kuntze; Chamaeriphes turbinata
(H. Wendl.) Kuntze; Corypha africana Lour.; Hyphaene baronii Becc.; Hyphaene
beccariana Furtado; Hyphaene hildebrandtii Becc.; Hyphaene natalensis Kuntze;
Hyphaene oblonga Becc.; Hyphaene parvula Becc.; Hyphaene pileata Becc.;
Hyphaene pleuropoda Becc.; Hyphaene spaerulifera Becc.; Hyphaene tetra-
gonoides Furtado; Hyphaene turbinata H. Wendl.; Hyphaene wendlandii Dammer
Hyphaene thebaica (L.) Mart.: Chamaeriphes crinita (Gaertn.) Kuntze;
Chamaeriphes thebaica (L.) Kuntze; Corypha thebaica L.; Cucifera thebaica (L.)
Delile; Douma thebaica (L.) Poir.; Hyphaene baikieana Furtado; Hyphaene crinita
Gaertn.; Hyphaene cuciphera Pers.; Hyphaene dahomeensis Becc.; Hyphaene
R. W. Bussmann (*)
G. N. Njoroge

dankaliensis Becc.; Hyphaene nodularia Becc.; Hyphaene occidentalis Becc.;

Hyphaene santoana Furtado; Hyphaene sinaitica Furtado; Hyphaene togoensis
Dammer ex Becc.; Hyphaene tuleyana Furtado; Palma thebaica (L.) Jacq.
Local Names
Hyphaene compressa: Swahili: Mkoma, Mkoche, Mlala; Turkana: Eng’oli,

Eeeng’ol; Boni: Mede; Borana: Kone; Gabbra: Meetti; Giriama: Mlala; Ilelewa:
Mezi, Mkoma; Kamba: Mukoma; Oromo: Meti, Kone; Somali: Dabell, Bar;
Sanya: Auwaki; Taveta: Irara (Beentje 1994; Kokwaro 2009).
Hyphaene coriacea: Swahili: Mkoma, Mkoma wa pwani; Borana: Medi; Boni:
Ona (Beentje 1994; Kokwaro 2009).
Hyphaene thebaica: English: Doum palm, Egyptian doum palm, Gingerbread
palm, Gingerbread tree, Dum
Botany and Ecology
Hyphaene compressa H. Wendl.: Massive tree palm eventually reaching heights of

up to 20 m. Trunk solitary and erect, or apparently forking below ground and giving
a group of 2–4 curving trunks of about the same size, sometimes with clusters of
young shoots below, these most probably representing seedlings, trunk about 40 cm
in diameter at the base, decreasing to about 30 cm just below the first dichotomy and
decreasing further at each successive dichotomy to about 20 cm below the ultimate
crowns, the dichotomies in well-grown mature individuals 4–5 in number, giving
16–32 crowns, rarely 6 with 64 crowns, exceptionally branched once only or
unbranched, trunk surface gray with close nodes about 2 cm apart, cracked vertically.
Leaves, unless disturbed, long persisting as a tangled spiny skirt covering the trunk,
usually burnt off leaving the trunk clean, about 15 green living leaves in each crown,
leaf base sheathing, split centrally to form a triangular cleft about 50 cm long, petiole
from 80 cm long in exposed leaves to 1.25 m in juvenile plants, to 3.5 cm wide,
semicircular in cross-section, yellowish in color, except at edges where black, and a
central black line abaxially, armed with fierce distally pointing black spines to 2 cm
long, 6 mm wide, much smaller in juvenile leaves, lamina strongly costapalmate, the
costa sometimes twisted, to 80 cm long, with a spread of about 1.25 m, generally
folded along the costa, divided to about 2/3 the radius into about 30 segments on
each side of the costa, the segments further divided to about 1/3, hastula black,
irregular, or rarely symmetrical, usually lop-sided, 3–4 mm high, with a ragged crest
of hairs, segments stiff, up to 4 cm wide proximally, tapering gradually to the
2 pointed tips, glaucous green with yellowish ribs, dark green inter-leaflet filaments
prominent, black scales abundant, scattered over the lamina surface, fluffy pale
brown scales present on ribs of young leaves. Male inflorescence arcuate to 1.5 m
long, 5 3 cm in diameter at the base, peduncular bracts about 25 cm long, covered
in buff brown hairs and dark scales, partial inflorescences up to 10, each with 1–3
Hyphaene compressa H. Wendl. . . . 601
( 5) rachillae held horizontally, axis of first order branch about 25 cm long by

1.5 cm wide, rachillae about 25 cm long by 1 cm in diameter, bracts on rachillae with
an area about 4 mm wide by 1 mm high exposed, greeny brown, pit hairs dull buff
brown. Female inflorescence as the male but rachillae usually solitary, sometimes
paired, rarely in threes, to 25 cm long by 1.5 cm in diameter, pit about 10 mm wide
by 8 mm long at anthesis, densely filled with pale brown hairs, pedicel about 5 mm
long, lengthening to over 10 mm long at fruit maturity, about 6 mm in diameter,
including the densely packed hairs, sepals and petals about 5 mm long by 7 mm
wide, bluntly triangular, bright green, ovary about 5 mm in diameter, pinkish brown
or pale straw-colored, stigma nectariferous. Male flowers with pedicel about 1 mm
long, sepals 1.5 0.5 mm, petals rounded, about 2 mm long, pale green, stamens
bright canary yellow, filaments about 1.5 mm long, anthers rounded, about 0.5 mm
long, no detectable scent. Male flowers with pedicel about 1 mm long, sepals
1.5 0.5 mm, petals rounded, about 2 mm long, pale green, stamens bright canary
yellow, filaments about 1.5 mm long, anthers rounded, about 0.5 mm long, no
detectable scent. Male inflorescence arcuate to 1.5 m long, 5 3 cm in diameter
at the base, peduncular bracts about 25 cm long, covered in buff brown hairs and
dark scales, partial inflorescences up to 10, each with 1–3( 5) rachillae held
horizontally, axis of first order branch about 25 cm long by 1.5 cm wide, rachillae
about 25 cm long by 1 cm in diameter, bracts on rachillae with an area about 4 mm
wide by 1 mm high exposed, greenish brown, pit hairs dull buff brown. Female
inflorescence as the male but rachillae usually solitary, sometimes paired, rarely in
threes, to 25 cm long by 1.5 cm in diameter, pit about 10 mm wide by 8 mm long at
anthesis, densely filled with pale brown hairs, pedicel about 5 mm long, lengthening
to over 10 mm long at fruit maturity, about 6 mm in diameter, including the densely
packed hairs, sepals and petals about 5 mm long by 7 mm wide, bluntly triangular,
bright green, ovary about 5 mm in diameter, pinkish brown or pale straw-colored,
stigma nectariferous. Young fruit dull maroon-brown to straw-colored with greenish-
cream pits, often with a waxy bloom. Mature fruit rich orangey brown to deep
chestnut, rarely pale golden brown, extremely variable in shape and size from (6–)7–
10( 12) cm tall by (4–)5–8( 9) cm wide at the widest point, varying from almost
cylindrical to oblong, obovoid, or sometimes much deformed, almost always with
2 distinctly compressed, flattened or sunken lateral faces, epicarp dimpled with
conspicuous pock-marks, moderately shiny, flecked with sinuous pock markings,
tough except on very old rotted fruit, mesocarp pale golden brown, usually richly
aromatic (like gingerbread) in ripe fruit, to 1 cm or more thick, pyrene generally
rather oblong to obovoid with 2 flattened faces, endocarp about 4–8 mm thick at
equator and about 10 mm thick round the terminal pore, pore 10–13 mm in diameter,
endosperm about 7 mm thick. Coastal lowlands and extending inland along water-
courses, 0–1400 m, apparently receiving a rudimentary cultivation in cleared areas
apparently extending north to Somalia and southwards to Mozambique (Dransfield
1986) (Figs. 1 and 2).
Hyphaene coriacea Gaertn.: Clustering or solitary palm, occasionally already
mature while still a rosette, tending to form shrubby thickets with decumbent trunks,
up to 5 m tall and very rarely tree-like. Stems suckering, hence building up clumps of
Fig. 1 Hyphaene compressa (Arecaceae), Samburu, Kenya. (Photo R.W. Bussmann)
Fig. 2 Hyphaene compressa (Arecaceae), Samburu, Kenya. (Photo R.W. Bussmann)

uneven aged trunks, dichotomizing once or twice, rarely more, when engulfed by
sand dunes the dichotomies close, producing a clump of crowns on the surface of the
dune, stems gray, to 25 cm in diameter, usually less, usually covered by long-
persistent leaf-bases, occasionally destroyed by fire, leaving very close leaf-scars.
Crown consisting of 8–15 green leaves, petiole to 70 cm long, often much shorter, to
3 cm wide, widening at the very base, the triangular cleft about 20 cm long, petiole
grayish green, armed throughout its length with distally pointing black triangular
spines to 1 cm long and rich brown scales and thin to very dense wax, especially
when young, hastula regular or very asymmetrical. Lamina strongly costapalmate,
the costa 30–80 cm long, usually curved, gray-green to very glaucous, with a dense
covering of wax in young leaves especially in sand dune forms and scattered black
scales, lamina divided to 1/2 the radius in mid-leaf, to 3/4 radius near the edge, into
15–20 segments, to 3.5 cm broad, rather stiff, rarely exceeding 40 cm long. Male
flowers with sepals very narrow, about 3 mm long by 0.5 mm wide, corolla stalk
about 2 mm long, the lobes somewhat spathulate, 3 mm long by 1.5 mm wide,
filaments about 2 mm long, anthers about 1.5 mm long, bright yellow. Male
inflorescence to 1 m, pendulous or arching with 5–7 partial inflorescences, peduncle
about 3 cm in diameter, peduncular bracts about 20 cm long, covered in abundant
brown hairs and white wax, rachillae 1–3 in a group, slender, rarely exceeding 15 cm
long by 0.7 cm wide, rachilla-bracts 1 mm high by 4 mm wide, exposed at anthesis.
Female inflorescence as the male, but rachillae usually 1–2 only in each partial
inflorescence, pits about 4 7 mm at anthesis. Male inflorescence to 1 m, pendulous
or arching with 5–7 partial inflorescences, peduncle about 3 cm in diameter, pedun-
cular bracts about 20 cm long, covered in abundant brown hairs and white wax,
rachillae 1–3 in a group, slender, rarely exceeding 15 cm long by 0.7 cm wide,
rachilla-bracts 1 mm high by 4 mm wide, exposed at anthesis. Female flower with
pedicel about 3 mm long by 1.5 mm wide at anthesis, sepals and petals similar, about
2 mm long by 2 mm wide at the base, triangular, ovary bright green, about 2 mm in
diameter. Male flowers with sepals very narrow, about 3 mm long by 0.5 mm wide,
corolla stalk about 2 mm long, the lobes somewhat spathulate, 3 mm long by 1.5 mm
wide, filaments about 2 mm long, anthers about 1.5 mm long, bright yellow. Female
flower with pedicel about 3 mm long by 1.5 mm wide at anthesis, sepals and petals
similar, about 2 mm long by 2 mm wide at the base, triangular, ovary bright green,
about 2 mm in diameter. Female inflorescence as the male, but rachillae usually 1–2
only in each partial inflorescence, pits about 4 7 mm at anthesis. Fruit extremely
variable in size and shape, from ovoid and pyriform to extreme, cottage-loaf shaped,
rarely 6 cm long and 4 cm wide at the apex, usually much less and occasionally as
small as 3 2.5 cm, usually with a distinct ridge on one side, and a distinct widening
toward the apex, the epicarp generally matt or very finely dimpled, without prom-
inent warts or pockmarks, pale green when immature, ripening mid to dark brown,
mesocarp about 4 mm thick, faintly aromatic, dark brown, endocarp about 3 mm
thick except at the apical pore. Seed somewhat polyhedral, conforming to the shape
Fig. 3 Hyphaene coriacea

(Arecaceae), flowers,
Analavelona, Madagascar.

(Arecaceae), flowers detail
of the endocarp, endosperm about 5 mm thick. Commonest in coastal regions,

especially in sand dunes, and beside creeks behind mangrove, much rarer inland,
in sand dunes it sometimes acts as a pioneer stabilizer, sea level to about 300 m.
Somalia to Mozambique, South Africa, and Madagascar (Dransfield 1986) (Figs. 3,
4, 5, 6, 7, 8, and 9).
Hyphaene thebaica (L.) Mart.: Dioecious tree up to 20 m tall, with dichotomous
branching, trunk solitary, up to 40 cm in diameter, soon dividing into 2 branches,
which may divide again to give 8( 16) crowns, most of the trunk covered with leaf
bases breaking up into fibers near the base of the trunk. Leaves spirally arranged into
dense crowns, with 8–20 leaves per crown, fan-shaped, up to 1.5( 2) m long,
sheathing at the base, petiole up to 140 cm long, margins brown or black with spines
up to 1 cm long, hastula up to 1.5 cm long, costa up to 3.5( 8) cm long, blade
55–90 cm long, costapalmately divided for up to 75% of its length, with up to
40 segments, with a single fiber between each segment, blue-green, segments single-
fold, apex divided, outer half with brown or pale scales. Inflorescence unisexual,

(Arecaceae), leaf,

(Arecaceae), leaf base,
axillary, between the leaves, male inflorescence up to 120 cm long, first erect, later
pendulous, with a tubular prophyll opening out to a lanceolate blade, partial inflo-
rescences 25–30 cm 1 cm, rachillae up to 20 cm about 1 cm, bracts

(Arecaceae), petiole,

(Arecaceae), petiole,
7–8 mm 3 mm and united into a pit with a small opening, flowers 3 per pit,
arranged as a cincinnus, female inflorescence similar to male, up to 80 cm long,
rachillas usually solitary, rarely 2 together, 12–25 cm 1–2 cm, flowers 1 in each

(Arecaceae), leaf base,
pit. Flowers unisexual, 3-merous, male flowers with minute green bracts, calyx with
tubular base and 3 acute, hooded lobes, corolla with a basal stalk and 3 overlapping,
obovate and hooded lobes, stamens 6, on the base of the corolla lobes, female
flowers much larger than male ones, pedicel short and wide, sepals 3, free, triangular,
acute and leathery, petals 3, rounded, somewhat smaller than sepals, scarious,
staminodes 6, ovary globose, 3-locular, but only 1 locule fertile, stigmas 3, sessile.
Fruit a drupe, highly variable in shape and size, ovate to globose to obovate, usually
with some irregular bumps around the base and/or shoulders towards the apex, 4.5–
9 cm 4.5–7( 8) cm, sessile or with short and wide pedicel, smooth, yellowish
brown to orange, red or dark red-brown, dull or shiny, 1-seeded, mesocarp 4–8 mm
thick, dry, fibrous, endocarp 3–8 mm thick, bony. Seed shaped like fruit, 3–
4 cm 2.5–4 cm, endosperm white, with hollow center. Hyphaene thebaica is
distributed from Senegal and Gambia eastwards to Somalia, and is especially
common between latitudes 8°N and 12°N. It also occurs in Libya, Egypt, Israel,
the Arabian Peninsula, and western India. Hyphaene thebaica is often planted. It was
already cultivated in ancient Egypt, where it was considered sacred (Dransfield
1986) (Figs. 10 and 11).
Hyphaene compressa: The fruit pulp is used as anthelminthic and for malaria
(Kokwaro 2009).
Hyphaene coriacea: The seeds are eaten for stomachache (Kokwaro 2009). Young
leaves used to treat diarrhea (Randrianarivony et al. 2016).
Hyphaene thebaica: In traditional medicine in Mali, a paste of the root is massaged
on the chest to relieve chest pain. A maceration of the root bark is taken for the
treatment of intestinal colic and inguinal hernia. In Sudan, an infusion of the leaf
fiber is used as an eyewash for the treatment of conjunctivitis, and the fruit is eaten
Fig. 10 Hyphaene thebaica (Arecaceae), Meru, Kenya. (Photo R.W. Bussmann)
against stomach pain and bladder infection. The mesocarp is credited with diuretic
properties, and a root extract is drunk in case of blood in the urine.
Local Food Uses
Hyphaene coriacea: The seeds are edible (Kokwaro 2009).
Hyphaene compressa: Leaves are used as thatch and fibers for weaving. The seeds
are used as vegetable ivory (Beentje 1994).
Hyphaene coriacea: Leaves are used as thatch and fibers for weaving. The seeds are
used as vegetable ivory (Beentje 1994). In Madagascar, a preferred material for
domestic utensils like baskets and floor mats. Also have cultural uses
(Randrianarivony et al. 2016).
Hyphaene thebaica: Strips from the young, unexpanded leaves are widely used for
weaving mats, bags, baskets, hats, fans, strainers, bowls, rope, string, nets, and
coarse textiles. Older leaves are also used for weaving mats, hats, baskets, ropes,
containers, and other items. In the Sahel, ropes for wells are often made from the
species. The midveins of the leaf segments are used as frames for woven objects and
tied together they are used as brooms. Whole leaves are used for thatching. In
Fig. 11 Hyphaene thebaica

(Arecaceae), Meru, Kenya.
Eritrea, the petioles are woven into bed mats and used in the construction of houses,
fences, and bridges. Fibers from the leaf blade are used for making rough bags, but
the extraction is laborious and the quality is not high. Waste material from fiber
extraction is used in Eritrea for stuffing and reinforcing cement. Fiber from the
petiole is used for making sponges and brushes. The roots yield a fiber used for
making snares and fishing nets and traps. The trunk is used for house construction,
fences, railway sleepers, and canoes. Cut into planks, it is made into canoes and
water wheels. Hollowed trunks are used as water troughs and irrigation pipes. In
Mali, the wood is used for poles, shafts, and harpoons. The wood is also used as fuel
and for making charcoal. The leaves, rachis, and fruits are used as fuel as well. Ash
from the stem is used as a vegetable salt. The apical bud is eaten as a vegetable
(“palm cabbage”). The heart at the base of the trunk is sometimes eaten cooked, and
the hypocotyl of seedlings is also cooked and eaten. Sap is extracted from the tree
just before flowering for the production of palm wine. Young leaves and stems are
browsed by livestock. In Sudan, the young leaves are cut and dried to be used as
fodder during the dry season. Older leaves are bitter and unpalatable. The male
inflorescence is a fodder in Sudan. The fruit mesocarp of some trees is inedible, but
that of other trees is very palatable, with a sweet, gingerbread-like taste. It is

sometimes made into syrup or ground into meal, which is made into cakes and
sweetmeat. The fruits have a hard endocarp, and the “nuts” are made into balls, toys,
and weapons. The endocarp is made into small containers. The hard kernel of mature
seeds was formerly used as vegetable ivory for the production of buttons, beads, and
small carvings. The endosperm of unripe seeds is soft and has a cavity holding a
liquid which is a much-savored drink in northern Nigeria. The endosperm of unripe
seeds is eaten raw or boiled. Dry fruits yield a black dye, which is used for dyeing
leather.
References
Dransfield J. Flora of tropical East Africa: Palmae. Rotterdam: Balkema; 1986.
Hypoestes aristata (Vahl) Sol. ex Roem. &
Schult.
Hypoestes forsskaolii (Vahl.) R. Br.
ACANTHACEAE

Grace N. Njoroge
Synonyms
Hypoestes aristata (Vahl) Sol. ex Roem. & Schult.: Hypoestes antennifera

S. Moore; Hypoestes aristata var. barteri (T. Anderson) Benoist; Hypoestes aristata
var. insularis (T. Anderson) Benoist; Hypoestes aristata var. kikuyensis Benoist;
Hypoestes aristata var. letestui Benoist; Hypoestes aristata var. macrophylla Nees;
Hypoestes aristata var. staudtii (Lindau) Benoist; Hypoestes barteri T. Anderson;
Hypoestes insularis T. Anderson; Hypoestes staudtii Lindau; Hypoestes verticillaris
(L.f.) Sol. ex Roem. & Schult.; Hypoestes verticillaris var. modesta Benist; Justicia
aristata Vahl
Hypoestes forsskaolii (Vahl.) R. Br.: Hypoestes paniculata Schweinf.; Justicia
forsskaolii Vahl; Justicia paniculata Forssk.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Hypoestes aristata: Kikuyu: Githuia, Coco; Shambaa: Kumbukwa; Kipsigis:

Turkwot, Turwot (Gachati 1989; Kokwaro 2009).
Botany and Ecology
Hypoestes aristata (Vahl) Sol. ex Roem. & Schult.: Perennial herb or subshrub to
1 m tall. Branches usually hairy, the hairs in longitudinal bands. Leaves broadly
ovate to lanceolate, lamina 3–13 cm long, 1.5–9 cm wide, apex acuminate, base
attenuate, margins entire, both surfaces usually sparsely hairy, petiole up to 6 cm
long. Inflorescences axillary, flowers appearing clustered due to shortening of
branches, involucre 5–9 mm long. Calyx shorter than involucre. Corolla pink to
mauve-violet or purple except for center band of upper lip white with crowded
purple markings, tube c. 2 cm long, lips 6–14 mm long. Capsule 8–12 mm long,
glabrous. Flowering May–August (Vollesen 2008; Darbyshire et al. 2010). Common
in the undergrowth of dry Afromontane forests (Bussmann 2002).
Hypoestes forsskaolii (Vahl.) R. Br.: Like H. aristata but bracts subspathulate,
flowers white, sometimes mauve, in slender axillary clusters, not appearing verticil-
late and calyx 5-fid, to 50 cm tall. Leaves narrow, 24–36 4.5–13.0 mm; short
cystoliths frequent. Inflorescence not appearing verticillate. Bracts obtuse, strigose.
Corolla lip in lower position frullate. Calyx 5-lobed. Seeds asperous. Flowers white,
pink or light lilac with dark purple honey guides. Flowers: in spikes; bracts with apex
obtuse, strigose; calyx 5-lobed; corolla with lip in lower position trullate, white, pink
or light lilac with dark purple honey guides, flowering February–June (Vollesen
2008; Darbyshire et al. 2010). Common in the undergrowth of dry Afromontane
forests (Bytebier and Bussmann 2000; Bussmann 2002) (Fig. 1).
Hypoestes aristata: The whole plant boiled in water is used for breast problems. The
roots are chewed for sore throat, cold, flu, and cough (Kokwaro 2009). Also used for
stomachache (Njoroge et al. 2004).
Hypoestes forsskaolii is used for bloody diarrhea, chest pain, cough with blood, and
stomachache (Teklehaymanot and Giday 2010; Wondimu et al. 2007).
Local Food Uses
Hypoestes forsskaolii: Leaves are boiled and eaten as famine food (Balemie and
Kebebew 2006).
Hypoestes aristata (Vahl) Sol. ex Roem. & Schult. . . . 613
Fig. 1 Hypoestes forsskaolii

(Acanthaceae), Mt. Kenya
R.W. Bussmann)
Hypoestes aristata and Hypoestes forsskaolii: Browsed by cattle and wildlife

(Bussmann 2006; Bussmann et al. 2011).
References
Darbyshire I, Vollesen K, Kelbessa E. Flora of tropical East Africa: Acanthaceae (part 2). Kew:
Vollesen K. Flora of tropical East Africa: Acanthaceae (part 1). Kew: Royal Botanic Gardens; 2008.
Ilex mitis (L.) Radkl.
AQUIFOLIACEAE

Grace N. Njoroge
Local Names
Ilex mitis: Kikuyu: Muthumara, Munyamati, Musumura; Meru: Murati;

Marakwet/Sebei: Seger; Ogiek/Kipsigis/Ndorobo: Tongotuet; Samburu:
Lchenioibor (Beentje 1994; Gachati 1989; Kokwaro 2009); English: African
holly, Cape holly, Watertree; Swahili: Msaira
Botany and Ecology
Ilex mitis (L.) Radkl.: Dioecious evergreen shrub or small to medium-sized tree up
to 30( 40) m tall, bole straight and cylindrical or short and poorly shaped, up to
100 cm in diameter, bark surface smooth, sometimes becoming slightly warty or
fissured, whitish to grey or pale brown, crown fan-shaped, twigs often short-hairy,
purplish. Leaves alternate, simple, stipules minute, up to 1 mm long, petiole up to
1 cm long, channeled above, blade elliptical to oblong-elliptical or obovate-
elliptical, 2.5–14 cm 1.5–5 cm, cuneate to obtuse at base, obtuse to acute or
R. W. Bussmann (*)
G. N. Njoroge

short-acuminate at apex, margins entire or sometimes spiny-toothed, leathery, gla-

brous, dark green, pinnately veined with looping lateral veins. Inflorescence an
axillary, few-flowered cyme up to 3 cm long, short-hairy, with peduncle up to
1.5 cm long. Flowers unisexual, regular, (4–)5–6-merous, sweet-scented, pedicel
up to 1( 1.5) cm long, calyx c. 1.5 mm long, lobes rounded to acute, short-hairy at
margins, corolla up to 3.5 mm long, white, with short tube, male flowers with
stamens inserted on the corolla tube and alternating with lobes, ovary rudimentary,
female flowers with rudimentary stamens and superior, globose ovary, stigma
sessile. Fruit a globose drupe 4–7 mm in diameter, becoming dark red upon ripening,
containing 4–7 stones, each stone 1-seeded. Seeds 2– mm long, brownish. Ilex mitis
is widespread, occurring in mountain areas from Sierra Leone east to Eritrea,
Ethiopia, and Kenya, and south to South Africa, Swaziland, and Lesotho; it also
occurs in eastern Madagascar (Verdcourt 1968). Character species of Cassipourea
malosana and Olea capensis dominated broad-leaved Afromontane forests
(Bussmann 2002a, b, 2006) (Figs. 1 and 2).
Ilex mitis: In East Africa, bark decoctions are administered for the treatment of
sterility, and in southern Africa as an emetic and as an enema to treat fever and colic.
Powdered bark is applied externally to treat rashes and sores. Influenza patients are
washed with bark and leaf macerations. Roots are used to treat fits and swellings. In
Madagascar, bark and root decoctions are taken against gonorrhea, whereas decoc-
tions of leaves, stems, roots, and bark are used to clean wounds. A pomade is made
from Ilex mitis (containing 0.75% of purified extract) which is popular in Madagas-
car under the name “fanaferol” as a skin care product to treat burns, sores, wounds,
and scars, and as a skin lightener. The bark decoction is used to treat infertility in
women. The bark is chewed for abdominal pain (Kokwaro 2009). Leaves are applied
Fig. 1 Ilex colchica

(Aquifoliaceae), sapling,
Adjara, Georgia. (Photo
R.W. Bussmann and
Ilex mitis (L.) Radkl. 617
Fig. 2 Ilex colchica

(Aquifoliaceae), Adjara,
Georgia. (Photo
R.W. Bussmann and
to remedy backpain (Rabearivony et al. 2015). Leaves and stem bark extract are
applied topically for dermal problems (Yineger et al. 2008).
Local Food Uses
Ilex mitis: The fruits are edible, being slightly sweet to slightly bitter, and boiled
roots are also reported to be edible. Excellent honey source (Beentje 1994). The seed
oil is used to grease baking plates before baking (Bussmann et al. 2011).
Ilex mitis: The flowers are an important source of nectar for honeybees, and the
honey is whitish and of good quality. The tree is planted in parks and large gardens
for its decorative glossy leaves and red fruits. It is also planted in hedges. The wood
is suitable for light construction, light flooring, interior trim, ship building, vehicle
bodies, furniture, toys, novelties, musical instruments such as drums and guitars, tool
handles, agricultural implements, railway sleepers, boxes, crates, vats, beehives,
matches, veneer, plywood, hardboard, particle board, and pulpwood. It has been in
demand for buckboards and spokes in wagon construction, and for the heels of
ladies’ shoes. It is also used as firewood and for charcoal production, although for the
last purpose it seems less suitable. Hard wood (Beentje 1994). The leaves are used to
envelope dough before putting it in the oven, so that it does not burn. The leaves are
used as amulet against bad luck (Razafindraibe et al. 2013).
References
2006;111:41–66.
2013;9:73.
Verdcourt B. Flora of tropical East Africa: Aquifoliaceae. London: Crown Agents for Overseas
2008;26:132–53.
Juniperus procera Hochst. ex Endl.
CUPRESSACEAE

Grace N. Njoroge
Synonyms
Juniperus procera Hochst. ex Endl.: Juniperus hochstetteri Antoine; Sabina pro-

cera (Hochst. ex Endl.) Antoine
Local Names
Juniperus procera: Kikuyu: Mutarakwa; Swahili: Mwangati; Borana: Aru;

Gabbra: Arru; Kamba: Ndalakwa, Tarakwet; Samburu: Tarakwa; Kipsigis/
Marakwet/Nandi: Tarakwet; Maa: Oltarakwa; Meru: Murana; Pokot: Tarakit,
Pokot; Tugen: Tarokwa, Torokwa; Rendile: Hal; Turkana: Eminet; Watende:
Teet; Chagga: Matarakwa (Beentje 1994; Gachati 1989; Kokwaro 2009); English:
African pencil cedar, East African cedar, East African juniper, Pencil cedar
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Juniperus procera Hochst. ex Endl.: Straight evergreen trees up to 40 m tall, bark

papery, reddish-brown. Leaves on young plants in threes, subulate, spine-tipped,
with an elongate gland abaxially, mature foliage of decussate scale-like leaves,
varying in size as the individual ages down to 1 mm long, abaxial gland elliptic.
Male cones 3 mm long, elliptic, scales about 10, each bearing 2–3 pollen sacs. Male
cones 3 mm long, elliptic, scales about 10, each bearing 2–3 pollen sacs. Female
strobili of 6–8 scales. Ripe fruit 6 mm in diameter, globose, blue-black, with the tips
of the scales just distinguishable. Mature seeds 5 mm long, 1 or 2, brown, aborted
seeds 1–3, smaller. Juniperus procera occurs wild from Sudan, Eritrea, and Ethiopia
southwards through East Africa and eastern DR Congo to Malawi and Zimbabwe; it
also occurs wild in Saudi Arabia and Yemen (Melville 1958). Juniperus procera
grows rather fast in comparison to other primary forest species (Bussmann 1999).
Pencil Cedar needs light to germinate and does not germinate without scarification,
which naturally is done by fire. The seeds of the species stores exceptionally well for
years (Bussmann and Lange 2000). The species forms the most extensive
Afromontane forests (Bussmann 1996; 2002a, b, c, 2004, 2006a, b; Bytebier and
Bussmann 2000) (Figs. 1, 2, 3, 4, 5, 6, 7, 8, 9, and 10).
The dry and semi-deciduous broad-leaved Afromontane forests fall into two
groups, dominated by Cassipourea malosana and Juniperus procera. The climax
type for both types is, however, characterized by Juniperus procera, albeit they
regenerate differently.
No indications of a regenerative cycle can easily be observed in the Cassipourea
malosana dominated forests. In general, the regeneration follows a dynamic pattern.
Trees of all age classes were found in the Cassipourea forests suggesting a mosaic
climax. Due to the environmental conditions, especially the high rainfall, fires are
rare in the Cassipourea forests and, whenever occurring, they were limited to small
Fig. 1 Juniperus procera

(Cupressaceae), branch of
sapling, Mt. Kenya National
Park, Sirimon Route, Kenya.
Juniperus procera Hochst. ex Endl. 621

(Cupressaceae), branch of
mature tree, Mt. Kenya
National Park, Sirimon Route,
Kenya. (Photo R.W.
Bussmann)
areas. Juniperus procera occurs only sporadically in the Cassipourea forests, as its
germination and regeneration is suppressed by heavy shade and the dense vegetation
covering the forest floor for this reason. Regenerating Cedar was only found in gaps
where the bright light, requiring seedlings compete with those of the broad-leaved
species.
In contrast to the Cassipourea malosana dominated forests, fire apparently plays
an important role in the regeneration cycle of the Juniperus procera dominated
forests. The African Pencil Cedar typically grows in the drier montane, submontane,
and subalpine areas of the East African mountains between 1000 and 3000 m
(Kerfoot 1961). Temperature as well as rainfall apparently limit its spreading
(Winiger 1979).
In his study of the South Aberdare Range, Wimbush (1937) suggested that
without fire, the Juniperus forest is only a seral stage in the succession to a broad-
leaved forest, proposing a fire-induced regeneration cycle.
After fire, herbaceous pioneer species and Juniperus procera seedlings invade the
gap (1), leading to a regenerating forest (2), where broad-leaved species as Olea
capensis ssp. hochstetteri, Olea europaea ssp. cuspidata, and Olinia rochetiana
become established too. After about 20 years, the Juniperus trees have grown about
9 m high. When a closed canopy is established, the Cedar trees start to outgrow the
other species, reaching about 30 m after 80–90 years. If the number of Juniperus
trees established after a fire was high, a pure Cedar forest (3) results by suppression
Fig. 3 Juniperus procera (Cupressaceae), very young regenerating Juniperus, Mt. Kenya National
Park, Sirimon Route, Kenya. (Photo R.W. Bussmann)
of other trees. After about 200 years, the maximum height of about 40–46 m is
reached. Although frequently infected by the fungus Fomes juniperinus, thinning the
crowns, the Juniperus trees may live for another 300–700 years. In case a burnt area
is mainly invaded by broad-leaved species, these create the canopy and only
scattered Juniperus specimens emerge. As Cedar cannot regenerate without fire,
natural gaps are filled with broad-leaved species. Therefore, pure broad-leaved
forests built the climax community (4) after some time. These broad-leaved forests
represent a stable climax, if not destroyed by fire (Fig. 11) (Bussmann and Beck
1995; Bussmann 2002b).
The Cassipourea malosana dominated forests can be seen as part (4) of the
mentioned cycle, although the impact of the rare fires appears to be very limited.
Scattered Juniperus trees regenerate in natural gaps without fire, especially so at the
upper limit.
In areas where Juniperus procera has been removed by logging, more or less pure
stands of broad-leaved trees were encountered with only few Cedar trees left.
However, most Cedar forests were in stage 4 of the succession cycle. Absolutely
pure stands of Cedar (stage 3) were never found at lower and medium altitudes.
Forests clearly dominated by Juniperus procera were seen only at higher altitudes
forming the upper borderline of these forests. From several large burnt areas on
Mount Kenya, the following regeneration process was concluded (Fig. 12).
After fire, a pioneer stage with a very high initial cover of Juniperus and Myrsine
africana develops (1), where soon broad-leaved trees start growing, producing

(Cupressaceae), young tree,
Sirimon Route, Kenya. (Photo
R.W. Bussmann)
various independent climax subassociations (2), due to different ecological condi-

tions and different seed intake. However, Juniperus procera remains always the
dominant canopy species. These climax stages are open forests with only scattered
shrubby undergrowth, allowing regeneration of all tree species. Thus, seedlings and
young trees of Juniperus were often encountered in these forests, even when fire had
been absent for a longer period (3). This indicates that at higher altitudes broad-
leaved forests in sensu Wimbush (1937) cannot replace the Juniperus forests, even if
burning has not occurred for long time. If a fire occurs, a new regeneration cycle
takes place.
Juniperus procera: In traditional African medicine, an infusion of ground young

twigs is taken against intestinal worms. People with rheumatism are treated by
exposure to the smoke of burnt twigs and seed cones. The smoke is also inhaled
as an expectorant. Ground dried leaves are applied on wounds of humans and
animals. A hot bath to which the leaves are added is used in the treatment of fever.
The resin is used as a stimulant and applied to ulcers. Bark macerations are drunk
and applied as a vaginal wash as birth-control agents. A decoction of the seed cones
is used as a sudorific and emmenagogue. Young twigs ground, soaked in water, and

(Cupressaceae), huge, mature
tree, see red clad person to the
right, Mt. Kenya National
drunk as anthelminthic (Kokwaro 2009). The fruits are chewed for sore throat
(Bussmann 2006a, b). The leaf extract is used against vomiting (Giday et al. 2007;
Teklehaymanot et al. 2007). The leaves are used to make a tea to alleviate cold and
flu (Luizza et al. 2013; Njoroge and Bussmann 2006). The dried cones are used to
treat intestinal problems (Sher et al. 2016).
The genus Juniperus is very widely used in medicine. Juniperus sabina is used
for diseases of the skin. A powder from the leaves is used externally for the treatment
purulent ulcers. Fresh crushed leaves are pulverized and applied with sweet butter on
scabies and for the elimination of warts (Batsatsashvili et al. 2017). A water infusion
is used against worm in small doses (Batsatsashvili et al. 2017). The fruits are used as
diuretic, for prostate problems, urinary problems, and to treat toothache. The leaves
are used to treat skin problems. The roots are employed for blood cleansing, as
detoxicant, for gallbladder ailments, and urinary system problems (Batsatsashvili
et al. 2017; Bussmann et al. 2017, 2018; 2020a, b, c; Bussmann 2017). Juniperus
semiglobosa: The fruits are used to increase appetite, aid in digestion, as diuretic,
disinfectant of the urinary system, anti-inflammatory, for pneumonia, and as anal-
gesic and expectorant. The oils are used for skin problems (Eisenman et al. 2012).
Juniperus seravschanica: Smoke from burning branches is used in Middle Asia for
rheumatism. Powdered material is inhaled to treat headaches. The oil is used for
wounds and skin diseases. Internally an infusion of dry fruits is used for urogenital
problems, scurvy, liver disease, rheumatism, edema, and nervous disorders. Fresh

(Cupressaceae), large
Juniperus among Olea
europaea ssp. africana
(grayish foliage), Mt. Kenya
National Park, Sirimon Route,
Kenya. (Photo R.W.
Bussmann)
fruits are eaten to treat stomach ulcers and to increase the appetite. The bark
decoction is used as choleretic to treat jaundice. The root infusion is used to remedy
stomach ulcers, bronchitis, tuberculosis, kidney stones, treat arthritis, and male
impotency. A decoction of the fruits and green branches is used as bath for rheuma-
tism (Eisenman et al. 2012). Juniperus indica is used for kidney problems (Kunwar
et al. 2006). Juniperus communis and Juniperus squamata serve to treat skin
problems (Ur-Rahman et al. 2018).
Juniperus procera has ceremonial and religious significance, as in some parts of

Ethiopia, where it is used especially in September during the traditional orthodox
ceremony of Meskel. In veterinary medicine, chopped and finely ground leaves
mixed with water are used as a drench for horses and mules with stomach disorders,
whereas a decoction of dry young branches is a medicine against itch of camels. The
wood of Juniperus procera is widely used for building (both construction and
lining), joinery, flooring (strip and parquet), furniture, and all sorts of outdoor
work such as roofing shingles, fence posts, water flumes, and transmission poles.
In Kenya, the wood is also used for making fire sticks, beehives, and salt troughs.
Formerly the wood was exported to Europe and North America for the manufacture
Fig. 7 Juniperus procera (Cupressaceae), Juniperus procera in open landscape, Bale Mountains
National Park, Odo Bulu, Ethiopia. (Photo R.W. Bussmann)
Fig. 8 Juniperus procera (Cupressaceae), Juniperus procera in open agricultural landscape, Bale
Mountains National Park, Odo Bulu, Ethiopia. (Photo R.W. Bussmann)

(Cupressaceae), small natural
fire in Juniperus forests,
Sirimon Route, Kenya. (Photo
R.W. Bussmann)

(Cupressaceae), large,
man-made fire in Juniperus
forest, Mt. Kenya National
of pencils and penholders, while small quantities were used for wardrobe linings. It
is also suitable for ship and boat building, agricultural implements, musical instru-
ments, carving, vats, toys and novelties, turnery, draining boards, and food con-
tainers. It can be used for making veneer and plywood, hardboard and particle board,
and as pulpwood. The wood is used as firewood and to make charcoal. The bark is
used for roof shingles and for covering beehives. Essential oil distilled mainly from
Fig. 11 Regeneration cycle of Afromontane, Cassipourea malosana dominated broad-leaved

forests. (Bussmann and Beck 1995; Bussmann 2002b)
Fig. 12 Regeneration cycle of Afromontane xeromorphic forests dominated by Juniperus procera

(Bussmann and Beck 1995; Bussmann 2002b)
the sawdust (“cedar wood oil,” “cedar oil”) is used in the cosmetic industry in soaps
and perfumes. Since Juniperus procera can grow in extreme conditions, it is
replanted in deforested areas for soil conservation or improvement and for erosion
control, e.g., in Eritrea, Ethiopia, and Kenya. It is also a useful shade tree and is
frequently planted as an ornamental tree and in windbreaks. Excellent timber tree for
construction, poles, and beehives (Beentje 1994). Juniperus is used to fend off evil
eye in livestock (Assefa et al. 2010). The branches are browsed by livestock and can
be used to fill mattresses (Bussmann 2006a, b). The timber is excellent for construc-
tion and firewood (Bussmann et al. 2011; Luizza et al. 2013; Mekonnen et al. 2015),
and for household tools (Mekonnen et al. 2015). Sometimes used for veterinary
purposes (Yineger et al. 2007). Extracts are used to remedy ectoparasites in livestock
(Wanzala 2017).
Juniperus sabina is used for dyeing: A dye solution is prepared from needless
young shoots and immature fruits to obtain olive color. The dye is used for dyeing
wool and silk in yellow and gray shades, and for scented soaps (Batsatsashvili et al.
2017). The leaves serve as diuretic and for dermatological problems in veterinary
medicine (Batsatsashvili et al. 2017; Bussmann et al. 2016a, b, 2017, 2018;
Bussmann 2017).
References
10.1186/1746-4269-6-20.
Batsatsashvili K, Mehdiyeva N, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S,
Tchelidze D, Alizade V, Paniagua Zambrana NY, Bussmann RW. Juniperus communis L.,
Juniperus sabina L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer
International Publishing; 2017.
Bussmann, R.W. 2002c. Vegetation ecology and regeneration of tropical mountain forests. In:
Ambasht RS, Ambasht NK, editors. Modern trends in applied terrestrial ecology. New York:
Kluwer. p 195–223.
Bussmann RW. Vegetation zonation and nomenclature of African mountains – an overview. Lyonia.
2006a;111:41–66.
Bussmann RW, editor. 2017. Ethnobotany of the Caucasus. Springer International Publishing,
Cham. XXVII, 746 p. ISBN 978-3-319-49411-1.
Bussmann RW, Beck E. Regeneration- and succession processes in the cedar-forests Juniperion
procerae of Mount Kenya. Ecotropica. 1995;1:79–84.
Nat Hist. 2000;89(1–2):101–11.
(Sakartvelo), Caucasus. J Ehnobiol Ethnomed. 2016a;12:43. https://doi.org/10.1186/s13002-
016-0110-2.
2167-0412.1000266.
gia), Caucasus. Indian J Tradit Knowl. 2017;16(1):7–24.
Bussmann RW, Batsatsashvili K, Kikvidze Z. Juniperus pseudosabina var. turkestanica (Kom.)
Silba., Juniperus sabina L., Juniperus semiglobosa Regel, Juniperus seravschanika Kom. In:
Batsatsashvili K, Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions Central
Asia and Altai. Cham: Springer International Publishing; 2020a.
Bussmann RW, Batsatsashvili K, Kikvidze Z, Paniagua-Zambrana NY, Khutsishvili M, Maisaia I,
Sikharulidze S, Tchelidze D. Juniperus communis L., Juniperus depressa Raf., Juniperus
hemispherica J. Presl & ABOUT Presl, Juniperus oblonga M. Bieb., Juniperus sabina L. In:
Batsatsashvili K, Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions far
eastern Europe. Cham: Springer International Publishing; 2020b. https://doi.org/10.1007/978-3-
319-77088-8_77-2.
Sikharulidze S, Tchelidze D. Juniperus communis L., Juniperus depressa Raf., Juniperus
hemispherica J. Presl & C. Presl, Juniperus oblonga M. Bieb., Juniperus sabina L. In:
Batsatsashvili K, Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions far
eastern Europe. Cham: Springer International Publishing; 2020c. https://doi.org/10.1007/978-3-
319-77088-8_77-2.
Eisenman SW, Zaurow DE, Struwe L, editors. Medicinal plants of Central Asia: Uzbekistan and
Kyegyzstan. Stuttgart: Springer Scientific; 2012.
Kerfoot O. Juniperus procera Endl. (The African Pencil Cedar) in Africa and Arabia –
I. Taxonomic affinities and geographical distribution. E Afr Agr For J. 1961;26:170–7.
study from Dolpa, Humla, Jumla and Mustang districts of Nepal. J Ethnobiol Ethnomed.
2006;2:27.
Res Appl. 2013;11:315–39.
practices. J Ethnobiol Ethnomed. 2015:11(64).
Melville R. Flora of tropical East Africa: Gymnospermeae. London: Crown Agents for Overseas
livestock industry with focus on Acari ticks. Evidence-Based Complement Alternat Med .
Article ID 8647919, 33 pages. 2017; https://doi.org/10.1155/2017/8647919.
Wimbush SH. Natural succession in the Pencil Cedar forests of Kenya Colony. Emp For
J. 1937;16:49–53.
Winiger M. Bodentemperaturen und Niederschlag als Indikatoren einer klimatisch-ökologischen
Gliederung tropischer Gebirgsräume. Geomethodica. 1979;4:121–50.
Kalanchoe densiflora Rolfe
Kalanchoe petitiana A. Rich.
CRASSULACEAE

Grace N. Njoroge
Local Names
Kalanchoe densiflora: Kikuyu: Mahuithia, Mukondori; Shambaa: Sopolwa;

Lhuya: Linyolonyolo (Beentje 1994; Gachati 1989; Kokwaro 2009).
Kalanchoe petitiana: Kikuyu: Mahuithia-Meru (Beentje 1994; Gachati 1989;
Kokwaro 2009).
Botany and Ecology
Kalanchoe densiflora Rolfe: Glabrous perennial herb, stem erect, 0.3–3 m high.
Episepalous stamens with anthers included within the tube by 1.5–2.5 mm, epipetalous
stamens with tip of anthers almost attaining the throat of the tube or included with the
tube by up to 1 mm, anthers with apical glands 0.05–0.1 mm in diameter, anthers of both
whorls nearly the same size, broadly oblong, 0.7–0.85 mm long, 0.5–0.7 mm wide.
Nectary scales linear-subulate, 1–3.5 mm long. Leaves petiolate, blade oblong, ovate or
R. W. Bussmann (*)
G. N. Njoroge

suborbicular, up to 18 cm long, 10 cm wide, apex obtuse, base cuneate, margins crenate,

light green, petiole up to 3 cm long, flattened and grooved above, slightly broadened at
the base. Carpels linear-lanceolate, 5–9 mm long, terminating in styles 1–3.5 mm long.
Corolla-tube pale to greenish yellow, limb yellow to dark orange, reddish orange or red,
tube 8–12 mm long, lobes oblong-lanceolate to broadly obovate, 2–5 mm long, 1.5–
4 mm wide, truncate or retuse, mucronate (mucro 0.2–1 mm long). Calyx-lobes
triangular-subulate, 2–8 mm long, 1–1.5 mm wide, long-attenuate, connate at the base
for 0.1–0.3 mm, glabrous, rarely sparsely pilose (var. subpilosa). Inflorescence a densely
flowered paniculate cyme up to 30( 70) cm long, pedicels 1–4 mm long, conspicuously
broadened below the flower, glabrous, rarely sparsely pilose (var. subpilosa Cuf. from
Ethiopia) (Wickens 1987). Found in clearings and open spaces in dry Afromontane
forests (Bussmann 2002) (Figs. 1, 2, and 3).
Kalanchoe petitiana A. Rich.: Root thick, fibrous. Stem 2–6 ft. high, glabrous through-
out or hispidulous in the upper portion. Leaves 2–3 in. long, oblong or roundish ovate or
spathulate, coarsely crenate, obtuse, cuneate at the base, glabrous, all but the uppermost
distinctly petiolate, petioles 1–1 1/2 in. long; upper leaves sometimes entire. Inflores-
cence glabrous or hispidulous in trichotomous many-flowered axillary and terminal
Fig. 1 Kalanchoe densiflora

(Crassulaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
Kalanchoe densiflora Rolfe. . . 635
Fig. 2 Kalanchoe densiflora

(Crassulaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
cymes; peduncles of axillary cymes long, the lowest 6–7 in. in length; internodes 3–6 in.
long. Bracts narrow, lanceolate, petiolate; bracteoles setaceous. Calyx-lobes glabrous or
hispidulous and viscid, lanceolate, acute, very slightly cohering at the base, 2–3 lines
long; pedicels slender, 2–3 lines long. Flowers bright yellow or orange; tube 1/2 in. long,
glabrous or pubescent; limb-segments oblong-lanceolate, acute. Squamulæ linear, 1–1
1/2 lines long. Carpels narrow, 3 lines long. Found in clearings and open spaces in dry
Afromontane forests (Bussmann 2002) (Figs. 4, 5, and 6).
Kalanchoe densiflora: Root and leaf decoction is used for abortion. The leaf juice is
squeezed on septic wounds. Hot leaf decoction for massages in rheumatism, stiff
joints, and swellings (Kokwaro 2009).
Kalanchoe petitiana: The roots are used to treat constipation (Yineger et al. 2007),
stomachache (Teklehaymanot 2009), and swellings (Teklehaymanot and Giday
2007). The species is also used as anthelmintic, especially for tapeworms, and to
Fig. 3 Kalanchoe lanceolata

(Crassulaceae), Bale
treat trachoma and syphilis, and has shown antibacterial and antifungal properties
(Tadeg et al. 2005). It is also used for skin itch (Giday et al. 2010), and headache
(Giday et al. 2007). Heated leaves are applied to swellings and sores (Bussmann
et al. 2011).
Kalanchoe glaucescens is used for rheumatic problems (Nankaya et al. 2019).
Kalanchoe laciniata serves to remedy tonsillitis (Yineger et al. 2008). Kalanchoe
quartiniana has antibacterial properties (Desta 1995).
Kalanchoe petitiana: Heated leaves are wrapped around broken limbs in livestocks
Fig. 4 Kalanchoe petitiana



References
Tadeg H, Mohammed E, Asres K, Gebre-Mariam T. Antimicrobial activities of some selected
traditional Ethiopian medicinal plants used in the treatment of skin disorders. J Ethnopharmacol.
2005;100:168–75.
peninsula, northwestern Ethiopia. J Ethnobiol Ethnomed. 2007;3:12. https://doi.org/10.1186/
1746-4269-3-12.
Wickens GE. Flora of tropical East Africa: Crassulaceae. Rotterdam: Balkema; 1987. 66 pp. ISBN
90-6191-331-4.

ailments at Bale Mountains National Park, southeastern Ethiopia. J Med Plants Res.
2008;26:132–53.
Kigelia africana (Lam.) Benth.
BIGNONIACEAE

Grace N. Njoroge
Synonyms
Bignonia africana Lam.; Crescentia pinnata Jacq.; Kigelia abyssinica A. Rich.;

Kigelia acutifolia Engl. ex Spreng.; Kigelia aethiopica Decne.; Kigelia aethiopicum
(Febzl) Dandy; Kigelia elliottii Sprague; Kigelia elliptica Sprague; Kigelia impressa
Sprague; Kigelia pinnata (Jacq.) DC.; Kigelia spragueana Wernham; Kigelia
talbotii Hutch. & Dalziel; Kigelia tristis A. Chev.; Sotor aethiopicum Fenzl;
Tanaecium pinnatum (Jacq.) Willd.
Local Names
Kigelia africana: Kikuyu: Kiratina, Muratina; Taita: Mwasina, Masina,

Mvopngnia; Lhuya: Mnikya, Murabe, Morabe; Luganda: Mussa; Meru:
Murantina; Maa: Ol-Sunguroi, Ol-Darpoi; Luo: Yago; Boni: Shelolo; Ilelewa:
Mobwoka; Kamba: Muatine, Kiatine; Giriama: Muratina; Kipsigis: Ratuinet;
Marakwet: Rotio; Nandi: Ratinuet; Oromo: Bogh; Pokot: Rotin; Rendile: Muun;
R. W. Bussmann (*)
G. N. Njoroge

Samburu: Imombi; Somali: Bukorola; Taveta: Mukisha; Turkana: Edot (Beentje

1994, Gachati 1989, Kokwaro 2009); English: Sausage tree, Cucumber tree; Swahili:
Mvungunya, Mwegea, Mwicha, Mranaa
Botany and Ecology
Kigelia africana (Lam.) Benth.: Tree often with wide spreading crown, 2.5–18
(–24) m tall and possibly larger. Bark usually gray, occasionally brown, smooth to
rough or ridged, scaly or flaking. Leaves opposite or ternate. Leaflets thin to very
coriaceous, (2–)3–8-jugate, oblong, elliptic, ovate or obovate, (6–)10–20
(–30) (4–)6–13(–16) cm, acute to cuneate or rounded, truncate, or emarginate at
the base, entire to serrate (particularly sucker shoots), rounded, obtuse or acute to
distinctly acuminate at the apex, glabrous to pubescent or tomentellous, smooth or
often slightly to very scabrid with raised pale dots, peltate and punctate glands
sometimes present; venation plane or sometimes raised or impressed, with 7–12
pairs of veins. Flowers said to smell unpleasant, in lax terminal hanging panicles
30–80(–150) cm long including peduncle up to 40 cm long. Bracts lanceolate, 1 cm
long. Individual cymes 1–3-flowered. Pedicels hooked, 1–4(–5) cm long. Calyx
campanulate, 2–5 cm long, 2-lipped, irregularly lobed, the lobes about 1 cm long,
tomentose to glabrous, with scattered glands. Corolla at first with tube yellow
outside, turning orange or red and with dark red to blackish inside, 3–9 cm long,
cylindrical at base for a very variable length; lobes yellow-green outside suffused
with crimson, dark crimson inside, ovate, 3–4.5 cm long and wide. Fruits
cylindrical, 30–90 cm long, 7.5–10 cm wide, tough and woody; stalks up to 50 cm
long. Seeds ovoid, 11 mm long, 7 mm wide, and 4 mm thick. Kigelia africana occurs
throughout tropical Africa, particularly in the drier regions. It is also found in
South Africa (Northern Provinces, Kwazulu-Natal) and Swaziland, but does not
occur in Mauritania, São Tomé and Principe, or the Indian Ocean islands. It has been
introduced as an ornamental to Cape Verde and Madagascar, as well as to Iraq,
Pakistan, India, China, South-East Asia, Australia, Hawaii, and Central and South
America (Bidgood et al. 2006). Mostly a savannah species, but found at the edges of
disturbed, dry, Afromontane forests in transition to savannah (Bussmann 2002)
(Figs. 1, 2, 3, 4, 5, 6, and 7).
Kigelia africana is widely used throughout Africa in local medicine. The diversity of
illnesses against which the plant is used includes fainting, anemia, sickle-cell
anemia, epilepsy, respiratory ailments, hepatic and cardiac disorders, and nutritional
illnesses such as kwashiorkor, rickets, wasting, and weakness. The leaves are
sometimes used to prepare a general tonic for improved health and growth. Aqueous
fruit preparations are applied as a wash or rub to promote weight gain in infants. The
roots, bark, leaves, stems, twigs, and fruits are used to treat digestive disorders.
Kigelia africana (Lam.) Benth. 643
Fig. 1 Kigelia africana (Bignoniaceae), Maasai Mara, Kenya. (Photo R.W. Bussmann &
Fig. 3 Kigelia africana

(Bignoniaceae), Maasai Mara,
Kenya. (Photo R.W.
Bussmann & N.Y. Paniagua-
Zambrana)
Administration is typically by oral ingestion or as an enema. The roots, bark, and

ripe or unripe fruits are taken as a laxative or emetic, to treat chronic and acute
digestive disorders and against gastric infections. Infections of the genitourinary
tract, particularly venereal diseases, are treated both internally and externally with
preparations of the roots, bark, leaves, stems, and twigs. In West and Central Africa
the dried and ground bark is macerated in palm wine and taken against syphilis.
Venereal diseases in children are treated simultaneously with a drink and wash
prepared from decocted bark. In Côte d’Ivoire, renal and bladder ailments are treated
with medicaments containing the bark and leaves of Kigelia africana and several
other medicinal plants. Aqueous preparations of the roots, fruits, and flowers are
administered orally or as a vaginal pessary. The fruits and bark are used to promote
breast development in young women, or in contrast to reduce swelling and mastitis
of the breasts. The fruits are further employed as a galactogogue. The bark and
leaves are decocted and administered as an abortifacient. Sexual complaints such as
infertility, poor libido, sexual asthenia, and impotence are treated with medicines
containing the fruits, roots, or leaves. A small amount of unripe fruit is chewed, or an
aqueous preparation is taken orally as a sexual stimulant, and the intoxicating
Fig. 5 Kigelia africana (Bignoniaceae), fruits, Maasai Mara, Kenya. (Photo R.W. Bussmann &

(Bignoniaceae), flower,
Guayaquil, Ecuador. (Photo
R.W. Bussmann &

(Bignoniaceae), flower,
Guayaquil, Ecuador. (Photo
R.W. Bussmann &
traditional beer to which they are added is drunk as an aphrodisiac. Excessive use of
Kigelia africana to treat male sexual complaints is said to induce scrotal elephanti-
asis, although in some regions the fruits are used to remedy this condition. Powders
and infusions of the bark, leaves, stems, twigs, or fruits are used to clean and dress
flesh wounds and open sores. Many dressings, topical treatments, and infusions are
also used for their analgesic and anti-inflammatory properties. The bark, stems,
twigs, leaves, and fruits are infused and taken orally, or applied locally, to relieve
rheumatism, sprains, hematoma, and bruising; a decoction of the fruit and bark is
used to relieve toothache and headache. Snake bite antidotes are made with an
infusion of the fruits, stems, leaves, twigs or bark, taken orally or rubbed onto the
bite. A fruit decoction is used to treat edema of the legs. Kigelia africana is used to
treat infectious diseases including leprosy, impetigo, and worm infestations in the
blood. Dermal complaints and infections, such as whitlows, cysts, acne, and boils,
are treated with traditional medicines containing the fruits, and less frequently, the
bark. Sore eyes are treated with drops made from flower sap mixed with water.
Fungal infestations such as ringworm, mycosia, and athlete’s foot are washed with
the water in which bark has been macerated, and preparations containing the leaves
and fruits applied locally. A root decoction is administered against internal parasitic
infestations, especially tapeworm. Kigelia africana is also used in both traditional
and allopathic medicines to treat malignant neoplasms such as skin melanoma,
tumors, and breast cancer. Traditional preparations include extracts, poultices, and
powders of the bark or fruits; topical creams containing extracts of the fruits are
produced commercially. Beer made from the fruits is used to wash children with
measles. The bark decoction is used for epilepsy, headache, venereal diseases, and
rheumatism. A leaf decoction serves for malaria, and the root decoction for gastro-
intestinal problems. Roasted leaves with honey are used to lower blood pressure
(Kokwaro 2009). Also used to treat skin diseases, ulcers, diabetes, as purgative and
for diarrhea (Jeruto et al. 2008), for common cold (Njoroge and Bussmann 2006),
and to treat measles (Muthee et al. 2011). Kigelia pinnata has shown antileishmanial
activity (Rocha et al. 2005).
Local Food Uses
Kigelia africana: Both ripe and unripe fruits are poisonous to humans, but the fruits
can be dried and used to start fermentation and used along with the bark to enhance
the flavor of traditional beers (Kokwaro 2009). The seeds are sometimes roasted and
eaten in times of food shortage.
Kigelia africana: The wood makes good quality timber for fences, planking, boxes,
and canoes. It is also used for making boxes, drums, stools, yokes, tool handles,
mortars, and large bowls for watering cattle. Bows are made from branches, and
smaller branches are hollowed to administer enemas to children. Wood and fruits are
carved into mousetraps, dolls, and various items of crockery and cutlery. The wood
is used as fuel. A black dye is obtained from the tannin-rich fruit pulp. The fruits, and
sometimes other plant parts, are also much used in ethnoveterinary medicine to treat
digestive system disorders, leg edemas, dermal irritations and infections, mastitis
and retained placenta, and for Brucellosis and Newcastle disease. The fruit pulp is
said to be inedible and toxic and may cause blistering of the tongue and skin.
However, fallen fruits along with leaves and flowers are browsed or foraged by
livestock and game. Due to the unusual fruits and large flowers, Kigelia africana is
considered a striking ornamental plant, and the fruits are used as florists’ material.
The thick stem is an attractive feature for bonsai. The tree is sometimes planted as a
boundary marker, but usually at roadsides and for shade. Due to its occurrence along
watercourses, it is suitable for erosion control and riverbank stabilization. Kigelia
africana is regarded as sacred in many regions; religious meetings are held under-
neath the tree, and the flowers and fruits are regarded as fetish. Fruits are commonly
sold in markets as charms to promote wealth and prosperity, to impart strength and
courage on warriors prior to, to increase crop yields, and as a fetish for fecundity, or
to avert whirlwinds. Bark and root decoction is given to calves for intestinal
problems (Kokwaro 2009) and to livestock for East Coast fever (Kioko et al.
2015). The species has antibacterial activity (Malik et al. 2015).
References
Bidgood S, Vollesen K, Verdcourt B. Flora of tropical East Africa: Bignoniaceae & Cobaeaceae.
Appendices 1–7. www.naturekenya.org/JournalEANH.htm.
borne diseases among Maasai people in Northern Tanzania. Vet World. 2015;8(6):755–62.
Rocha LG, Almeida JRGS, Mace RO, Barbosa-Filhob JM. A review of natural products with
antileishmanial activity. Phytomedicine. 2005;12:514–35.
Kyllinga bulbosa P. Beauv.
CYPERACEAE

Grace N. Njoroge
Synonyms
Kyllinga bulbosa P. Beauv.: Cyperus cyperinus (Retz.) Srringar; Kyllinga nana

Nees
Local Names
Kyllinga bulbosa: Kikuyu: Ngonda, Ngothe (Gachati 1989).
Botany and Ecology
Kyllinga bulbosa Beauv.: Perennials. Rhizomes short. Culms scattered, 15–70 cm

tall, slightly stout, acutely triquetrous, smooth, base many leaved. Leaves shorter
than culm; sheath purplish red; leaf blade 5–7 mm wide, slightly flaccid or stiff,
folded at basal part and spreading at apical part, abaxially midvein and margin
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Kyllinga
sp. (Cyperaceae). Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
scabrous. Involucral bracts 6–10, leaf-like, longer to slightly shorter than inflores-
cence, margin scabrous. Inflorescence a simple anthela; rays 6–10, short or to
4.5 cm, unequal in length, with a single spike at apex. Spikes oblong-obovoid,
often appearing to taper toward base or rarely cylindric, 1–1.8[ 3] 0.8–1.2 cm,
with many densely arranged spikelets, base cuneate to subtruncate. Spikelets erect to
erect-spreading, narrowly linear-ovoid to linear-oblong, 4–6.5 ca. 1 mm, each
(1 or)2–4-flowered; rachilla wings lanceolate, broad. Glumes crowded, brownish to
purplish red on both surfaces, elliptic, ca. 3.5 mm, papery, veins several with middle
3 conspicuous and green, keeled, apex obtuse to acute and not mucronate. Stamens
3; anthers broadly linear, 1–1.5 mm; connective prominent beyond anthers. Style of
medium length; stigmas 3. Nutlet dark grayish brown, narrowly oblong, 2–
2.5 0.6–0.9 mm, ca. 2/3 as long as subtending glume, 3-sided, slightly curved
in dorsiventral view, with dense punctate processes. Flowering and fruiting April–
September (Wu et al. 1994–2013). In Afromontane forests (Bussmann 2002)
(Fig. 1).
Kyllinga bulbosa P. Beauv. 651
Kyllinga species are boiled and the infusion is applied topically for heart problems
and luck (Bussmann and Sharon 2006).
References
Landolphia buchananii (Hallier f.) Stapf
Landolphia kirkii DyIer
APOCYNACEAE

Grace N. Njoroge
Synonyms
Landolphia buchananii (Hallier f.) Stapf: Clitandra buchananii Hallier f.;

Clitandra kilimandjarica Warb.; Clitandra semlikiensis Robyns & Boutique;
Jasminochyla ugandensis (Stapf) Pichon; Landolphia cameronis Stapf; Landolphia
kilimandjarica (Warb.) Stapf; Landolphia rogersii Stapf; Landolphia swynnertonii
S. Moore; Landolphia ugandensis Stapf
Landolphia kirkii DyIer: Clitandra stapfiana A. Chev.; Landolphia delagoensis
(Dewèvre) Pierre; Landolphia dondeensis Busse; Landolphia polyantha K. Schum.
Local Names
Landolphia buchananii: Kikuyu: Muugu; Kamba: Kiongao; Embu: Nakamwe;

Kipsigis: Ngigichet, Tuneiyet; Luo: Rikikini; Maa: Entiangege; Nandi:
Ngengichet, Ngunyet; Samburu: Seebit; Tugen: Mugugugwa (Beentje 1994;
R. W. Bussmann (*)
G. N. Njoroge

Landolphia kirkii: Swahili: Mpira; Giriama: Mungo, Mtongasi (Beentje 1994).
Botany and Ecology
Landolphia buchananii (Hallier f.) Stapf: Liana 3–40 m long, rarely sarmentose
shrub 0.9–7 m high with copious white latex, stems deeply fluted, to 22 cm in
diameter, dark brown, branchlets glabrous or pilose. Leaves petiolate, blade elliptic
to ovate or obovate, 1.9–14.5 cm long, 0.8–6 cm wide, obtuse to bluntly acuminate at
the apex with acumen up to 1 cm long, rounded to cuneate at the base, sometimes
slightly ciliate, glabrous above, lower surface completely glabrous or pubescent at the
base of midrib and lower secondary veins, petiole 1.5–8 mm long, glabrous or pilose,
with 4–8 triangular glands on each node, often with empty bracts in between nodes.
Inflorescence terminal, sometimes also axillary, 1.6–5.5 cm long, 2–20-flowered,
peduncle 5–33 mm long, sometimes tendril-like, elongate and curved, bracts ovate
to triangular, 1–2.5 cm long, pedicels 1.2–6.5 mm long. Flowers fragrant, sepals ovate,
1.2–3.1 mm long, rounded at the apex, corolla white, creamy or yellow, tube often
greenish, occasionally tinged pink or reddish, 6–12 mm long, lobes narrowly ovate to
elliptic, 5.3–15.5 mm long, 1.9–4 mm wide, glabrous, rounded, stamens inserted
1.8–3.6 mm from the base of the tube, included for 1.3–5.9 mm, ovary ovoid, glabrous
or with a few hairs. Fruits green with white or light brown verrucose spots, 1–50 mm
in diameter, globose or pyriform, 2.5–6 cm long, 2–20-seeded, pericarp 5–6.5 mm
thick, pulp fleshy, white, turning yellow when exposed, seed up to 17 mm long,
irregularly ovoid, laterally compressed, minutely pitted. Riverine or swamp forest,
450–2400 m (Bussmann 2002; Omino 2002; Goyder et al. 2012) (Fig. 1).
Landolphia kirkii Dyer: Straggling shrub or liane attaining 18–30 m in length, with
rough dark bark. Young stems brownish- or ferrugineous-pubescent, later glabrescent.
Upper leaf surface fairly glossy, almost always puberulent when young, glabrescent, or
less often glabrous except for the pubescent midrib, very rarely lamina completely
glabrous, midrib channelled, lateral and tertiary nerves more or less raised, never
impressed, lower surface crispate-pubescent or glabrous except for the appressed- or
spreading-pubescent midrib, midrib prominent, other nerves raised or level with
surface, tertiary reticulation easily seen with a hand-lens. Leaves membranous to
subcoriaceous, usually drying discolorous with the upper surface much darker than
the lower, petiole 2–7 mm long, pubescent, lamina 2.4–9.2 1–3.2 cm, oblong to
narrowly ovate or rarely oblong-obovate, the apex attenuate or cuspidate-attenuate into
a short to long acumen, or acumen more or less absent, the base rounded to cuneate.
Tendrils common. Inflorescences terminal, many-flowered paniculate cymes forming
dense clusters or sometimes the branches elongating and becoming tendrillous, axes
densely rufous-tomentose or rufous-pubescent. Flowers scented, each subtended by
two sepal-like bracteoles and the inflorescence branches by bracts. Calyx 2 mm long,
sepals almost free, oblong, grayish-brown, not glossy, ciliate, dorsally glabrous or
puberulent, with hispid midrib. Corolla white, cream-colored or yellow, darker on the
tube and reverse of petals, corolla tube 3.5–4 mm long, clavate, the wall thickened in
the upper half and the mouth very restricted, exterior surface puberulous especially
Landolphia buchananii (Hallier f.) Stapf. . . 655
Fig. 1 Landolphia buchananii (Apocynaceae), Mt. Kenya National Park, Ocotea usambarensis
dominated forest, Kenya. (Photo R.W. Bussmann)
above the calyx, corolla lobes narrowly elliptic, more or less equalling the tube,
glabrous. Stamens inserted at or just above the middle of the corolla tube, anthers
subsessile, 0.75 mm long, reaching to base of constricted part of corolla tube. Ovary
0.75–1 mm long, glabrous, style, clavuncle and stigma 1–1.5 mm long. Fruit globose
or pyriform, attaining 15 cm in diameter, green with white flesh, edible. Seeds 8.5–
10 mm long (Omino 2002; Goyder et al. 2012).
Landolphia buchananii: Fruits are eaten to clean infected tongue. Root decoction is
used to treat bilharzia and gonorrhea (Kokwaro 2009). Leaf infusion is topically
applied for wounds and gonorrhea (Jeruto et al. 2008).
Local Food Uses
Landolphia buchananii: Fruits are edible (Beentje 1994).

Landolphia kirkii: Fruits are edible (Beentje 1994).
Landolphia buchananii: Young stems are used as ropes (Beentje 1994, Bussmann
et al. 2006). The leaf infusion serves as molluscicide (Jeruto et al. 2008).
References
Goyder d, Harris T, Masinde S, Meve U, Venter J. Flora of tropical East Africa: Apocynaceae (part
E. Africa, Apocyn. (Part 2)].
Omino EA. Flora of tropical East Africa: Apocynaceae (part 1). Rotterdam: Balkema; 2002. 114 pp.
ISBN 90-5809-409-X.
Lantana camara L.
VERBENACEAE

Grace N. Njoroge
Synonyms
Lantana camara L.: Camara aculeata (L.) Kuntze, Camara aculeata fo. obtusifolia
Kuntze, Camara aculeata var. subinermis Kuntze, Camara vulgaris Benth., Lantana
aculeata L., Lantana antillana Raf., Lantana armata S. Schauer, Lantana armata
var. guianensis Moldenke, Lantana camara fo. macrantha (Loes.) Moldenke, Lan-
tana camara fo. mista (L.) Moldenke, Lantana camara fo. urticulata (Mill.)
I.E. Méndez, Lantana camara subsp. articulata (L.) Moldenke, Lantana camara
var. alba Moldenke, Lantana camara var. crocea Bailet, Lantana camara var. flava
(Medik.) Moldenke, Lantana camara var. macrantha Loes., Lantana camara var.
mista (L.) L.H. Bailey, Lantana camara var. moritziana (Otto & a. Dietr.) López-
Pal., Lantana camara var. nivea (Vent.) L.H. Bailey, Lantana camara var. parviflora
(Moldenke) Moldenke ex D.N. Gibson, Lantana crocea Jacq., Lantana crocea var.
guatemalensis Loes., Lantana flava Medik., Lantana foetida Rusby, Lantana for-
mosa C. Koch & Fintelm., Lantana frutilla var. obtusifolia Moildenke, Lantana
glanulosissima fo. albiflora Moldenke, Lantana glanulosissima fo. parviflora
Moldenke, Lantana glandulosissima Hayek, Lantana glanulosissima var. grandis
R. W. Bussmann (*)
G. N. Njoroge

Moldenke, Lantana hirsuta M. Martens & Galeotti, Lantana hirsuta var. pubescens
Moldenke, Lantana hirta fo. ternata Moldenke, Lantana hispida fo. alba Moldenke,
Lantana hispida fo. parviflora Moldenke, Lantana hispida Kunth, Lantana hispida
var. ternata Moldenke, Lantana horrida fo. inermis Moldenke, Lantana horrida
Kunth, Lantana horrida var. grandiflora Schauer, Lantana horrida vra. parviflora
Schauer, Lantana mexicana Turner, Lantana minasensis var. hispida Molenke,
Lantana mista L., Lantana moritziana Otto & A. Dietr., Lantana nivea Vent.,
Lantana scabrida Sol., Lantana scandens Moldenke, Lantana scorta Moldenke,
Lantana spinosa L. ex Le Coimte, Lantana tiliifolia Cham., Lantana tiliifolia
fo. albiflora Moldenke, Lantana triplinervia var. hispida (Moldenke) Moldenke,
Lantana urticifolia Mill.
Local Names
Lantana camara: Kikuyu: Mukigi, Kagiri, Mukenia; Digo: Mjasasa; Kamba:

Kitavisi; Luo: Nyamrih; Tugen: Getipkamoskon (Beentje 1994; Gachati 1989).
Botany and Ecology
Lantana camara L.: Branching shrubs, up to 2 m high, stems and branches mostly
unarmed or slightly prickly, usually minutely and inconspicuously pubescent.
Leaves decussate-opposite, the blades chartaceous, ovate to oblong-ovate,
2–12 cm long and 2–4.5 cm wide, acute or short-acuminate (rarely obtuse apically),
acutely narrowed or abruptly rounded to a subcuneate acumination basally, crenate-
serrate, more or less reticulate-rugose and decidedly scabrous or scabrellous above,
usually only sparsely pilosulous or strigillose beneath (mostly only on the venation)
with canescent or brownish trichomes, sometimes glabrescent, petioles 7–12 mm
long. Inflorescences axillary, shorter than, equaling, or surpassing the subtending
leaves, the heads always capitate, hemispheric, to 3 cm wide, not elongating after
anthesis, many-flowered, peduncles slender, 2–9 cm long, more or less appressed-
pilose or puberulent, bractlets usually not conspicuous, oblong to lanceolate, equal,
Lantana camara L. 659
4–7 mm long, and 1–1.5 mm wide, subulate or acute, rarely a few larger ones also
present, appressed strigose-pubescent, usually about as long as the corolla-tube.
Flowers with the calyx thin, about 3 mm long, corolla hypocrateriform, mostly
orange-yellow or orange, changing to red or scarlet in age, the tube about 10 mm
long, barely enlarged above the middle, slightly curved, puberulent, the limb
6–8 mm wide. Drupes fleshy, black, about 3 mm in diameter.
Lantana camara: Introduced, aggressive invasive species around the tropics. Often
incorporated in the local pharmacopoeia (Kumar et al. 2013; Kunwar et al. 2015).
Used in Ethiopia to treat fever (Giday et al. 2010), colds, malaria, flu, diarrhea
(Njoroge et al. 2004; Njoroge and Bussmann 2006a; Tabuti 2008), cough, otitis
media (Njoroge and Bussmann 2006b), tetanus, and inflammation (Muhammad et al.
2019). Used in Pakistan to treat ringworm, headache, aerodontalgia, malaria, fever,
rheumatoid arthritis, cuts, wounds, cough, and cold (Umair et al. 2019). In Mada-
gascar even sold in local markets as remedy for hemorrhages and hypertension
(Randriamiharisoa et al. 2015), epilepsy, malaria (Randrianarivony et al. 2016;
Rabearivony et al. 2015), and diarrhea (Rakotarivelo et al. 2015). In some areas
for treating sexual problems and intestinal parasites (Rabearivony et al. 2015). Also
used to remedy cough and cold in India (Raj et al. 2018), and for indigestion (Luizza
et al. 2013).
Lantana ukambensis and Lantana trifolia are used as medicinal species in Ethiopia
(Bekalo et al. 2009).
Local Food Uses
Lantana camara: The fruits are edible (Randrianarivony et al. 2016). The leaves are
used as spice (Luizza et al. 2013) (Figs. 1 and 2).
Fig. 1 Lantana
sp. (Verbenaceae), Laikipia
plateau, Samburu, Kenya.
Lantana camara: Sometimes planted as ornamental (Mekonnen et al. 2015).

Lantana camara L. 661
Fig. 2 Lantana cf. camara

(Verbenaceae), Laikipia
plateau, Samburu, Kenya.
References
org/10.1186/1746-4269-5-26.
Kumar M, Sheikh MA, Bussmann RW. Effect of fire on soil nutrients and under storey vegetation in
Chir pine forest in Garhwal Himalaya, India. Acta Ecol Sin. 2013;33(1):59–63.
Res Appl. 2013;11:315–39.
plants known by men at Ambalabe. 2015.
018-0208-9.
org/10.1186/s13002-015-0046.
2013;9:73.
10.1186/s13002-019-0285-4.
Lasianthus kilimandscharicus K. Schum.
RUBIACEAE

Grace N. Njoroge
Local Names
Lasianthus kilimandscharicus: Kikuyu: Mukwethi, Murundi; Kipsigit: Ruandet

Botany and Ecology
Lasianthus kilimandscharicus K. Schum.; Shrub or small tree 1.2–7.5 m tall, with

smooth grey bark, fresh wood reported in Uganda to smell unpleasant, shoots
glabrous or finely pubescent, drying black above but with pale yellowish bark, the
nodes often with some persistent indumentum below the stipules but older stems
quite glabrous. Leaf-blades oblong, narrowly oblong-elliptic or oblong-lanceolate,
(4–)9–17( 22) cm long, (1.2–)2–6( 7) cm wide, narrowly acuminate at the apex,
cuneate at the base, firmly papery but not coriaceous, glabrous above and usually
R. W. Bussmann (*)
G. N. Njoroge

beneath save for fine sparse adpressed pubescence on the nerves or rarely hairy,
lateral nerves (5–)8–10 on each side, petiole 0.5–1( 1.8) cm long, stipules narrowly
to broadly triangular, 1.5–6 mm long, hairy, particularly along the margins. Fruit
intense cobalt blue, subglobose, prominently 4–6-lobed in dry state but not grooved
when fresh, 4.5 (dry) to 10 (fresh) mm in diameter, finely puberulous, the persistent
calyx-lobes oblate, very obtuse, constricted at the base, 2 mm long, 2 mm wide,
pyrenes chestnut-brown or straw-colored, basically pyriform but with a marked
ventral notch extending from the pointed end for about half the length of the pyrene,
3.2–4 mm long, 2.3–2.5 mm wide, 2 mm thick. Ovary 4–6-locular, style 4.5–5.5 mm
long in long-styled flowers, 2.5 mm long in short-styled flowers, stigma-lobes 4–5,
oblong or subcapitate, 0.5–0.6 mm long. Calyx pinkish white or tinged purple,
particularly on the lobes, glabrous or puberulous, tube turbinate, 2 mm long, lobes
very convex in living state 0.5–3 mm long, 2 mm wide, sometimes with pubescent
traces of intermediate accessory lobes. Flowers few, sessile in the axils of the leaves,
heterostylous, bracts ovate to lanceolate, 2–6 mm long, 1–2 mm wide (? sometimes
stipules of reduced shoots), with distinctly hairy margins. Stamens with anther-tips
just exserted in long-styled flowers but with anthers and 1 mm of filaments
exserted in short-styled flowers. Corolla glistening white or violet outside, white
inside, tube cylindrical, 2.5–4( 5) mm long, glabrous to slightly pubescent outside,
with densely hairy throat but tube glabrous inside or hairy only in upper half, lobes
4–5, ovate-oblong, 2.2–3.7 mm long, 1.5–2 mm wide, densely hairy inside with
white hairs, finely hairy or glabrous outside. (Verdcourt 1976; Bridson and Verdcourt
1988, 1991). Commonly found in Afromontane forests (Bussmann 2002a, b, 2006)
(Figs. 1 and 2).
Lasianthus kilimandscharicus: Sometimes used as firewood.

Lasianthus kilimandscharicus K. Schum. 665
Fig. 1 Lasianthus
kilimandscharicus
(Rubiaceae), Mt. Kenya
National Park, Chogoria
forest. (Photo R.W:
Bussmann)
Fig. 2 Lasianthus
kilimandscharicus
National Park, Chogoria
forest. (Photo R.W:
Bussmann)
References
Bridson DM, Verdcourt B. Flora of tropical East Africa: Rubiaceae (Part 2). Rotterdam: Balkema;
1988. p. 415–747. ISBN 90-6191-337-3. [Fl. Trop. E. Africa, Rub. (Part 2)].
1991. p. 749–956. ISBN 90-6191-357-8 [Fl. Trop. E. Africa, Rub. (Part 3)].
appendices 1-7 @ www.naturekenya.org/JournalEANH.htm.
2006;11(1):41–66.
Verdcourt B. Flora of tropical East Africa: Rubiaceae (Part 1). London: Crown Agents for Overseas
Governments & Administrations; 1976. 414 pp. ISBN 0-85592-043-32. Invalid ISBN [Fl. Trop.
E. Africa, Rub. (Part 1)].
Lippia javanica (Burm. f.) Spreng.
VERBENACEAE

Grace N. Njoroge
Synonyms
Lippia javanica (Burm. f.) Spreng.: Lippia galapiniana H. Pearson, Lippia

asperifolia A. Rich. ex Marthje, Verbena javanica Burm. f.
Local Names
Lippia javanica: Kikuyu: Muthoroti, Muthiriti, Mucohi; Kipsigis: Mwokyot, Kisii:

Onyinkwa; Luo: Ang’we-rao, Mweny; Marakwet: Mwokio; Nandi:
Chepngosoriet; Samburu: Sunoni; Sebei: Karnet; Kamba: Kyulu, Mutithi; Maa:
Ol-Sinoi; Acholi: Orwo; Luhya: Sulasula; Iraqw: Ufani (Beentje 1994; Gachati
1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Lippia javanica (Burm. f.) Spreng.: Much-branched shrub 0.6–4.5( 6) m. tall, stems
brown with adpressed or spreading short stiff tubercle-based hairs and small glands.
Nutlets brown, half-ovoid, 1.3 mm long, 1 mm wide. Corolla white, yellowish white or
greenish (rarely pink), usually with a yellow throat, tube narrowly funnel-shaped from a
very narrow base, 1.2–2 mm long, pubescent and glandular above, limb 1.2–2 mm
wide, the largest lobe squarish, 0.5 mm long, 0.8 mm wide. Leaves opposite or in whorls
of 3, blades lanceolate, elliptic- or ovate-lanceolate or oblong-elliptic, 1–10
( 13.5) cm long, 0.3–4.3( 5) cm wide, obtuse to acute at the apex, rounded then
cuneate at the base, closely serrulate, densely pubescent with tubercle-based hairs and
shorter hair above, slightly to distinctly scabrid and rugulose bullate, softly and densely
often velvety adpressed pubescent beneath and nerves raised, also with small glands.
Calyx 1 mm long, 2-lobed, densely spreading pubescent. Flowers in conical or oblong
spikes 0.45–1.5( 2.2) cm long, (3–)5–8 mm wide, 1–5 per axil, sessile or with
peduncles 0.3-3 cm long, lower bracts of spikes ovate, 4 mm long, 2 mm wide, long-
acuminate, upper smaller, 1.8 mm long, 1.2 mm wide, acuminate, densely spreading
white pubescent and glandular. (Smith 1987; Smith and Carter 1988) (Figs. 1, 2, and 3).
Fig. 1 Lippia javanica

(Verbenaceae), Sekenani
Camp, Maasai Mara, Kenya.
Lippia javanica (Burm. f.) Spreng. 669

Lippia javanica: The leaf infusion is used for fever. Leaves and flowers are snuffed
for congestion after rubbing them between the hands. A decoction of boiled leaves
serves for malaria and is both drunk and used as bath. Pounded leaves are applied to
wounds. The leaf juice is used as anthelminthic and for indigestion (Fassil 2005;
Kokwaro 2009). Used for colds, flu, and anaplasmosis (Njoroge et al. 2004).
Employed to treat chest pains, for cleaning wounds, colds, constipation, general
body health, heartburn, induce vomiting, joints, skin rashes, stomachache, and as
anthelminthic (Nankaya et al. 2019, 2020; Njoroge and Bussmann 2006). Lippia
kivuensis is used as purgative (Muthee et al. 2011). Lippia adoensis is chewed for
stomach problems, constipation, and stomach pain (Teklehaymanot 2009; Yineger
et al. 2008). The species has shown antimicrobial activity (Tadeg et al. 2005).

Local Food Uses
Lippia adoensis is used as spice and fragrance in Ethiopia (Mekonnen et al. 2015).
Lippia javanica: The plant is used ceremonially after circumcision – it is put on the
parent’s bed for its good smell. Also browsed by sheep and goats (Bussmann et al.
2006).
Lippia javanica (Burm. f.) Spreng. 671
References
Fassil H. Beyond plants, professionals & parchments: the role of home-based medicinal plant use
and traditional health knowledge in primary health care in Ethiopia. Ethnobot Res Appl.
2005;3:37–49.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Medicinal plants of the maasai of Kenya: a review.
Plant. 2020;9:44. https://doi.org/10.3390/plants9010044.
1988. p. 409–597. ISBN 90-6191-338-1.
2005;100:168–75.
2008;26:132–53.
Lobelia giberroa Hemsl.
CAMPANULACEAE

Grace N. Njoroge
Local Names
Lobelia gibberoa: Kikuyu: Muhehe; Elgeyo: Chepkaset; Kamba: Ewewe;

Kipsigis: Serenguet, Chepkurbet; Kisii: Omomoa; Maa: Embenyoi-dabas, Oljani-
Oseba; Nandi: Chepkwaret; Taita: Mukirombosha (Beentje 1994; Gachati 1989).
Botany and Ecology
Lobelia giberroa Hemsl.: Large herb. Stem erect, 6–12 cm thick and woody at the base,
terete, hollow, more or less pubescent above. Inflorescence 1–3 m. long, dense, cylin-
drical, unbranched, pedicels 4–8 mm long, pubescent, with 2 min linear bracteoles,
bracts linear to narrowly lanceolate, 25–75 mm long, 2–4 mm wide. Rosette-leaves of
nonflowering plant sessile, narrowly oblanceolate to oblong-ovate, 25–90 cm long,
4–19 cm wide, acute to acuminate at the apex, attenuate at the base, pubescent beneath
especially on the veins or glabrous, weakly pubescent or glabrous above, margin doubly
R. W. Bussmann (*)
G. N. Njoroge

serrate, dentate or denticulate, venation protruding beneath with main lateral nerves in
more or less 30–40 pairs arising at an angle of 60–90° to the midrib. Filaments connate
for most of their length, more or less densely papillose at the margins, free from corolla.
Corolla 25–35 mm long, greenish or greenish white, often tinged with blue or purple,
split to the base on the back, pubescent on the outside at least at the apex, glabrous
inside, two lateral petals linear, more or less united with middle petals below, three
middle petals united to form a lip with linear lobes. Calyx-lobes narrowly triangular to
linear, 10–28 mm long, entire, more or less pubescent especially on the outside.
Hypanthium obovoid, more or less truncate at the base, 10-nerved, pubescent. Capsule
10–15 mm long, 10-nerved, pubescent, crowned by the hardening calyx-lobes and with
2 short valves. Anther-tube (6–)9–15 mm long, 2 lower anthers apically barbate,
otherwise glabrous, or occasionally pubescent on connectives. Ovary subinferior.
Seeds elliptic-oblong in outline, compressed, narrowly winged, more or less 0.8 mm
long, finely striate, pale brown. (Thulin 1984). As companion species in Afromontane
forests, often close to the timberline (Bussmann 2002, 2006a, b) (Figs. 1 and 2).
Fig. 1 Lobelia gibberoa

(Campanulaceae), Mt. Kenya
National Park, Maro Moru
Roite, Hagenia abyssinica
R.W. Bussmann)
Lobelia giberroa Hemsl. 675
Fig. 2 Lobelia gibberoa

(Campanulaceae), Mt. Kenya
National Park, Maro Moru
Roite, Hagenia abyssinica
R.W. Bussmann)
The roots of Lobelia sp. are boiled, and the decoction used to treat malaria (Giday
et al. 2007). The sap of leaves and flowers of Lobelia pyramidalis is applied
externally for body ache. Leaves and inflorescence are antispasmodic and used for
asthma, bronchitis and fever. Leaves and flowers are anti-spasmodic and they are
used as an expectorant. The plant is also used for sciatica and back pain (Kumar et al.
2011, Kunwar et al. 2010). The crushed roots of Lobelia rhynchopetalum, one of the
very few Afro-alpine plants used for medicine, are used to treat the condition
“Gubbaa” (Teklehaymanot 2009; Yineger et al. 2007).
Lobelia holstii is eaten by livestock and used to fill mattresses (Bussmann 2006a, b).
References
Bussmann RW. Succession and regeneration patterns in East African Mountain forests – A Review.
Systematics and Geography of Plants. 2002;712:959–974.
2006b;11(1):41–66.
Giday M, Teklehaymanot T, Animut A, Mekonne Y. Medicinal plants of the Shinasha, Agew-awi
habitation in Garhwal Himalayan, India. Journal of Medicinal Plant Research. 2011;511:
2252–2260.
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in far-West Nepal: a
Thulin M. Flora of tropical East Africa: Lobeliaceae. Rotterdam: Balkema; 1984.
Macaranga kilimandscharica Pax
EUPHORBIACEAE

Grace N. Njoroge
Local Names
Macaranga kilimadscharica: Kikuyu: Mukuhakuha; Cherangani: Erwa;

Ndorobo: Legumeta, Lugumeta; Kamba: Mukalati; Luhya: Mukaho,
Mukuhakuha; Marakwet: Kibgetoyoa; Meru: Mukarati; Nandi: Sebsebet; Sebei:
Kaptemba; Kipsigis/Ndorobo: Logomaita (Beentje 1994; Gachati 1989; Kokwaro
2009).
Botany and Ecology
Macaranga kilimandscharica Pax: A small to medium tree 4.5–18 m, or large tree

up to 27 m., often with a pyramidal crown, much branched, the branches ascending,
or with a broad, spreading crown. Fruit transversely ovoid, subglobose or bilobate,
4–6 mm long, 5–11 mm diameter, densely granulate-glandular, dull green, styles
laterally displaced in the monococcus fruits. Seeds subglobose, (3.5–)4–4.5 mm
diameter, shallowly malleate-rugulose or smooth, dull or shiny, dark brown to
R. W. Bussmann (*)
G. N. Njoroge

blackish. Petioles (1–)5–10( 14) cm long, pulvinate at the base, indumentum as

in M. capensis, leaf-blades triangular-ovate to rhombic-lanceolate, (3–)5–15
( 17) cm long, (1.5–)3–10 cm wide, acutely acuminate, entire, cuneate, rounded,
truncate, occasionally cordulate, peltate or not, usually biglandular at the petiole-
insertion, firmly chartaceous to thinly coriaceous, 3–7( 9)-nerved from the base,
lateral nerves (5–)7–12 pairs, not prominent above, prominent beneath,
brochidodromous, tertiary nerves parallel or subparallel, not or scarcely concentric
at the base, densely minutely uniformly glandular-punctate beneath, the glands
drying blackish, ferruginous tomentellous above and beneath and occasionally
whitish hirsute towards the bases of the midrib and main nerves beneath at first
(rarely over the entire surface), later usually glabrescent, dark or very dark green
with pallid nerves above, usually somewhat glaucous with yellowish brown nerves
beneath. Wood soft, heavy. Young shoots and inflorescence-axes densely ferrugi-
nous tomentellous at first, later glabrescent. Bark green at first, later becoming
greyish white, light or dark grey, pale reddish brown or dark greyish brown, thin,
smooth or longitudinally striated or fluted, with prominent lenticels. Inflorescences
axillary, solitary. Stipules narrowly lanceolate to linear-lanceolate, 8–15 mm long,
2–3 mm wide, acutely acuminate, entire, densely ferruginous tomentellous, cadu-
cous. Male flowers subsessile, calyx-lobes 3, elliptic-ovate, 1 mm long, puberulous
without, glabrous within, yellowish green or cream, stamens 2, filaments united at
the base, 1 mm long, whitish, anthers obscurely 4-thecous, 0.5 mm across, yellow.
Male inflorescences paniculate, (3–)5–10 cm long, axes zigzag towards the apex,
bracts ovate-lanceolate to triangular-ovate, (2–)3–4( 5) mm long, rarely longer, the
proximal ones a little larger than the distal, the distal ones thickened, reflexed, entire,
biglandular at the base, uniglandular at the apex, rarely multiglandular, 20–30–
flowered. Female flowers with pedicels 1–2 mm long, calyx cupular, splitting into
2–3 lobes and becoming flattened as the fruit matures, fulvous tomentellous or
puberulous without, glabrous within, greenish, ovary 1–2-lobate, 1–1.5 mm long,
1–2 mm wide, densely yellowish granulate-glandular, styles 1–2, recurved, united at
the base when 2, 1–2 mm long, adaxially densely papillose, abaxially puberulous,
tinged with red or purple. Female inflorescences racemose to subpaniculate, 2–6
( 7.5) cm long, axis straight, bracts more or less as in the male inflorescences, but
only 1–3-flowered. (Smith 1987; Smith and Carter 1988). The species is an aggres-
sive pioneer and germinates very easily under light conditions (Bussmann and Lange
2000). One of the fastest growing species in Afromontane forests (Bussmann 1999),
and the main species closing gaps after treefall and logging in Ocotea usambarensis
dominated humid lower montane forests, forming successional associations of its
own. Macaranga is dominant in gaps where Ocotea is absent, i.e., has died naturally
or has been logged out. Saplings of Ocotea are successfully completed by
Macaranga, which has a 50% higher growth rate than that of the Camphor
(Bussmann 1996, 2002a, b, c, 2004, 2006) (Figs. 1, 2, 3, and 4).
Macaranga kilimandscharica Pax 679
Fig. 1 Macaranga
kilimanscharica
(Euphorbiaceae), germinating
Macaranga, Mt. Kenya
National Park, Chogoria,
Kenya. (Photo
R.W. Bussmann)
Macaranga kilimandscharica: The root extract is used to treat bilharzia, and cough.
A leaf decoction is drunk for stomach problems (Kokwaro 2009). Macaranga
sp. are used in India for bone injuries (Reang et al. 2016). Macaranga obovata is
used to treat diarrhea in Madagascar (Rakotoarivelo et al. 2015), Macaraga
oblongifolia for malaria and diarrhea (Razafindraibe et al. 2013), and Macaranga
alnifolia for diarrhea (Rabearivony et al. 2015).
Fig. 2 Macaranga kilimanscharica (Euphorbiaceae), Mt. Kenya National Park, Chogoria, Kenya.
Macaranga kilimandscharica: Wood used for boxes and crates (Beentje 1994).
Fig. 3 Macaranga
kilimanscharica
(Euphorbiaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
Fig. 4 Macaranga
kilimanscharica
(Euphorbiaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
References
reference to those of Mount Kenya. J East Afr Nat Hist. 1999;881(2):69–78.
Nat Hist. 2000;891–2:101–11.
Ambasht NK, editors. Modern trends in applied terrestrial ecology. Dordrecht: Kluwer; 2002c.
p. 195–223.
2006;11(1):41–66.
2013;9:73.
Aromatic Plants. 2016;5:2.
1988.
Maerua angolensis DC.
Maerua triphylla A. Rich.
CAPPARACEAE

Grace N. Njoroge
Synonyms
Maerua angolensis DC.: Maerua bukobensis Gilg & Gilg-Ben., Maerua currorii
Hook. f., Maerua emarginata Schinz, Maerua lucida Hochst. ex A. Rich., Maerua
retusa Hochst. ex A. Rich., Maerua senegalensis R.Br., Maerua thomsonii
T. Anderson, Maerua tomentosa Pax
Maerua triphylla A. Rich.: Crateva avicularis Burch. ex DC., Crateva caffra
Burch. ex DC., Maerua dupontii Hemsl., Maerua jasminifolia Gilg & Gilg-Ben.,
Maerua trachycarpa Gilg, Maerua trichocarpa Gilg & Gilg-Ben., Maerua variifolia
Gilg & Gilg-Ben.
Local Names
Maerua angolensis: Sukuma: Msurula (Kokwaro 2009); Borana: Kaikai;

Gabbra: K’alkacca; Kamba: Mukolekya; Luo: Bware; Maa: Ol-Amaloki;
Pokot: Mandas; Samburu: Luliondo, Loitarkini; Somali: Hamaloshi; Tugen:
R. W. Bussmann (*)
G. N. Njoroge

Chebosiewa; Tiriki: Erreng, Esuwat (Beentje 1994); English: Small bead bean;
Swahili: Msingizi, Mukasi, Mukayi, Mkuturu, Mlala-mbuzi.
Maerua triphylla: Kikuyu: Gathandika, Mukuriundu, Mutumburu; Pokot:
Chokotwa, Ohokowa, Mandas; Hehe: Kipegero; Zigua: Milamila; Maa:
Ol-Amalogi; Borana: Kalkaj, Dumao; Giriama: Mushindano; Ilwana: Klaqacha;
Kamba: Mulingula; Kipsigis: Borowet-Abiteta; Luo: Bware; Ndorobo: Lito;
Oromo: Kalkalch-hare; Samburu: Mulingula; Somali: Dumei; Turkana: Ebele
Botany and Ecology
Maerua angolensis DC.: Shrub or small tree up to 8 m tall, young branches

pubescent or glabrescent, becoming densely lenticelled. Leaves simple, lamina
2.5–7 1.3–5.5 cm, lanceolate, broadly elliptic or ovate to broadly ovate, apex
rounded or emarginate, mucronulate, base broadly cuneate or rounded, glabrous or
pubescent on both sides, nerves in 5–6 pairs, slightly prominent on both sides,
petioles up to 3 cm long, often slightly swollen just below the lamina. Inflorescence
of short, terminal, corymbose racemes on the main branches or on short side
branches or flower solitary in the axils of the upper leaves, pedicels up to 2.5 cm
long, glabrous or pubescent, bracts subulate, trifid with the longer middle lobe up to
3 mm. long or similar to the leaves but smaller. Receptacle 8–15 mm long, 3 mm
wide, cylindric, slightly wider at the mouth, glabrous or pubescent outside, with a
fimbriately lobed disk, 2.5 mm long at its margin within. Sepals 1–1.7 0.5–0.7 cm,
narrowly oblong-obovate or elliptic, apex rounded to a shortly apiculate apex,
pubescent on both sides or glabrous on both sides except at the margins.
Androgynophore the same length as the receptacle. Stamens 50, filaments 4 cm
long, anthers 2 0.75 mm, narrowly oblong, arcuate. Ovary on a gynophore up to
4 cm long, narrowly oblong, ovules 80 on 2 placentas. Fruit 2–16 1 cm, narrowly
cylindric, torulose. Seeds from several to numerous, 6 mm in diam., subglobose,
with a smooth, pale brown testa. Erect or scandent shrub or small tree, 5–7.5 m tall.
Twigs glabrous or pubescent. Leaves petiolate, simple or 1–3-foliolate, blade lance-
olate, ovate, elliptic or obovate, 1.5–10 cm long, 0.6–4.5 cm wide, apically obtuse,
acute or rounded, emarginate, usually mucronate, glabrous or pubescent, petiole up
to 3 cm long. Inflorescence a simple or branched corymbose raceme, pedicels
1–2 cm long. Sepals 5–9 mm long, hairy or glabrous outside, with the margin
generally densely white-puberulent, receptacle narrowly cylindrical, 2–5 mm long,
disk with a free rim, usually uneven-lydentate, sometimes subentire. Petals obovate,
3–5 mm long, clawed, whitish or greenish. Androphore about as long as the
receptacle, stamens (10–)15–28, 15–20 mm long. Gynophore 1.5–2.5( 3) cm
long, ovary narrowly cylindrical to ovoid or globose, 1–3 mm long, glabrous or
pubescent, stigma sessile and flattened-capitate. Fruits globose or more usually
ellipsoid or cylindrical, commonly rather clearly constricted between the seeds, up
to 5 cm long, 0.7–1.5 cm in diameter.
Maerua angolensis DC. . . . 687
Maerua triphylla A. Rich.: Erect or scandent shrub or small tree, 5–7.5 m tall. Twigs
glabrous or pubescent. Leaves petiolate, simple or 1–3-foliolate, blade lanceolate,
ovate, elliptic or obovate, 1.5–10 cm long, 0.6–4.5 cm wide, apically obtuse, acute or
rounded, emarginate, usually mucronate, glabrous or pubescent, petiole up to 3 cm
long. Inflorescence a simple or branched corymbose raceme, pedicels 1–2 cm long.
Sepals 5–9 mm long, hairy or glabrous outside, with the margin generally densely
white-puberulent, receptacle narrowly cylindrical, 2–5 mm long, disk with a free
rim, usually unevenly dentate, sometimes subentire. Petals obovate, 3–5 mm long,
clawed, whitish or greenish. Androphore about as long as the receptacle, stamens
(10–)15–28, 15–20 mm long. Gynophore –l15–2.5( 3) cm long, ovary narrowly
cylindrical to ovoid or globose, 1–3 mm long, glabrous or pubescent, stigma sessile
and flattened-capitate. Fruits globose or more usually ellipsoid or cylindrical, com-
monly rather clearly constricted between the seeds, up to 5 cm long, 0.7–1.5 cm in
diameter. Maerua triphylla occurs from Ethiopia and Somalia south to Zimbabwe
and Mozambique. It also occurs in Mayotte, Madagascar and the Seychelles. Besides
tropical Africa, it is also found in peninsular Arabia (Elffers et al. 1964) (Fig. 1).
Fig. 1 Maerua triphylla

(Capparaceae), Bale
Mountaimns National Park,
Maerua angolensis: Roots are used for flu and toothache (Kokwaro 2009), as well
as for treating gonorrhea (Wondimu et al. 2007). The species is also used to treat
wounds (Bussmann et al. 2006).
Maerua triphylla: In East Africa a root infusion is drunk to cure dizziness and
headache and it is also used as an aphrodisiac or to treat venereal diseases. A root
infusion is drunk, fresh roots are chewed and the wound washed with it to treat
snakebites. The Luo people of Kenya drink decoctions and infusions of leaves, bark
and roots as a cure for stomach-ache and diarrhea. Chewing a piece of stem helps to
overcome a dry cough. The ash of burned leaves is taken as an antidote for poison.
The Dorobo people of northern Kenya chew the leaves, mix them with ashes, and
use this mixture as a dressing for boils. Leaves are soaked in water and the water is
applied as an eye lotion (Kokwaro 2009). Also used to treat a variety of sexually
transmitted diseases (Njoroge and Bussmann 2009), and for wound treatment
(Bussmann 2006). The species has anthelminthic activity, also reported for Maerua
edulis and Maerua subcordata (Muthee et al. 2011). Maerua subcordata is also
used to treat bone and joint pain, glandular problems, stomachache, toothache, and
women’s problems (Delbanco et al. 2017).
Maerua nuda is used to treat infected wounds (Randrianarivony et al. 2016, 2017),
and Maertua caffra for gynecological problems (Mohagheghzadeh and Faridi
2006).
Local Food Uses
Maerua angolensis / Maerua triphylla: In Ethiopia the leaves are cooked and eaten
during food-shortage. In Kenya during periods of famine the roots were carefully
boiled to remove toxic substances and prepared as porridge (Teklehaymanot and
Giday 2010).
Maerua angolensis: Used for furniture (Beentje 1994) and as fish poison
(Neuwinger 2004). The same use has been reported for Maerua decumbens.
Maerua triphylla: The leaves and fruits yield a yellow dye. The Maasai and
Samburu people consider the foliage a good fodder, especially for donkeys and
goats (Bussmann et al. 2006; Bussmann 2006). The wood is used for poles, tools,
building, and beehives. The branches are used for firewood, the trunk is used to
produce charcoal (Bussmann 2006). Maerua triphylla is planted especially in rocky
sites for soil reclamation and is often spared when clearing land. Branches and roots
are used to clarify water, but this should be discouraged as the plants could be toxic.
The flowers attract many bees and butterflies (Beentje 1994). Also used for con-
struction of huts and to make containers (Bussmann 2006).
Maerua angolensis DC. . . . 689
References
Elffers J, Graham RA, DeWolf GP. Flora of tropical East Africa: Capparidaceae. London: Crown
2009;82:262–9.
2017;13:1. https://doi.org/10.1186/s13002-017-0147-x.
Mangifera indica L.
ANACARDIACEAE

Grace N. Njoroge
Synonyms
Mangifera indica L.: Mangifera austroyunnanensis H.H. Hu; Mangifera indica var.
armeniaca Bello; Mangifera indica var. intermedia Bello; Mangifera indica
var. leiosperma Bello; Mangifera indica var. macrocarpa Bello; Mangifera indica
var. viridis Bello; Rhus laurina Nutt.
Local Names
Mangifera indica: Kikuyu: Mwiembe (Gachati 1989).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Mangifera indica L.: Large evergreen tree up to 20 m tall, with a dark green,
umbrella-shaped crown. Trunk stout, up to 90 cm in diameter; bark brown,
smoothish, with many thin fissures; thick, becoming darker, rough and scaly or
furrowed; branchlets rather stout, pale green and hairless. Inner bark light brown and
bitter. A whitish latex exudes from cut twigs and a resin from cuts in the trunk.
Leaves alternate, simple, leathery, oblong-lanceolate, 16–30 3–7 cm, on flowering
branches, up to 50 cm on sterile branches, curved upward from the midrib and
sometimes with edges a little wavy. Young leaves red, aging to shiny dark green
above, lighter below, with yellow or white venation; petioles 4.5 cm long, striate and
swollen at the base. Inflorescence 16 cm or more in length, a much-branched panicle
bearing many very small (4 mm) greenish-white or pinkish flowers. Flowers radially
symmetrical, usually have 5 spreading petals, 3–5 mm long, 1–1.5 mm broad,
streaked with red, imbricate, with the median petal prolonged like a crest at the
base, finely hairy and fragrant, partly male and partly bisexual; stalk short; 5 stamens,
1 fertile, the other 4 shorter and sterile, borne in a disc. The flower has a conspicuous
5-lobed disc between the petals and stamens. Calyx yellow-green, very short, deeply
5-lobed; 5 sepals, each 2–2.5 mm long 1–1.5 mm broad, green with whitish
margin, or yellowish-green, hairy outside. Fruit an irregularly egg-shaped and
slightly compressed fleshy drupe, 8–12 (max. 30) cm long, attached at the broadest
end on a pendulous stalk. The skin smooth, greenish-yellow, sometimes tinged with
red. The underlying yellow-orange flesh varies in quality from soft, sweet, juicy and
fiber-free in high-quality selected (clonal) varieties to turpentine flavored and fibrous
in wild seedlings. The single, compressed-ovoid seed is encased in the white fibrous
inner layer of the fruit (Macbride and Weberbauer 1936–1995) (Figs. 1, 2, 3, and 4).
Fig. 1 Mangifera indica (Anacardiaceae), Pusac, Peru. (Photo R.W. Bussmann & N.Y. Paniagua-
Zambrana)
Mangifera indica L. 693
Zambrana)
Zambrana)
Fig. 4 Mangifera indica

(Anacardiaceae), Pusac, Peru.
(Photo R.W. Bussmann &
Fig. 5 Mangifera indica (Anacardiaceae), production of mango jam, Pusac, Peru. (Photo
R.W. Bussmann & N.Y. Paniagua-Zambrana)
Mangifera indica: In Kenya used to treat respiratory disorders (Njoroge and

Bussmann 2006). In Madagascar mango is used to treat dental cavities, hemor-
rhoids, and diarrhea (Rabearivony et al. 2015), as well as evacuation of the placenta,
leucorrhoea and gonorrhea (Razafindraibe et al. 2013). In India for indigestion,
dysentery, cough and cold, as anthelminthic, infection, hypertension, heat stroke
and digestion (Raj et al. 2018), as well as stomachache and diarrhea, especially in
children (Raj et al. 2018). The bark is used to treat fever and malaria; leaves are used
to treat afts, inflammations, mouth infections, strengthen the gums, and for toothache
Mangifera indica L. 695
Fig. 6 Mangifera indica (Anacardiaceae), production of mango jam, Pusac, Peru. (Photo
R.W. Bussmann & N.Y. Paniagua-Zambrana)
(Bussmann et al. 2018), and also boiled for toothache (Debbarma et al. 2017). Dried
leaves are used for bronchitis, colds, and Inflammation (chest) (Bussmann and
Sharon 2006, 2015). The seed extract is used to treat stomachache, diarrhea, and
dysentery in children (Kunwar et al. 2010; Singh et al. 2017, 2019). In Uganda used
as antimalarial (Tabuti 2008).
Local Food Uses
Widely used as food (Mekonnen et al. 2015) (Figs. 5, and 6).
References
978-0-9960231-2-2.
Bussmann RW, Paniagua-Zambrana NY, Romero C, Hart RE. Astonishing diversity - the medicinal
plant markets of Bogotá, Colombia. J Ethnobiol Ethnomed. 2018;14(1):43. https://doi.org/
10.1186/s13002-018-0241-8.
tribes of Tripura, Northeast India. Afr J Tradit Complement Alternat Med. 2017;14(4):156–68.
indigenous uses and pharmacological validity. Med Aromatic Plant Sci Biotechnol. 2010;4
Njoroge GN, Bussmann RW. Traditional management of Ear, Nose and Throat (ENT) diseases in
018-0208-9.
women from Agnalazaha littoral forest (Southeastern Madagascar). J Ethnobiol Ethnomed.
2013;9:73.
itants of Jakholi Block, Rudraprayag district, Western Himalaya, India. J Ethnobiol Ethnomed.
2017;13:1–29. https://doi.org/10.1186/s13002-017-0178-3.
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;18. https://
doi.org/10.32859/era.18.14.1-11.
Markhamia lutea (Benth.) K. Schum.
BIGNONIACEAE

Grace N. Njoroge
Local Names
Markhamia lutea: Kikuyu: Muu, Mho, Muho; Embu: Moo; Luo: Siala; Kamba:
Kyoo; Luhya: Shisimbali, Lusiola; Meru: Mogu, Mungwani; Nandi: Mobet;
Swahili: Mtarawanda; Luganda: Nsambya; Haya: Msambia (Beentje 1994;
Botany and Ecology
Markhamia lutea (Benth.) K. Schum.: Tree (5–)15–21( 24) m tall or sometimes

shrubby 1.2–4.5 m tall, regenerating and coppicing easily. Trunk sometimes divided
from base, sometimes fluted. Bark grey-black or reddish brown, smooth or rough,
ridged. Branches usually densely lenticellate, wood reddish. Leaves up to 35 cm long.
Leaflets (2–)3–6-jugate, elliptic to obovate, 4.5–21 4–9 cm, cuneate to rounded at the
base, acuminate at the apex, glabrous but densely lepidote above, more sparsely so
R. W. Bussmann (*)
G. N. Njoroge

beneath, often with pubescent axillary domatia beneath. Petiole 6–12 cm long, petiolules
up to 5( 10) mm long or absent, pseudostipules, round, 2–3 cm diameter. Flowers
scented, in often narrow terminal panicles up to 20 cm long and wide. Pedicels about 5
( 10) mm long. Bracts triangular, 1–5 mm long, 1 mm wide. Calyx spathaceous, 1.8–
2.6( 3) cm long, 0.7–1.4 cm wide, woolly when young, becoming glabrous, lepidote
and with glands on the margin opposite the split, often very distinctly uncinate at the tip.
Corolla golden yellow with brownish purple or red veins or spots at the throat, tube (2–)
3–4.5 cm long, up to 2.5 cm wide at the throat, limb 2-lipped, 5-lobed, the lobes (1–)
1.5–2.5 cm long and wide, with distinct glands but varying in number. Capsule curved,
35–80( 105) cm long, 1–2 cm wide, compressed, glabrous but lepidote. Seeds 2.5–
3.5 cm long, 6–8 mm wide including the wing. Wooded grassland, submontane and
riverine forest, evergreen rain forest, 1500–1900 m. (Bidgood et al. 2006; Bussmann
2002). One of the easiest Afromontane forest species to germinate, both in light and dark
conditions, and even after long storage (Bussmann and Lange 2000).
Markhamia lutea: Young shoots and leaves are chewed to treat throat problems.
Bark is chewed for toothache, the bark decoction drunk for syphilis. A fresh leaf
infusion is drunk, and the leaves applied to treat snakebites (Kokwaro 2009). The
leaves and wood are burnt, and the smoke inhaled to remedy depression and other
mood disorders (Mohagheghzadeh and Faridi 2006).
Markhamia lutea: The wood is hard and termite-resistant, for construction, poles,
utensils (Beentje 1994), and sometimes used as firewood (Kiefer and Bussmann 2008).
References
Bidgood S, Vollesen K, Verdcourt B. Flora of tropical East Africa: Bignoniaceae & Cobaeaceae.
appendices 1-7 @ www.naturekenya.org/JournalEANH.htm
Nat Hist. 2000;891(2):101–11.
Maytenus senegalensis (Lam.) Exell
Maytenus undata (Thunb.) Blakelock
CELASTRACEAE

Grace N. Njoroge
Synonyms
Maytenus senegalensis (Lam.) Exell: Catha grossulariae Tul., Catha montana (Roth
ex Roemer & Schultes) G. Don, Catha senegalensis (Lam.) G. Don, Celastrus
coriaceus Guill. & Perr., Celastrus europaeus Boiss., Celastrus montanus Roth ex
Roemer & Schults, Celastrus saharae Batt., Celastrus senegalensis Lam., Celastrus
senegalensis ver inermis A. Rich., Gymnosporia baumii Loes., Gymnosporia
crenulata Engl., Gymnosporia dinteri Loes., Gymnosporia eremocusa Loes.,
Gymnosporia grossulariae (Tul.) Loes., Gymnosporia montana (Roth ex Roemer &
Schultes) Benth., Gymnosporia paniculata Baker, Gymnosporia senegalensis (Lam.)
Loes., Gymnosporia senegalensis fo. chartacea Loes., Gymnosporia senegalensis
fo. coriacea (Guill. & Perr.) Loes., Gymnosporia senegalensis fo. macrocarpa
Loes., Gymnosporia senegalensis var. inermis (A. Rich.) Loes., Maytenus baumii
(Loes.) Exell & Mendonça.
Maytenus undata (Thunb.) Blakelock: Catha fasciculata Tul., Celastrus
fasciculatus (Tul.) Boivin ex O. Hoffm., Celastrus fasciculatus Drake, Celastrus
lancifolius Thonn., Celastrus laurofolius A. Rich., Celastrus luteolus Delile,
R. W. Bussmann (*)
G. N. Njoroge

Celastrus undatus Thunb., Celastrus zeyneri Sond., Gymnosporia albata (N.E. Br.)
Sm., Gymnosporia deflexa Sprague, Gymnosporia fasciculata (Tul.) Loes.,
Gymnosporia goetzeana Loes., Gymnosporia ilicina (Burch. ex Loudon) Loes.,
Gymnosporia lancifolia (Thonn.) Loes., Gymnosporia luteola (Delile)
Loes., Gymnosporia malinensis Schnell, Gymnosporia rehmannii Szyszyl.,
Gymnosporia undata (Thunb.) Szyszyl., Gymnosporia zeyheri (Sond.) Szyszyl.,
Maytenus fasciculata (Tul.) Loes., Maytenus lancifolia (Thonn.) Loes.
Local Names
Maytenus senegalensis: Kikuyu: Muthuthi, Muenyuke; Swahili: Mdunga-Ndewe,

Mnyonyao; Digo: Mtsokolangongo; Embu: Mvenyuke; Giriama: Ndugu-Tundu;
Ilwana: Baagsa; Kamba: Kivinda-Ngiti; Kipsigis: Kaguruet; Luo: Mathari,
Nyandema; Maa: Ol-Lalmoronaili; Nandi: Shibe, Ngarawi; Oromo: Koboch;
Pokot: Akwichanian; Samburu: Laimurunyai; Sebei: Chemetio, Chemetyo,
Munyandetap; Somali: Mandaruk; Tugen: Aburuwa; Turkana: Ekaburu; Sukuma:
Luwenje, Mchomafisi; Nguru: Mrambangoma; Luhya: Muruli; Haya: Mweza
Maytenus undata: Kikuyu: Muthuthioi, Muthuthi; Elgeyo: Cheptoya; Giriama:
Muriakitui; Kipsigis: Kalpuguet; Maa: Olgerosha; Ndorobo: Cheptuyetumo,
Eburua; Pokot: Kerepurwa; Samburu: Itui; Sebei: Kwonget, Chemoiwa; Tugen:
Katuya, Chepkendu; Turkana: Emuthi; Shambaa: Kabere (Beentje 1994; Gachati
1989; Kokwaro 2009).
Botany and Ecology
Maytenus senegalensis (Lam.) Exell: Shrub or tree or rarely a shrublet, (0.15–)

1–9( 15) m high, unarmed or with spines up to 4 cm long, axillary or terminating
short axillary branches, glabrous, without latex, branches unlined, flattened, reddish
purple to reddish brown, often densely glaucous, and sometimes with numerous pale
indistinct lenticels at first, becoming terete and eventually grey-brown. Sepals
5, subequal, oblong-lanceolate to ovate-triangular or subcircular, 0.3–1.2 mm long,
obtuse to rounded, with margin ciliolate. Cymes 1–3.5 cm long, slender, dichasial at
first, becoming monochasial, solitary and axillary or 1–6 on short axillary shoots,
with peduncle 1–16 mm long, pedicels 0.7–4( 6) mm long, articulated in the lower
1/3( 1/2), flowers scented, 3–60 or more in each cyme, 2–6 mm in diameter,
unisexual (plants dioecious or rarely monoecious). Male flowers: stamens 5, shorter
than the petals, with filaments 0.5–1 mm long, slender, arising below disc, disc
relatively broad, flat or slightly concave, shallowly 5–10-lobed, pistillode small,
globose. Petals 5, white or greenish white to pale yellow, sometimes tinged pink,
elliptic to oblong-elliptic or oblanceolate, 1–3.5 mm long, with margin finely
ciliolate. Capsule pink to deep red, globose to pyriform, 2–6 mm long, coriaceous,
smooth. Female flowers: staminodes 5, shorter than the ovary, disc as in ♂, ovary
2( 3)-locular, globose, scarcely immersed in the disc, style as long as the ovary,
Maytenus senegalensis (Lam.) Exell. . . 701
Fig. 1 Maytenus arbutifolia

(Celastraceae), Bale

shorter than the petals, 2( 3)-fid at the apex, with small stigmas. Seeds 1–2, dark
reddish brown, glossy, with a fleshy smooth white aril obliquely covering the lower
1/3–2/3. Leaves fasciculate or not, petiolate, lamina pale green, usually glaucous,
sometimes with reddish midrib, often mottled (at least when dry), oblong or rarely
ovate to obovate or oblanceolate or oblong-elliptic or rarely subcircular,
2–11.6 0.4–5.7( 6.3) cm, rounded or retuse to obtuse or apiculate at the apex,
with margin densely regularly obtusely serrulate or serrulate-crenulate to subentire
or very rarely entire, cuneate to attenuate at the base, coriaceous or subcoriaceous,
with lateral nerves and relatively lax reticulate venation slightly prominent or not
prominent, petiole 3–13( 20) mm long. (Robson 1994). Found in drier, often
Juniperus procera dominated, Afromonatne forests (Bussmann 2002) (Figs. 1, 2,
3, 4, 5, 6, 7, and 8).

Maytenus undata (Thunb.) Blakelock: Shrub or tree 1.5–10( 12) m high, much
branched, sometimes scandent, unarmed, glabrous, without latex, very variable,
branches markedly lined or angled when young, usually becoming terete. Seeds
orange-brown, a quarter to almost completely enclosed by a thin, white to orange,
oblique aril. Capsule white or pinkish red to yellow or orange, obovoid to sub-
globose or 3-gonous, 4–7( 10) mm long, with lobes rounded or keeled, smooth, not
fleshy. Ovary 3-locular, subglobose, 1/3 immersed in the disc, style elongate or
almost absent, stigma shortly 3-lobed. Disc 5–10-lobed or subentire, flat or with thin
free upturned margin or rarely somewhat convex. Stamens 5, shorter than the petals,
with filaments 0.7–2 mm long, slender, not or scarcely flattened, united with the base
of the disc. Petals 5, white to cream or pale yellow, oblong or elliptic to obovate or
oblanceolate, 1.5–3( 5) mm long, sessile or shortly clawed, with margin entire or
eroded-denticulate, often crisped. Sepals 5, unequal, ovate to semicircular, rounded,
0.7–1.5( 2) mm long, sometimes tinged red, with margin entire to eroded-
denticulate. Leaves not fasciculate, petiolate, lamina pale green or silver-green to
dark green and sometimes glossy above, concolorous or sometimes paler beneath or
whitish-glaucous beneath or on both surfaces, ovate or oblong to elliptic or
Fig. 4 Maytenus
cf. senegalensis
Fig. 5 Maytenus
cf. senegalensis
Fig. 6 Maytenus
cf. senegalensis
oblanceolate or circular or oblate, (1.1–)3.2–13( 20) (0.9–)1.2–7.5( 11) cm,

acute to rounded at the apex, with margin glandular-denticulate to spinulose-dentate
or subentire, rarely reflexed or indurated, cuneate-angustate or rarely truncate to
subcordate at the base, coriaceous to chartaceous, with 6–9 lateral nerves and
reticulate venation varying in density and prominence, petiole 2–8( 15) mm long.
Pedicels 2–9( 15) mm long, articulated at the base or in the lower third, flowers
fragrant, 2–30 in each fascicle or rarely solitary, 4–8 mm in diameter, bisexual
(Robson 1994). Found in drier, often Juniperus procera dominated, Afromonatne
forests (Bussmann 2002) (Figs. 9, 10, 11, 12, and 13).
Maytenus senegalensis: Roots are used to treat snakebites, rheumatism, tuberculo-

sis, diarrhea, and fever. Crushed leaves are soaked in water and the extract is used as
eye-drops. The concoction is also drunk for high blood pressure and applied
externally to wounds and sores. The root decoction is used for anthrax, kidney
Fig. 7 Maytenus
cf. senegalensis
Fig. 8 Maytenus cf. undata



problems, and gastro-intestinal affections (Kokwaro 2009). Also used for gyneco-
logical problems (Muthee et al. 2011). Also used to treat fever (Wondimu et al.
2007), malaria (Kunwar et al. 2010), (Njoroge and Bussmann 2006a), and tonsillitis
(Njoroge and Bussmann 2006b).
Maytenus undata: The bark decoction is used as tonic (Beentje 1994; Kokwaro
2009). The root decoction is taken for syphilis and urinary diseases (Kokwaro 2009).
Maytenus gracilipes is employed for epilepsy and headache (Yineger et al. 2008).
Maytenus arbutifolia for liver problems (Yineger et al. 2008).


Local Food Uses
Maytenus heterophylla: The fruits are eaten, and the leaves boiled for strengthening
soups (Bussmann 2006).
Maytenus senegalensis: Wood used for utensils (Beentje 1994), and to make
charcoal (Giday et al. 2003).
Maytenus undata: Wood used for construction and utensils (Beentje 1994).

Maytenus serrata is used as fishing poison (Neuwinger 2004). Maytenus arbutifolia

is used to treat evil eye (Yineger et al. 2008). Maytenus heterophylla is used to
smoke calabashes, and browsed by livestock (Bussmann 2006).
References
Kunwar RM, Shrestha KP, Bussmann RW. Traditional herbal medicine in far-West Nepal: a
Robson N. Flora of tropical East Africa: Celastraceae. Rotterdam: Balkema; 1994.

2008;26:132–53.
Moringa oleifera Lam.
MORINGACEAE

Grace N. Njoroge
Synonyms
Moringa oleifera Lam.: Anoma moringa (L.) Lour.; Guilandia moringa L.; Hyper-
anthera decandra Willd.; Hyperanthera moringa (L.) Vahl); Moringa amara Durin;
Moringa domestica Buch.-Ham.; Moringa edulis Medik.; Moringa erecta Salisb.;
Moringa moringa (L.) Millsp.; Moringa nux-eben Desf.; Moringa octogona Stokes;
Moringa ovalifolia Dinter & Berger; Moringa polygona DC.; Moringa
pterygosperma Gaertn.; Moringa sylvestris Buch.-Ham.; Moringa zeylanica
Burman.
Local Names
Moringa oleifera: English: Drumstick tree, Ben oil tree, Horseradish tree; Swahili:
Mzunze, Mronge, Mlonge (Beentje 1994).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Moringa oleifera Lam.: A fast-growing, deciduous tree. It can reach a height of

10–12 m and the trunk can reach a diameter of 45 cm. The bark has a whitish-grey
color and is surrounded by thick cork. Young shoots have purplish or greenish-white,
hairy bark. The tree has an open crown of drooping, fragile branches, and the leaves
build up feathery foliage of tripinnate leaves. The flowers are fragrant and bisexual,
surrounded by five unequal, thinly veined, yellowish-white petals. The flowers are
about 1.0–1.5 cm long and 2.0 cm wide. They grow on slender, hairy stalks in
spreading or drooping later flower clusters which have a length of 10–25 cm. The
flowers are fragrant and bisexual, surrounded by five unequal, thinly veined,
yellowish-white petals. The flowers are about 1.0–1.5 cm long and 2.0 cm wide.
Moringa oleifera is indigenous in northern India and Pakistan. It has been introduced
throughout the tropics and subtropics and has become naturalized in many African
countries (Figs. 1, 2, and 3).
Fig. 1 Moringa oleifera

(Moringaceae), preparando
Moringa leaves for food,
Mahaboboboka,
Southwestern Madagascar.

Moringa leaves for food,
Mahaboboboka,
Moringa oleifera Lam. 713

flowers, Mahaboboboka,
Moringa oleifera: Almost all parts have traditional medicinal applications, as an

anodyne, anthelmintic, antispasmodic, and disinfectant (bactericidal, fungicidal) are
widespread. Used to treat malaria (Randrianarivony et al. 2017; Tabuti 2008). Also
to alleviate blood pressure issues, gastroenteritis, cold and cough, body pain, clear-
ing stool, tumors, leucoderma, liver disorder, snake bite, piles, cough, stomach
worm, diarrhea, and dysentery (Bekalo et al. 2009; Raj et al. 2018). The fruits are
used in Nepal for liver disorders (Kunwar et al. 2008, 2009, 2013, 2016; Kunwar and
Bussmann 2009). Roots are purgative. Leaves are a galactogogue. Gum is useful in
dysentery. The plant is antispasmodic, expectorant, cardiotonic and diuretic. The seeds
are used for bronchitis, asthma, and leprosy. Bark and fruit are taken for epilepsy,
urinary tract infection, and rheumatism. The plant is astringent, aphrodisiac, carmina-
tive, and stomachic (Kunwar et al. 2009). In India, the species is used as abortifacient,
anti-fertility, appetizer, for baldness, caries dental, catarrh, cholera, delivery, asthma,
ear complaints, eczema, epilepsy, fever, headache, intestinal worms, liver complaints,
madness, menstrual complaints, paralysis, pneumonia, rheumatism, scabies, scorpion
bite, scurvy, snake bite, sores, spleen enlargement, stomach ache, syphilis, tonic,
typhoid, urine complaints, and wounds (Verma et al. 2007).
Moringa stenophylla is used for flu (Teklehaymanot and Giday 2010).
Local Food Uses
Moringa oleifera: Whereas in Asia the fruits are the most important part of Moringa
oleifera, the leaves are preferred in Africa. The leaves are eaten as a salad, cooked,
and in soups and sauces (Balemie and Kebebew 2006; Debbarma et al. 2017). In the
Mascarene Islands, it is known as “brède mouroungue” or “brède médaille.” Flowers
are sometimes eaten as a vegetable, added to sauces or used to make tea. In Sudan the
flowers are made into a paste by crushing and then fried. The young fruits are eaten
as a vegetable (“drumsticks” or “bâtons mouroungue”), older fruits are added to
sauces. In West Africa some health projects fight malnutrition quite successfully by
promoting a number of measures including the use of leaf powder in the diet of
children and pregnant and lactating women. The tuberous root cores can be a
substitute for horseradish (Armoracia rusticana Gaertn., B. Mey. & Scherb.)
(Beentje 1994). The fried seeds are eaten in Nigeria and are said to taste like
groundnuts. The seeds are added locally to sauces for their bitter taste. The seed
oil, known as “Ben oil” or “Behen oil,” can be used for cooking. The bark exudes a
white to reddish gum (“Ben gum” or:Moringa gum”) with the properties of traga-
canth (Astragalus) oil, which is used for tanning, in calico printing, and is sometimes
added to sauces to make them thicker.
The whole or pounded seeds have long been used to purify water in Sudan, and this
practice is promoted elsewhere in Africa. The seed cake, a residue from oil extrac-
tion, can also be used for water purification. The seed oil is used in hair-dressing, as a
lubricant, and in the perfume industry as a base for fragrant volatile compounds in
perfumes. “Moringa acid oil,” consisting of fatty acids from the seed oil, is used as a
lubricant and to make soap. Moringa oleifera is used for living fences, in alley
cropping, and as a source of nectar for bees. The leaves are eaten by livestock,
especially goats, camels, and donkeys. The seed cake is considered unsuitable as
animal feed because of the high content of alkaloids and saponins and is mainly used
as fertilizer. The soft wood burns smoke-free and yields a blue dye. In India its pulp
has been used to make paper. For cuts and wounds of domestic animals, and as snake
repellent (Raj et al. 2018). Used also as aphrodisiac (Delbanco et al. 2017).
References
doi.org/10.1186/1746-4269-5-26.
tribes of Tripura, Northeast India. Afr J Tradit Complement Alternat Med. 2017;14(4):156–68.
Biodiversität, Naturausstattung im Himalaya, vol. III. Erfurt: Verein der Freunde & Förderer
des Naturkundemuseums; 2009. p. 475–89.
Moringa oleifera Lam. 715
Kunwar RM, Chowdhary CL, Bussmann RW. Diversity, utilization and management of medicinal
macology of medicinal plants in far-West Nepal. Ethnobot Res Appl. 2009;7:5–28.
and management in far-West Nepal. J Ethnobiol Ethnomed. 2013;9:24.
Western Nepal. PLoS One. 2016;11:e0167812. https://doi.org/10.1371/journal.pone.0167812.
018-0208-9.
2017;13:19. https://doi.org/10.1186/s13002-017-0147-x.
Myrianthus holstii Engl.
URTICACEAE

Grace N. Njoroge
Synonyms
Myrianthus holstii Engl.: Myrianthus mildbraedii Peter
Local Names
Myrianthus holstii: Kikuyu/Meru: Mutuya; Kipsigis: Saoune; Shambaa:

Mkonde; Luganda: Mugunga (Beentje 1994; Gachati 1989; Kokwaro 2009);
English: Giant yellow mulberry, Bush pineapple, Corkwood.
Botany and Ecology
Myrianthus holstii Engl.: Tree up to 20 m tall. Lamina 25 25 to 60 60 cm,

3–7( 8)–fid or -parted down to the petiole; segments sessile or stoutly petiolulate,
margin rather regularly serrate-dentate to subentire; upper surface hirtellous to
R. W. Bussmann (*)
G. N. Njoroge

puberulous on the main veins, lower surface brownish hirtellous to subsericeous on

the main veins; petiole 7–35 cm long, brownish hirtellous to subsericeous; stipules
1.5–4 cm long, brown to golden-yellow sericeous. Staminate inflorescences
4–15 cm in diameter; peduncle 3–13.5 cm long; glomerules of flowers spike-
like, 4–6 mm in diameter. Pistillate inflorescences 1–2 cm., in fruit 5–8 cm in
diameter; peduncle 1–5 cm long; flowers 20–40; apex of the perianth broadly conical;
endocarp-body 1.2 0.8 cm. Staminate inflorescences 4–15 cm in diameter;
peduncle 3–13.5 cm long; glomerules of flowers spike-like, 4–6 mm in diameter.
Pistillate inflorescences 1–2 cm, in fruit 5–8 cm in diameter; peduncle 1–5 cm long;
flowers 20–40; apex of the perianth broadly conical; endocarp-body 1.2 0.8 cm.
Mountain-forest, sometimes at edges, in regrowth and along rivers; 900–2100 m.
Common in afromontane forets (Bussmann 2002a, b).
Myrianthus holstii: In Congo chopped leaves are eaten raw with salt to treat heart
troubles, pregnancy complications, dysmenorrhea, and incipient hernia. A bark
decoction is drunk to treat malaria, fever, and cough. In DR Congo, a leaf decoction
is also used as an anticough medicine. In Tanzania, an infusion of the leaves is taken
to improve lactation in women. Sap from young leaves or terminal buds is applied
topically to treat toothache or to the chest against bronchitis and to the throat against
laryngitis or sore throat. The bark and root decoction is used as cough syrup. The leaf
decoction serves as galactogogue (Kokwaro 2009).
Local Food Uses
Myrianthus holstii: The fruit is edible (Beentje 1994).
Myrianthus holstii: The wood is soft, yellowish white, perishable, and difficult to
work, but is used for fencing and occasionally as a general-purpose wood. The wood
is also suitable for paper making. Its ashes are used in soap making in Guinea.
References
Myrianthus holstii Engl. 719
Myrsine africana L.
PRIMULACEAE

Grace N. Njoroge
Synonyms
Myrsine africana L.: Myrsine africana var. acuminata C.Y. Wu & C. Chen; Myrsine
africana var. bifaria (Wall.) Franch.; Myrsine africana var glandulosa J.M. Zhang;
Myrsine africana var. retusa A. DC.; Myrsine bifaria Wall.; Myrsine mirophylla
Hayata; Myrsine potama D. Don; Myrsine vaccinifolia Hayata; Rhamnus myrtillus
Lév.
Local Names
Myrsine africana: Kikuyu: Mugaita; Iraqwi: Maheli; Maa: Ol-Segetit; Marakwet:

Segatet; Kipsigis: Segetyet, Osegeteti; Pokot: Parapara; Samburu: Segeteta; Tugen:
Segetetwa; Turkana: Ekaiyi; Kamba: Muketa-munini (Beentje 1994; Gachati 1989;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Myrsine africana L.: Evergreen undershrub, shrub or tree from 1–6 m tall, often or
perhaps always subdioecious, the male plants producing no ovaries or fruit, the
flowers on female plants containing stamens which may or may not be functional;
bark grey or white with lighter streaks, shredding; branches brown or purplish,
densely minutely crisped hairy especially on the younger wood, older wood with
longitudinal ribs. Leaves alternate, crowded; blades ovate to lanceolate, rhomboid,
obovate or orbicular, 4.5–20 mm long, 3–11.5 mm wide, acute to acuminate,
cuneate, usually serrate in upper half, occasionally entire, coriaceous, glossy and
rugulose above, smooth, punctate beneath, glabrous except occasionally a few hairs
at base of midrib on upper surface, glands faintly pellucid, orange; nerves (2–)3–5
( 7) on either side of midrib; petiole 1–3 mm long, glabrous to densely glandular-
hairy. Inflorescence consisting of 1–5-flowered axillary clusters on young wood.
Flowers male or female, small, 4-merous, pendent, glabrous, gland-dotted; pedicels
up to 1.5 mm long, glabrous. Sepals united at base or rarely free, 1 mm long,
margins finely erose in male flowers, entire in female flowers. Corolla greenish white
or pink, campanulate with entire lobes to funnel-shaped with ciliate lobes, 2 mm.
long; lobes as long as tube in male flowers, and half as long in length in female
flowers, 2 mm long. Stamens exserted, subsessile or with filaments up to 1.5 mm
long, united into a collar which is adnate to the corolla-tube; anthers crimson to blue-
violet, ovate, apiculate, 1–2.5 mm long. Ovary ovoid; placenta stalked, ovoid,
apiculate; ovules few, uniseriate; style exserted to half its length, slender, 1–3 mm
long; stigma cream, discoid, fimbriate. Fruit an edible berry, pale green, red, purple,
finally black, globose with persistent style and calyx, up to 5 mm in diameter,
smooth except for raised gland-dots, glabrous, 1-seeded. Seeds globose. Upland
forest edges, open wooded grassland, stony hillsides in grassland and pasture,
sometimes riverine, on sandy soil over basement complex or on volcanic soil or
lava; 1200–3600 m. (Halliday 1984). Myrsine africana is a characteristic species in
dry, Juniperus procera dominated Afromontane forests, and is especially character-
istic for the regeneration stages after fire (Bussmann 2002a, b, c, 2004, 2006;
Bussmann and Beck 1995) (Figs. 1 and 2).
Myrsine africana: Fruits are eaten raw as anthelminthic, especially for hookworms
and roundworms. Also dried and ground as remedy for chest-pain and stiff joints.
Has purgative effect (Kokwaro 2009). The seed extract is rubbed on pimples and
skin rashes (Sher et al. 2016). Used to treat backache, chest pains, cold, deworming,
diarrhea, general body health, heartburn, high blood pressure, stomachache, as
anthelminthic, purgative and for constipation (Muthee et al. 2011; Nankaya et al.
2019, 2020; Yineger and Yewhalaw 2007). The plant is also used to treat gonorrhea
(Muthee et al. 2011), other sexually transmitted diseases and malaria (Gafna et al.
2017), bone and joint pain, kidney diseases and women’s health (Delbanco et al.
Myrsine africana L. 723
Fig. 1 Myrsine africana

(Primulaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
Fig. 2 Myrsine africana

(Primulaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)
2017), and as anti-inflammatory (Desta 1995). Traditionally also used to treat polio
and tuberculosis (Bussmann 2006).
Myrsine africana: The leaves are used to wrap dough for cooking (Bussmann et al.
2011; Luizza et al. 2013). The seeds are crushed, and the oil is used to grease the
baking plate before baking (Bussmann et al. 2011).
Myrsine melanophloeos is used for construction and firewood (Bussmann et al.
2011).
References
p. 195–223.
Bussmann RW, Beck E. Regeneration- and succession processes in the Cedar-Forests Juniperion
procerae of Mount Kenya. Ecotropica. 1995;1:79–84.
Desta B. Ethiopian traditional herbal drugs. Part I: Studies on the toxicity and therapeutic activity of
Halliday P. Flora of tropical East Africa: Myrsinaceae. Rotterdam: Balkema; 1984.
Res Appl. 2013;11:315–39.
Myrsine africana L. 725
Plants (Basel, Switzerland). 2020;9:44. https://doi.org/10.3390/plants9010044.
Neoboutonia macrocalyx Pax
EUPHORBIACEAE

Grace N. Njoroge
Local Names
Neoboutonia macrocalyx: Kikuyu: Mutundu; Cherangani: Kibatwa; Kamba:

Mwega; Kipsigis: Sabaitet; Meru: Mutuntuki; Sebei: Chebakwa; Taveta: Muhaa;
Taita: Dundu, Shawa (Beentje 1994; Gachati 1989).
Botany and Ecology
Neoboutonia macrocalyx Pax: A slender open-crowned tree up to 25 m tall, but

more commonly 10–15 m. Twigs at first densely scurfily stellate-pubescent, some of
the hairs occasionally with one ray much longer than the rest, later becoming
sparingly so. Bark fairly smooth, pale greyish brown or grey-green. Female inflo-
rescences up to 40 cm long and 30 cm wide, stiffly erect, less branched than the ♂,
but otherwise similar, bracts 4 mm long, 1-flowered. Male flowers: pedicels up to
5 mm long, glabrous, sepals 2( 3), ovate or elliptic-ovate, 2.5–3 mm long, 1.5 mm
R. W. Bussmann (*)
G. N. Njoroge

wide, acute, sparingly pubescent to subglabrous without, glabrous within, pale

yellowish green, disc-glands 10, cylindric-conic, 0.3 mm long, stamens 30–40,
2–3 mm long, filaments greenish cream, anthers 1.3 mm long, cream-yellow, with
a small white apical gland. Male inflorescences up to 50 cm long and 30 cm wide,
much-branched, often leafy at the base, axes sparingly to evenly stellate-pubescent,
bracts ovate, 1.5–2 mm long, acute, sparingly pubescent, (1–)2–3( 4)-flowered.
Stipules triangular-ovate, 1 cm long, 1 cm wide, subacute, cordate, densely scurfily
stellate-pubescent. Petioles (2–)10–20( 30) cm long, indumented as the twigs, leaf-
blade broadly ovate to suborbicular, 6–35 cm long and wide, acutely or subacutely
and very shortly acuminate or rounded-obtuse, deeply cordate, with 2 somewhat
irregular stipitate glands at the insertion of the petiole, usually entire or subentire,
occasionally denticulate, chartaceous, 7( 9)-nerved from the base, lateral nerves
5–8 pairs, nerves, veins, and reticulations not prominent above, prominent beneath,
sparingly scurfily stellate-pubescent above and only on the nerves, veins and retic-
ulations beneath, minutely gland-dotted above and/or beneath, dark green above,
paler beneath. Seeds ellipsoid, 7 mm long, 5 mm wide, brown-mottled, with an
adpressed bilobed caruncle. Female flowers: pedicels stout, 2–5 mm long, densely
pubescent, calyx-lobes 5( 6), unequal, lanceolate to linear-lanceolate, 5–7 mm
long, 1.5–2 mm wide, accrescent to 9 mm long in fruit, thick, reflexed, evenly
velutinous without and within, 3-nerved without, dull green, disc shallowly 5-lobed,
ovary trilobate, 1.5 mm long, 2 mm diameter, densely stellate-pubescent, styles
2 mm long, pubescent without, glabrous within. Fruits trilobate, 1–1.2 cm long,
1.2–1.3 cm diameter, evenly scurfily stellate-pubescent. (Smith 1987; Smith and
Fig. 1 Neoboutonia macrocalyx (Euphorbiaceae) Mt. Kenya National Park, Chogoria, Kenya.
Neoboutonia macrocalyx Pax 729
Fig. 2 Neoboutonia
macrocalyx (Euphorbiaceae)
R.W. Bussmann)
Carter 1988). Neoboutonia macrocalyx is a pioneer species of lower Afromontane

forests, characterized by very fast growth (Bussmann 1999), that fills especially gaps
created by logging of valuable timber species like Ocotea usambaransis (Bussmann
1996). Only found in the lowermost parts of the forest belt (Bussmann 2002a, b, c,
2004). Germinating well under light conditions, as one might expect from a pioneer
species (Bussmann and Lange 2000) (Figs. 1 and 2).
Wood used to make paper (Beentje 1994).

References
p. 195–223.
Nat Hist. 2000;891(2):101–11.
1988. p. 409–597. ISBN 90-6191-338-1.
Nicotiana rustica L.
Nicotiana tabacum L.
SOLANACEAE

Grace N. Njoroge
Synonyms
Nicotiana rustica L.: Nicotiana pavonii Dunal

Nicotiana tabacum L.: Nicotiana chinensis Fisch. ex Lehm.; Nicotiana mexicana
Schltdl.; Nicotiana mexicana var. rubriflora Dunal; Nicotiana pilosa Dunal
Local Names
Nicotiana tabacum: Kikuyu: Kibaki, Mbaki (Beentje 1994; Gachati 1989;

Kokwaro 2009)
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Nicotiana rustica L.: Annual. Stem over 1 m tall, herbaceous, glandular-pilose

branching almost from the base. Leaves all petiolate, soft, slightly fleshy, ovate,
usually obtuse, rarely acute, with cordate base. Flowers in racemose panicle on short
pedicels. Calyx broadly campanulate, 6–10 mm long, with broad triangular lobes.
Corolla 1.5–2 cm long, with broad whitish tube and greenish yellow, flat, narrow
limb with broad, ovate-triangular obtuse lobes. Capsule subglobose, with numerous
extremely small, brown, ovoid seeds. Flowering from July to September. Cultivated
in fields and kitchen gardens. South America (native in Peru); cultivated in all
countries with tropical and subtropical climate and the more southern regions of
the Temperate Zone (Shishkin and Boborov 1955).
Nicotiana tabacum L.: Annual. Stem 0.75–1.5 m tall, herbaceous, simple or with
few branches, with glandular pubescence. Leaves alternate, entire; basal leaves
narrowly elliptical, decurrent along stem; upper leaves narrowly lanceolate, long-
acuminate, sessile or short petiolate. Flowers in cymose panicles on short pedicels.
Calyx 1–2 cm long, campanulate, with acute narrowly triangular teeth. Corolla
5–6 cm long, red or pink, rarely white, infundibuliform, with long and broad
spreading limb, lobes broadly triangular, with sharply pointed tip. Capsule ellipsoi-
dal-ovoid, apiculate. Seeds ovoid-reniform, very small. Flowering July–September,
Fig. 1 Nicotiana tabacum

(Solanaceae), garden,
Chilchos, Peru. (Photo R.W:
Zambrana)
Nicotiana rustica L. . . . 733

(Solanaceae), garden,
Chilchos, Peru. (Photo R.W:
Zambrana)
fruiting from September. Cultivated in fields and kitchen gardens (Shishkin and
Boborov 1955) (Figs. 1, 2, 3, 4, 5, 6, and 7).
Phytochemistry
Alkaloids (nicotine, nicoteine, nicotelline, nicotinin, betanin, i-amigdalin,

pyrrolidine, n-methyl pyrrolidine), fatty acids (palmitic, butyric, and linoleic acids,
stearic) (Sokolov 1990).
Nicotiana rustica: The leaves are used for arthritis, bronchitis, sore throat, tonsillitis,
and wounds. Used as toothache remedy, and for respiratory tract diseases
(Mohageghzadeh and Faridi 2006).

(Solanaceae), garden, Adjara,
Georgia. (Photo R.W:
Zambrana)

(Solanaceae), drying for use,
Osh, Kyrgyzstan. (Photo R.W:
Zambrana)
Nicotiana tabacum: The leaves are used to treat arthritis, bronchitis, digestive
system problems, sinusitis, and skin problems. An infusion is sometimes used to
treat stomach infections (Delbanco et al. 2017). In Madagascar, the extract serves to

(Solanaceae), died smoking
tobacco, Pshavi, Georgia.
(Photo R.W: Bussmann &

tobacco, Pshavi, Georgia.
(Photo R.W: Bussmann &

tobacco, market, Dinsho,
Zambrana)
stop nosebleeds (Razafindraibe et al. 2013), and to treat bilharzia (Rabearivony et al.
2015). Also used for general gynecological disorders, narcotic, toothache remedy,
general skin diseases, and the treatment of ear problems (Mohageghzadeh and Faridi
2006).
Both species traditionally grown and widely used as smoking tobacco and snuff.
Nicotiana rustica: The leaves are used for protection, smoking, and snuff. The stems
are also used as masticant.
Nicotiana tabacum: The leaves are used for protection, smoking, and snuff
(Mekonnen et al. 2015). The whole plant is used as natural pesticide (Bussmann
et al. 2014, 2016a, b; 2017, 2018; Bussmann 2017; Wanzala 2017). Sometimes used
as fishing poison (Neuwinger 2004). The leaf extract serves to treat blackleg in
livestock (Yineger et al. 2007), and to remove leeches (Wondimu et al. 2007).
Sometimes used for respiratory problems and cough in cattle (Giday et al. 2009).
Locally grown and sold as stimulant (Giday et al. 2009).
References
Bussmann RW, editor. Ethnobotany of the Caucasus. Springer International Publishing XXVII;
2017. 746p. ISBN 978-3-319-49411-1.
2014;12:237–313.
(Republic of Georgia), Caucasus. Med Aromat Plants. 2016a;5:266. https://doi.org/10.4172/
2167-0412.1000266.
Khutsishvili M, Batsatsashvili K, Hart RE. Plant and fungal use in Tusheti, Khevsureti and
Pshavi, Sakartvelo (Republic of Georgia), Caucasus. Acta Soc Bot Pol. 2016b;86(2):3517.
https://doi.org/10.5586/asbp.3517.
Bussmann RW, Batsatsashvili K, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S,
Tchelidze D, Paniagua Zambrana NY. Ethnobotany of the Caucasus – Georgia. In: Bussmann
RW, editor. Ethnobotany of the Caucasus. Cham: Springer International Publishing; 2017.
Delbanco AS, Burgess ND, Cuni SA. Medicinal plant trade in Northern Kenya: economic impor-
tance, uses, and origin. Econ Bot. 2017;711:13–31.

2013;9:73.
Shishkin BK, Boborov EG. Flora of the USSR, volume 22: Solanaceae and Scrophulariaceae.
Leningrad: Akademia Nauk; 1955 (English 1993). 745 p.
Sokolov PD, editor. Plant resources of the USSR: flowering plants, their chemical composition, use.
Volume 5. Families of Caprifoliaceae – Plantaginaceae. Leningrad: Akademia Nauk; 1990,
Livestock Industry with Focus on Acari Ticks. Evid Based Complement Alternat Med 2017;
Article ID 8647919. 33 p. https://doi.org/10.1155/2017/8647919.
Nuxia congesta R. Br. ex Fresen.
LOGANIACEAE

Grace N. Njoroge
Synonyms
Nuxia congesta R. Br. ex Fresen.: Lachnopylis angolensis (Gilg) Philipson,

Lachnopylis annobonensis Mildbr., Lachnopylis breviflora (S. Moore) C.A. Sm.,
Lachnopylis compacta C.A. Sm., Lachnopylis congesta (R. Br. ex Fresen.)
C.A. Sm., Lachnopylis emarginata (Sond.) C.A. Sm., Lachnopylis flocculosa
C.A. Sm., Lachnopylis goetzeana (Gilg) Greenway, Lachnopylis guineensis Hutch.
& M.B. Moss, Lachnopylis heterotricha C.A. Sm., Lachnopylis mannii (Gilg)
Hutch. & M.B. Moss, Lachnopylis montana C.A. Sm., Lachnopylis odorata (Gilg)
Greenway, Lachnopylis platyphylla (Gilg) Greenway, Lachnopylis pubescens
(Sond.) C.A. Sm., Lachnopylis sambesina (Gilg) C.A. Sm., Lachnopylis saxatilis
C.A. Sm., Lachnopylis schistotricha C.A. Sm., Lachnopylis speciosa C.A. Sm.,
Lachnopylis ternifolia Hochst., Lachnopylis thomensis Philipson, Lachnopylis
R. W. Bussmann (*)
G. N. Njoroge

tomentosa (Sond.) C.A. Sm., Lachnopylis viscidulosa C.A. Sm., Lachnopylis

viscosa (Gibbs) C.A. Sm., Nuxia angolensis Gilg, Nuxia breviflora S. Moore,
Nuxia dekindtiana Gilg, Nuxia emarginata Sond., Nuxia gilletii De Wild., Nuxia
goetzeana Gilg, Nuxia keniensis T.C.E. Fr., Nuxia latifolia T.C.E. Fr., Nuxia mannii
Gilg, Nuxia odorata Gilg, Nuxia platyphylla Gilg, Nuxia pubescens Sond., Nuxia
rupicola Gilg, Nuxia sambesina Gilg, Nuxia siebenlistii Gilg, Nuxia tomentosa
Sond., Nuxia viscosa Gibbs
Local Names
Nuxia congesta: Kikuyu: Muchorowe, Mucorui; Kamba: Mukalaliki,

Mukalamu’u; Luhya: Lubambo, Inoyna; Maa: Ol-burin; Marakwet / Nandi /
Kipsigis / Sebei: Chorua; Pokot: Selta; Sebei: Morosuet; Tugen: Kerruwa;
Turkana: Akwanga, Enonochorie; Taita: Mgaraso, Mora (Beentje 1994; Gachati
1989; Kokwaro 2009); English: Common wild elder, Brittle wood
Botany and Ecology
Nuxia congesta R. Br. ex Fresen.: Evergreen or sometimes in Ethiopia deciduous

tree or shrub, 2–25 m high, often gnarled. Trunk in larger trees fluted and
irregularly ridged, 5–100(200) cm in diameter, bark pale grey-brown, fissured.
Branchlets glabrous to densely pubescent, not lenticellate, with 6(8) raised lines
and often sulcate when dry. Leaves ternate (occasionally 4-nate), petiolate, petiole
glabrous to densely pubescent, 3–20 mm long, lamina coriaceous or sub-
coriaceous, very variable in shape and size, elliptic, narrowly elliptic, obovate,
nearly rhomboid, or suborbicular, 1–3(5) times as long as wide, (1)2–15 (0.3)
1.2–7.5 cm, acuminate to emarginate at the apex, cuneate or less often rounded at
the base or decurrent into the petiole, entire or sometimes more or less distinctly
serrate-dentate, or crenate, glabrous to subtomentose with stellate hairs on both
sides. Leaves on sucker shoots (mostly in the shade) darker green, herbaceous,
serrate, and more hairy in tropical countries, but thicker in southern Africa.
Inflorescence variable in shape and size, seemingly umbellate to paniculate,
congested or rather lax, 3–15 3–15 cm, 3–7 times branched. Peduncle and
branches hairy like the branchlets. Pedicels short or obsolete. Flowers solitary or
three together, fragrant. Calyx green, often viscid, 3–8 1.5–2.2 mm, minutely to
manifestly pubescent with glandular and mostly also ordinary hairs outside, inside
sericeous, lobes variable in size, mostly about as long as wide or wider, 0.5–
2 1–1.5 mm, acute or less often obtuse, entire, valvate, often two coherent or
partially united. Corolla white, with recurved pubescence on the lobes which may
be reduced to scattered hairs or is rarely absent and with minute glandular hairs at
the base of the lobes and the apex of the tube (the latter sometimes on the middle
of the tube or on nearly the complete tube too), inside with curved hairs in the
Nuxia congesta R. Br. ex Fresen. 741
Fig. 1 Nuxia congesta (Loganiaceae), Mt. Kenya National Park, Naro Moru Route, Kenya. (Photo
R.W. Bussmann)
throat and sometimes also at the apices of the lobes and with minute glandular
hairs in the tube except for its glabrous base (hairs in throat or hairs in tube are
occasionally absent, although never both simultaneously), tube slightly shorter or
sometimes – when calyx very short only – longer than the calyx, lobes oblong,
2–3 as long as wide, 2–5 1–2 mm, obtuse or subacute at the apex. Stamens:
anthers glabrous, 1.2–2 mm long. Pistil (4.5–)7.5–13 mm. long, ovary ovoid or
ellipsoid, slightly laterally compressed, 1.5–2.5 1.2–2 1–1.5 mm,
appressedly hirto-pubescent, glabrous at the base. Capsule 0·5–1.5 mm. longer
than the calyx, appressedly hirto-pubescent. Nuxia congesta is the most wide-
spread species of the genus, occurring from Guinea eastward to Ethiopia and
Somalia and southward through East Africa to South Africa and Swaziland. It is
also found in southern Arabia (Bruce and Lewis 1960). Important species in
broad-leaved Afromontane forests (Bussmann 2002a, b, c, 2004, 2006a)
(Figs. 1, 2, 3, 4, and 5).
Nuxia congesta: In traditional medicine in Kenya, bark decoctions are taken to treat
indigestion and menstrual problems, and as an expectorant. Bark and leaves are also
chewed against indigestion, whereas ash of leafy twigs is taken in water against
Fig. 2 Nuxia congesta

(Loganiaceae), Mt. Kenya
Route, Kenya. (Photo
R.W. Bussmann)

R.W. Bussmann)
diarrhea. In Uganda the bark is applied against toothache and impotence (Kokwaro
2009). Used to treat common cold and flu (Gafna et al. 2017). For the treatment of
rheumatism and scabies (Yineger et al. 2008).
Nuxia congesta R. Br. ex Fresen. 743
Fig. 4 Nuxia congesta (Loganiaceae), Mt. Kenya National Park, Naro Moru Route, Kenya. (Photo
R.W. Bussmann)

R.W. Bussmann)
Nuxia congesta: The wood of Nuxia congesta is mainly used for poles for con-
struction, fence posts, and carpentry. It is suitable for furniture. It is also used as
firewood and for charcoal production, whereas branches are used as firesticks
(Beentje 1994). The abundance of fragrant flowers makes Nuxia congesta an
attractive ornamental. It is suitable as boundary, barrier, and support tree. The trees
prevent erosion on rocky sites and provide shade or shelter. The leaves are browsed
by livestock. Smoke from the leaves is applied to the udder to treat mastitis in cattle.
In Kenya and Tanzania, Nuxia congesta is an important honey plant. Mostly for
construction, firewood, as nectar tree for honey production, and fences. The leaves
can be eaten by livestock (Bussmann 2006b; Bussmann et al. 2011).
References
Ambasht NK, editors. Modern trends in applied terrestrial ecology. New York: Kluwer; 2002b.
p. 195–223.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002c; 91 1/2:27-79,
appendices 1–7 @. http://www.naturekenya.org/JournalEANH.htm.
2006a;111:41–66.
2008;26:132–53.
Ocimum gratissimum Lam.
Ocimum kilimandscharicum Guerke
LAMIACEAE

Grace N. Njoroge
Synonyms
Ocimum gratissimum Lam.: Ocimum guineense Schumach. & Thonn, Ocimum

suave Willd., Ocimum urticifolium Roth, Ocimum viride Willd.
Local Names
Ocimum gratissimum: Kikuyu: Makuri, Mukuri; Borana: Manjabbi, Anchabbi;

Digo: Vumba manga; Giriama: Vamba manga; Kamba: Mukandu; Luo:
Oluluruecha; Maa: Ol-emoran; Marakwet: Jemasat, Chesimia; Samburu:
Lemurranm; Taita: Mrumbawassi; Turkana: Loguru, Ichoke; Banyoro: Omwenti;
Lugbara: Mujaja (Beentje 1994; Gachati 1989; Kokwaro 2009).
Ocimum kilimandscharicum: Kikuyu: Makuri, Mugio, Mukuri; Luhya:
Esilokha, Mwonyi; Kamba: Mwenye; Embu: Makole; Meru: Makuru; Pokot:
R. W. Bussmann (*)
G. N. Njoroge

Supko; Turkana: Kameteber; Shambaa: Mzubngwa; Luo: Bwar, Okita (Beentje

1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Ocimum gratissimum Lam.: Subshrubs, erect, to 2 m, glabrous to softly pubescent

on the younger branches. Leaves ovate or elliptic, acute, crenate to subentire, the
bases rounded and weakly attenuate or obtuse, up to 7 cm long and 4.5 cm wide,
resin-dotted, glabrous above and sparsely pubescent on the veins beneath; petioles to
ca 4.5 cm long, pubescent. Inflorescences verticils, each ca 6-flowered, 0.5–1 cm
distant, the false racemes 10–15 cm long, the bracts deltoid, 2–3 mm long, pubes-
cent. Flowers with pedicels ca 1–2 mm long; bracteoles not apparent; calyx 3–4 mm
long, membranaceous, resin-dotted, pubescent on the veins and at the base, naked
within, the upper lip entire, 3-veined, lateral margins decurrent only on upper 1/3 of
tube, the lower lip 5–7-veined; corolla tubular, white and purple, ca 2.5 mm long
overall; stamens exserted, lower filament pair with conspicuously pubescent append-
ages at the base, the anthers ca 0.6 mm long; ovary 4-lobed, gynobase weakly lobed,
the style bifid near the-tip, the branches equal. Nutlets (3–)4, spherical or slightly
elongated, ca 1.2 mm long, the pericarp warty (Paton et al. 2009). In the undergrowth
of drier Afromontane forests (Bussmann 2002) (Figs. 1, 2, 3, and 4).
Fig. 1 Ocimum
sp. (Lamiaceae), Bale
Demaro, Ethiopia. (Photo R.
W: Bussmann)
Ocimum gratissimum Lam. . . . 747
Fig. 2 Ocimum
W: Bussmann)
Fig. 3 Ocimum
W: Bussmann)
Ocimum kilimandscharicum Guercke: Woody herb or shrub, 20–200 cm tall.

Leaves ovate-elliptic, with cuneate base, 1.5–6 0.6–3 cm, pubescent. Margins
minutely serrate. Flowers white to pale violet, sometimes purplish, corolla 4–7 mm
Fig. 4 Ocimum
W: Bussmann)
long, inflorescences 6–25 cm long (Beentje 1994). In the undergrowth of drier

Afromontane forests (Bussmann 2002).
Ocimum gratissimum: Leaves are boiled, and the decoction drunk for constipation
(Bekalo et al. 2009; Kokwaro 2009). Leaves are also used to treat cough (Beentje
1994), nose congestion, flu, and colds (Nankaya et al. 2019; Njoroge et al. 2004;
Tabuti 2008), as well as otitis media (Njoroge and Bussmann 2006a, b). Widely used
as medicine in the Himalayas, as diuretic, for asthma and rheumatism (Kunwar et al.
2012, 2015, 2018). In Madagascar used to expel the placenta after birth
(Randrianarivony et al. 2016a, b). Also used for severe headache, wounds as
abortifacient, colds, low calcium, dental problems, digestion, chest complaints,
diarrhea, vomiting, and as antiseptic (Rabearivony et al. 2015; Razafindraibe et al.
2013; Randriamiharisoa et al. 2015), and for rheumatism (Yineger et al. 2008).
Ocimum kilimandscharicum: Leaves are boiled, and the steam is inhaled for colds.
The leaves can be crushed and sniffed for colds and coughs, and the infusion applied
to sore eyes. The Luo use it as bath for measles. The leaf decoction serves for
diarrhea (Kokwaro 2009). The leaves are used for cough (Beentje 1994).
Ocimum canum is used for eye problems (Flatie et al. 2009). Ocimum lamiifolium
serves for diarrhea and cough (Gedif and Hahn 2003), and fever (Giday et al. 2007,
2009a, 2010; Mekonnen et al. 2010; Teklehaymanot et al. 2007; Teklehaymanot
2009; Wondimu et al. 2007), as well as cutaneous leishmaniasis (Giday et al. 2009b).
Ocimum urticifolium is used for fever (Giday et al. 2003), toothache, oral thrush, ear
infections, mouth infections in children, and tongue infections (Giday et al. 2010;
Mekonnen et al. 2010). Ocimum sanctum is one of the most widely used medicinal
plants on the Indian subcontinent f or cough, colds, gastroenteritis, neck-pain, and
cancer (Raj et al. 2018).
Local Food Uses
Ocimum gratissimum: Leaves used as tea (Beentje 1994). Ocimum obovatum is

used as spice in Ethiopia, to make “Berbere” (Bussmann 2011; Luizza et al. 2013).
Ocimum gratissimum: The leaf infusion is applied to the eyes of livestock infected
with Thelazia sp. (eyeworm) (Kokwaro 2009).
The leaves of Ocimum suave are smoked and serve as fodder (Bussmann 2006),
and as acaricide (Wanzala 2017). Ocimum lamiifolium is used for diarrhea and
hepatitis in livestock (Yineger et al. 2007), and to treat evil eye (Teklehaymanot et al.
2007).
References
doi.org/10.1186/1746-4269-5-26.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002; 91 1/2:27-79,
appendices 1–7 @. http://www.naturekenya.org/JournalEANH.htm
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Gedif T, Hahn H-J. The use of medicinal plants in self-care in rural central Ethiopia.
group of Ethiopia: an ethnobotanical investigation. J Ethnobiol Ethnomed. 2009a;5:34. https://
doi.org/10.1186/1746-4269-5-34.
ethnobotanical study. J Ethnopharmacol. 2009b;124:513–21.
species in far-West Nepal – a valuable resource being wasted. J Mountain Sci. 2012;9:589–600.
P. Cross-cultural comparison of plant use knowledge in Baitadi and Darchula Districts, Nepal
Res Appl. 2013;11:315–39.
Mekonnen T, Urga K, Engidawork E. Evaluation of the diuretic and analgesic activities of the
rhizomes of Rumex abyssinicus Jacq in mice. J Ethnopharmacol. 2010;127:433–9.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Sustainability of the loita maasai childrens’
ethnomedicinal knowledge. Sustain 2019;11:5530. https://doi.org/10.3390/su11195530.
Paton AJ, Bramley G, Ryding O, Polhill RM, Harvey YB, Iwarson M, Willis F, Phillipson PB,
Balkwill K, Lukhoba CW, Otieno DF, Harvey RM. Flora of tropical East Africa: Lamiaceae.
org/10.1186/s13002-015-0046.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016a. ISBN 978-0-9960231-4-6.

2013;9:73.
Raj AJ, Biswakarma B, Pala NA, Shukla G, Vineeta V, Kumar M, Chakravarty S, Bussmann RW.
Indigenous uses of ethno-medicinal plants among forest dependent communities of Northern
Bengal, India. J Ethnobiol Ethnomed. 2018;141:8. https://doi.org/10.1186/s13002-018-0208-9.
10.1186/1746-4269-4-11.
Ocotea usambarensis Engl.
LAURACEAE

Grace N. Njoroge
Local Names
Ocotea usambarensis: Kikuyu: Muthaiti, Musili; Meru: Muurta; Embu: Muthura;

Taita: Manyoda, Muganga, Mulongo. Kivumba; Chagga: Miseri; Swahili: Mkulo,
Mukongo (Beentje 1994; Gachati 1989; Kokwaro 2009); English: Camphorwood
Botany and Ecology
Ocotea usambarensis Engl.: A medium to large evergreen tree 15–30(40) m tall

with a dense crown and a stout straight bole up to 10 m tall, buttressed at the base.
Bark purplish-grey, rough when old and flaking in small rounded or rectangular
scales; wood aromatic with a camphor-like scent when freshly cut, slash a light
reddish-brown, becoming darker with exposure. Young branches and twigs slender,
R. W. Bussmann (*)
G. N. Njoroge

angular and finely longitudinally striate towards the ends, more or less densely
fulvous-pubescent or tomentose, sometimes glabrescent. Leaves aromatic with a
camphor-like scent, membranous to coriaceous, discolorous drying brownish above
and silvery-white or greyish on the lower surface; petiole 0.5–2.0 cm long, tomen-
tose to glabrescent; lamina (2.5)4.5–14(17) (2)2.5–6(7.5) cm, ovate to elliptic,
acuminate or sometimes acute or obtuse at the apex, rounded to truncate rarely
subcordate or cuneate at the base, entire; upper surface glossy dark green and more
or less puberulous especially on the midrib and nerves, or glabrous; lower surface
silvery-white pruinose and more or less puberulous, at least on the midrib and
nerves, rarely glabrescent; lateral nerves in 6–8(9) pairs, slightly raised above,
prominent beneath; tertiary nerves net-veined, more or less inconspicuous above.
Inflorescence a panicle of cymes in the axils of subterminal leaves, panicles 3–13 cm
long, more or less lax, few to many-flowered, densely yellowish-brown pubescent;
peduncle 2–8 cm long; bracts minute, ovate to ovate-lanceolate, soon falling.
Flowers small, yellowish, densely pubescent; pedicels 1–4 mm long; receptacle
1 mm long, broadly obconical, glabrous within; tepals 2–3 mm long, elliptic to
ovate-elliptic, obtuse to rounded at the apex, pubescent within. Stamens 9 in
3 whorls, with staminodes forming a fourth (innermost) whorl; filaments of the
first and second (the outer) whorls short and broad, flattened, their anthers 1.0–
1.5 mm long, elliptic-rectangular, obtuse at the apex and dehiscing introrsely;
filaments of the third whorl more or less square, pilose, with 2 shortly-stalked
globose to ovate glands, the anthers oblong, a little longer than the others, truncate
or emarginate, dehiscing extrorsely; staminodes (in the innermost whorl) c. 1 mm
long, ovate-triangular, sessile or subsessile. Ovary 1.0–1.5 mm long, ovoid, gla-
brous; style short thick glabrous; stigma subpeltate. Fruit orange-brownish, up to
10 6 mm, ellipsoid, the lower one third enclosed in the accrescent more or less
fleshy receptacle, becoming slim, almost cylindric (except the summit). In interme-
diate to montane evergreen wet forest; 900–2400 m ( 3000 m on Kilimanjaro)
(Verdcourt 1996).
The dioecious Ocotea usambarensis flowers mostly between early March and late
April, at the beginning of the long rains. A second flowering can be observed after
the start of the short rains in November. In most cases, however, only few trees can
be found flowering during these periods, whereas every 5–7 years mass-flowering
can be found. Most of the produced fruits are attacked by gall-insects when still
unripe or are eaten by birds. The few remaining seeds are only viable for a few days,
even under the most favorable conditions like high humidity and radiation. There-
fore, a regeneration of Ocotea by seedlings is very rare (Bussmann 2002a, b, c, 2004,
2006).
In natural Ocotea forests, only very old Ocotea trees of approximately 300–
600 years old (stem diameter 1.2–2.5 m) are found. Very few scattered young trees
(less than 100 years old) exist generally. Seedlings of Ocotea were found only in a
few places, obviously due to the problems described for the seed development and
germination of Ocotea.
Notwithstanding the uniform age structure, numerous Ocotea saplings, originat-
ing from old roots were observed. Seedlings and young specimens of the other tree
Ocotea usambarensis Engl. 755
40 40
m m
30 30 30 30
m m
20 20 20 20
10 10 10 10
0 10 20 30 40 50 60 m 0 10 20 30 40 50 60 m
10 10 10 10
0 10 20 30 40 50 60 m 0 10 20 30 40 50 60 m
5 Logging of Ocotea leads to large No regeneration of Ocotea. Dying

gaps with mass germination of of overaged secondary forest leads
Macaranga kilimandscharica or again to mass germination of
6 pioneer species. Development of
Neoboutonia macrocalyx.
a secondary forest cycle.
40 40
1 m m
3
40 40
4
m m 30 30
30 30
Dying of overaged 20 20
20
Macaranga trees, no
20
regeneration of the 10 10
10 10
pioneer species in
shade of young 0 10 20 30 40 50 60 m
0 10 20 30 40 50 60 m
Ocotea trees and
reestablishment of 10 10
primary forest
10 10
0 10 20 30 40 50 60 m
0 10 20 30 40 50 60 m
2
Secondary forest with high proportion
Camphor forest with Dying of overaged of Macaranga kilimandscharica in gaps
high proportion of Ocotea trees. and slow regeneration of Ocotea
equally old Ocotea seedlings in the protective shade of the
usambarensis trees pioneer species.
2
1 40
m
40
m
3
30 30 30 30
m m
20 20 Dying of overaged 20 20
Macaranga trees, no
10 10 regeneration of the 10 10
pioneer species in
0 10 20 30 40 50 60 m shade of young 0 10 20 30 40 50 60 m
Ocotea trees and

10 10
reestablishment of 10 10
primary forest
0 10 20 30 40 50 60 m 0 10 20 30 40 50 60 m
4
Aningeria adolfi-friederici Anthocleista zambesiaca Apodytes dimidiata Argomuellera macrophylla
Bersama abyssinica Blighia unijugata Canthium oligocarpum Casearia battscombei
Cassipourea malosana Celtis gomphophylla Chrysophyllum gorgunosanum Croton macrostachyus
Cordia africana Ehretia cymosa Ekebergia capensis Fagara macrophylla

Garcinia volkensii Lasianthus kilimandscharicus Lovoa swynnertonii Macaranga kilimandscharica
Myrianthus holstii Neoboutonia macrocalyx Ocotea usambarensis Olea capensis ssp. hochstetteri
Podocarpus latifolius Psychotria orophila Strombosia scheffleri Syzygium guineense
Tabernaemontana stapfiana Teclea nobilis Teclea trichocarpa Trichilia dregeana
Uvariodendron anisatum Vitex keniensis Xymalos monospora
Fig. 1 Ocotea usambarensis (Lauraceae), regeneration cycle of Ocotea dominated Afromontane

forests (R.W. Bussmann)
species characteristic of lower Afromontane forests can be found everywhere,

indicating the exceptive character of Ocotea itself, while the other species follow
normal regeneration patterns and therefore form a kind of mosaic structure. The
species is essentially impossible to germinate and store, as seeds have a very short
lifespan (Bussmann and Lange 2000). In addition, saplings are the slowest growing
of any species observed in Afgromontane forests (Bussmann 1999) (Fig. 1).
The equally old populations of Ocotea usambarensis trees in the Ocotetea
usambarensis die over an extended period of time (1), thus creating a mosaic of
gaps in which Macaranga kilimandscharica starts growing as secondary species (2).
This leads to a Euphorbeacee forest often dominated by Macaranga
kilimandscharica on small scale which does not change the structure of the forest
to a significant extent. Regeneration of Ocotea from seedlings is negligible but
numerous saplings from the roots of a fallen tree start growing not any longer
suppressed by the mother tree (3). Macaranga kilimandscharica seems to play an
important role as shade tree because young Ocotea saplings apparently do not
tolerate full sun. When the relatively short-living Macaranga trees break down,
the meanwhile well-established young Ocotea trees can close the gaps (4) and by
shading prevent further germination or establishment of Macaranga
kilimandscharica (5). Again, Ocotea stands of a more or less uniform age result in
which Ocotea trees generated from the roots frequently form large rings (6). Other
species fill the centers of these rings.
The removal of Ocotea usambarensis on a large-scale leads to numerous big gaps
in a very short time, especially as many additional trees are killed during felling,
removal, and transport of Ocotea (Bussmann 1996). Even worse, the current logging
Fig. 2 Ocotea usambarensis (Lauraceae), large Ocotea usambarensis to the right, Mt. Kenya
National Park, Ocotea forests on Chogoria Route. (Photo R.W. Bussmann)
Fig. 3 Ocotea usambarensis (Lauraceae), large felled Ocotea usambarensis, Mt. Kenya National
Park, Ocotea forests on Chogoria Route. (Photo R.W. Bussmann)
Fig. 4 Ocotea usambarensis

(Lauraceae), Ocotea
usambarensis saplings arising
from old root, Mt. Kenya
National Park, Ocotea forests
on Chogoria Route. (Photo
R.W. Bussmann)
practice often uses uprooting of the Ocotea trees and thus prevents the usual
propagation by root saplings. The gaps are soon closed by Macaranga
kilimandscharica and at higher altitudes by Neoboutonia macrocalyx, and totally
new vegetation types arise. Since Ocotea does not regenerate in these man-made
“gaps” after breakdown of the pioneer trees, these secondary Euphorbiacee forests
regenerate in a cycle of their own. The possibilities for a regeneration to primary
forest are unknown, and the regeneration of the mentioned secondary forests into the
original primary associations of the Ocotea forests would at least require a complete
stop of logging around the “gaps,” thus enabling the removed species to intrograde
from the margins. However, even then regeneration would require a very long time
period. As an alternative, replanting of Ocotea and other primary forest species on a
large scale appears possible and presumably would enable a faster regeneration
(Figs. 2, 3, 4, 5, and 6).

(Lauraceae), Ocotea
R.W. Bussmann)

(Lauraceae), Ocotea
R.W. Bussmann)
Ocotea usambarensis: The pounded bark of roots, bole and branches is applied to
swellings, boils and wounds. A bark decoction is given to treat whooping cough and
measles. Bark powder is taken against stomach-ache. Roots steeped in water are
taken to treat malaria and back pain (Kokwaro 2009). Ocotea bark is used to treat
back-pain in women (Njoroge and Bussmann 2009), and also for the treatment of
malaria (Njoroge and Bussmann 2006a), for cough, and neck and throat problems
(Njoroge and Bussmann 2006b). Ocotea bullata is sometimes used as analgesic
Ocotea usambarensis: The wood, often traded as “camphor,” is valued for joinery,
paneling, poles for building, doors, window frames, shutters, furniture, cabinet work,
vehicle bodies, sliced veneer, and plywood. It is used for flooring of local houses and
for implements. The wood is suitable for construction, ship building, boxes, crates,
vats, matches, and pulpwood (Beentje 1994). It is less suitable for draining boards
and kitchen utensils because of the camphor-like smell. It is also used as firewood
and for charcoal production. Sometimes planted as ornamental.
References
Bussmann RW. Growth rates of important east African montane forest trees, with particular
reference to those of Mount Kenya. J East Afr Nat Hist. 1999;881–2:69–78.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi. J East Afr Nat Hist 2002a;91 1/2:27–79.
p. 195–223.
2006;111:41–66.
Nat Hist. 2000;891–2:101–11.
2009;82:262–9.
Verdcourt B. Flora of tropical East Africa: Lauraceae. Rotterdam: Balkema; 1996. 19 p. ISBN
90-6191-367-5.
Oldeania alpina (K. Schum.) Stapleton
POACEAE

Grace N. Njoroge
Synonyms
Oldeania alpina (K. Schum.) Stapleton: Arundinaria alpina K. Schum.;

Arundinaria fischeri K. Schum.; Arundinaria tolange K. Schum.; Oxytenanthera
ruwenzorensis Chiov.; Sinarundinaria alpina (K. Schum.) C.C. Chao & Renvoize;
Yushania alpina (K. Schum.) W.C.Lin
Local Names
Oldeania alpina: Kikuyu: Murangi; Swahili: Mianzi, Mwanzi; Kamba: Muangi;

Maa: Ol-diani; Kipsigit/Marakwet: Tegaat, Terga; Nandi/Ndorobo: Tegat; Sebei:
Tegendet; Taita: Mrangi (Beentje 1994; Gachati 1989; Kokwaro 2009); English:
African alpine bamboo, mountain bamboo; French: Bambou creux.
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Oldeania alpina (K. Schum.) Stapleton: Perennial, caespitose. Rhizomes short,

pachymorph. Culms erect, 200–1950 cm long, 50–125 mm diameter, woody.
Culm-internodes terete, thick-walled, mid-green. Lateral branches dendroid.
Branch complement three, or several, in an irregular line, with subequal
branches. Culm-sheaths glabrous, or hispid, with red hairs, auriculate, setose
on shoulders. Culm-sheath blade linear, acute. Leaf-sheath oral hairs setose.
Ligule an eciliate membrane. Leaf-blade base with a brief petiole-like connection
to sheath. Leaf-blades deciduous at the ligule, lanceolate, 5–20 cm long,
6–15 mm wide. Leaf-blade venation with distinct cross veins. Leaf-blade surface
glabrous. Leaf-blade apex attenuate, filiform. Inflorescence a panicle. Panicle
open, elliptic, dense, or loose, 5–15 cm long. Spikelets solitary. Fertile spikelets
pedicelled. Pedicels 1–3 mm long, glabrous. Spikelets comprising 5–10 fertile
florets, with diminished florets at the apex. Spikelets linear, laterally compressed,
15–40 mm long, 3–4 mm wide, breaking up at maturity, disarticulating below
each fertile floret. Rhachilla internodes eventually visible between lemmas.
Floret callus pubescent. Spikelets comprising 5–10 fertile florets, with dimin-
ished florets at the apex. Spikelets linear, laterally compressed, 15–40 mm long,
3–4 mm wide, breaking up at maturity, disarticulating below each fertile floret.
Rhachilla internodes eventually visible between lemmas. Floret callus pubescent.
Glumes persistent, similar, shorter than spikelet, similar to fertile lemma in
texture. Lower glume ovate, 4–8 mm long, 0.8 length of upper glume, herba-
ceous, without keels, 5–9 -veined. Lower glume apex acuminate. Upper glume
elliptic, 4–8 mm long, 0.75 length of adjacent fertile lemma, herbaceous, without
keels, 5–9-veined. Upper glume apex acuminate. Fertile lemma ovate, 7–10 mm
long, herbaceous, without keel, 7–9-veined. Lemma lateral veins with cross-
veins. Lemma surface glabrous, or puberulous. Lemma apex acute. Palea 1 length
of lemma, 2-veined. Palea keels ciliolate. Apical sterile florets resembling fertile
though underdeveloped. Lodicules 3, obovate, membranous, ciliate. Anthers 3.
Stigmas 2, or 3 (rarely). Ovary glabrous. Caryopsis with adherent pericarp,
fusiform, apex unappendaged. Embryo 0.2 length of caryopsis. Hilum linear.
Africa: west-central tropical, northeast tropical, east tropical, and southern trop-
ical. African alpine bamboo occurs in scattered populations on mountains from
southern Sudan and Ethiopia southwards to Malawi. An interval of 2000 km
separates its occurrence in western Cameroon from those in eastern Africa. It is
frequently planted, e.g., in Ethiopia and Zimbabwe. In Afromontane forests
especially in the bamboo zone and in disturbed areas where animals rest
(Bussmann 1996, 2002a, b, c, 2004, 2006a, b) (Fig. 1).
Sambucus africana is an important species in the regeneration cycle of East
African Bamboo. The dense forests dominated by the African Bamboo, Oldeania
africana (in the following in this chapter we use the much better used synonym
Sinarundinaria alpina), which are found on various East African mountains, form a
clear-cut class (Sinarundinarietea alpinae Bussmann 1994, 1997, 1999a, b, 2002a,
b, c; Bussmann and Beck 1995, 1999; Bussmann and Lange 1999; Bytebier and
Oldeania alpina (K. Schum.) Stapleton 763
Fig. 1 Sambucus africana (Caprifoliaceae) Mt. Kenya National Park, Naro Moru route. (Photo
R. W. Bussmann)
Bussmann 2000) of their own. Sinarundinaria alpina (Poaceae), Impatiens hoehnelii

(Balsamiaceae), and Pseudocarum eminii (Umbelliferae) as well as Selaginella
kraussiana (Selaginellaceae), Cyperus dereilema, Anthriscus sylvestris
(Umbelliferae), and Cardamine obliqua (Cruciferae) are the most characterizing
species of this forest type.
soils, namely, humic Andosols. On Mount Kenya, the Sinarundinarietea alpinae are
and a stem diameter of up to 10 cm. New vegetative shoots are sprouting from old
4500–7000 culms produced per hectare require longer periods of favorable climate.
die, occurs in areas of about one to several hundred acres in more or less regular

(Poaceae), sapling arising
route. (Photo R. W.
Bussmann)

route. (Photo R. W.
Bussmann)
intervals of at least 40 years (Wimbush 1945). On Mount Elgon, Clayton (1970)

observed a flowering cycle of only 15 years.
Agnew (1985), referring to a long-term knowledge of the vegetation of the
1. Pioneer: Regeneration from sections of old rhizomes and rhizome fragments and
occasionally from fallen culms, lasting about 3–5 years during which dense
ever been reported from stands of Sinarundinaria (Figs. 2 and 3)
2. Building: During this period, the growing bamboo exceeds the average plant
getting momentum. The pH of the soil starts decreasing (Figs. 4, 5, and 6).

(Poaceae), young regrowing
bamboo, Mt. Kenya National
Park, Naro Moru route. (Photo
R. W. Bussmann)
3. Mature: No further increase in culm height takes place during this stage.
The culms show even distribution and the soil pH is decreasing further
(Figs. 7, 8, and 9).
4. Flowering and 5. Sambucus Stage: Within 2–3 years after mass-flowering the
bamboo therefore could be indicating the sites of frequent flowering (Figs. 10,
11, 12, 13, 14, and 15).
Kenya (Bussmann 1994; Bussmann and Beck 1995; Bussmann and Lange 1999),
Mt. Nyiro in Northern Kenya (Bussmann 2002a, b, c; Bytebier and Bussmann
2000), and the Harenna Escarpment in the Ethiopian Bale Mountains (Bussmann
1997). However, the vegetation showed considerable floristic differences on wet and
drier sites of bamboo forests, even if the areas compared were in the same stage of
the cycle.

R. W. Bussmann)
In the North of Mt. Kenya, young bamboo was apparently invading parts of the
Juniperion procerae with open undergrowth as a new habitat, forming dense, more or
less extended thickets. To clarify whether this represented a new colonization, very
young sprouts were excavated in several places, especially when forming outposts
far from young bamboo thickets. All samples indicated that the new shoots arose
from old rhizome fragments. Therefore, the areas which seemed to be newly
colonized, proved to be regenerating old bamboo stands, where Sinarundinaria
alpina had died many years ago, and very small vegetative bamboo sprouts had
been part of the heavily grazed pasture, so that the local people did not remember
bamboo in these locations (Mathenge pers. comm. 1993). Since no old culms were
found, they must have been decomposed a long time ago, as dead culms normally
stay for many years especially in the drier areas. Therefore, the time gap between two
subsequent bamboo cycles appears to be very long in this area, i.e., regeneration
seems to be very slow.

R. W. Bussmann)
Fig. 7 Sinarundinaria alpina (Poaceae), mixed bamboo – young re-growing bamboo stands,
Mt. Kenya National Park, Naro Moru route. (Photo R. W. Bussmann)

(Poaceae), mature bamboo,
Naro Moru route. (Photo
R. W. Bussmann)
– The region is fairly dry, i.e., Sinarundinaria alpina is at the limit of its distribution
– The Sirimon area in the Northwest has a very high population of herbivores,
In Ethiopia, no distinct Sambucus-stage could be observed. Apparently Sambucus

africana does not occur naturally in Ethiopia (Friis, pers. comm.); however, after

R. W. Bussmann)
Fig. 10 Sinarundinaria alpina (Poaceae), bamboo dye-back, Mt. Kenya National Park, Naro Moru
route. (Photo R. W. Bussmann)
Fig. 11 Sambucus africana (Caprifoliaceae) – Sinarundinaria alpina (Poaceae), stage after bam-
boo dye-back, with young re-sprouting bamboo in the background, Mt. Kenya National Park, Naro
Moru route. (Photo R. W. Bussmann)
dieback of the flowered bamboo, other light demanding species occur, until the
pioneer stage takes over again.
During the presented study, on Mount Kenya and the neighboring Aberdare Range,
bamboo stands were mainly found on the wetter western, southern, and southeastern
slopes and were absent in the Northern areas. From this distribution, it was concluded
that Sinarundinaria alpina requires precipitation of about 800–1000 mm per year
although arid periods are tolerated. In clear correspondence to the observations of
Wimbush (1945), new culms were mainly produced after prolonged rainy areas. During
the research period of 1992–1996, the main culm production occurred in large areas in
May/June 1995, after 9 months of unusually heavy rainfall. This, and the fact, that the
newly produced culms were mostly much larger and taller than the old ones clearly
depicts the dependency of bamboo growth on rainfall duration and intensity.
Mt. Nyiro, Harenna Escarpment (Bale Mts., Ethiopia) could be observed. In clear
climatic conditions. Therefore, it has to be concluded, that in contrast to many Asian
Although covering wide areas (Harenna, Bussmann 1997), Illubabor (Friis, pers.
Fig. 12 Sambucus africana

(Caprifoliaceae) –
Sinarundinaria alpina
(Poaceae), stage after bamboo
dye-back, with young
re-sprouting bamboo in the
background, Mt. Kenya
route. (Photo R. W.
Bussmann)
comm.) these forests were not recognized as specific forest types or not mentioned at
all by earlier authors. The large bamboo populations of South and Central Kenya
(Mt. Kenya, Aberdares, Mau, about 1000 km South of Harenna) are linked to
Harenna only by a small population occurring on top of Mt. Nyiro at the southern
end of Lake Turkana, which is also at least 500 km away from the Bale mountains.
As mentioned above, viable seeds of Sinarundinaria alpina have never been found,
and the species is known to regenerate only vegetatively, thus a propagation by
animals seems very unlikely. Therefore the large gaps between the main population
and even these outposts pose some interesting questions regarding the ecology and
former distribution of Sinarundinaria, as influenced by the palaeoclimate of the
region. This requires further fieldwork.
Fig. 13 Sinarundinaria
alpina (Poaceae), bamboo
dye-back, infrared image,
R. W. Bussmann)
route. (Photo R. W. Bussmann)
Fig. 15 Sinarundinaria
alpina (Poaceae), mature
bamboo with Podocarpus
latifolius, Mt. Kenya National
R. W. Bussmann)
mentioned by different authors (Agnew 1985; Schmitt and Beck 1992) are regarded
as clearly differentiated subassociations of the respective higher taxonomic units.
Local Food Uses
Young shoots are sometimes marketed as a vegetable (Mount Elgon, Kenya/Uganda

border).
The whole stems of African alpine bamboo are used for hut construction, particularly
as rafters, and for fencing (Beentje 1994). In the Poroto mountains in Tanzania entire
villages are constructed using this bamboo, and water pipes made from African
alpine bamboo supplied water to around 100,000 people in the 1980s. A few
specialized enterprises produce bamboo furniture. Split canes are woven into storage
pots and baskets in Tanzania and Kenya, in Uganda they are made into beehives. Dry
stems are used as fuel. The leaves and fine branchlets serve as cattle fodder.
Sinarundinaria alpina is also valued in soil conservation as a more effective
watershed cover than trees. The leaves are eaten by livestock, Bamboo makes strong
containers, and is used in circumcision ceremonies (Bussmann 2006a, b).
References
Agnew ADQ. Cyclic sequences in plant communities of the Aberdare mountains, Kenya. J East
Africa Nat Hist Soc Natl Mus. 1985;75:1–12.
management of a tropical mountain forest ecosystem. Dissertation Universität Bayreuth, Bay-
reuth; 1994.
Bussmann RW. Notes on the vegetation of Menagesha forest (Shoa Province, Ethiopia). In: BITÖK,
editor. Vegetationsökologische, ethnobotanische und -faunistische Beiträge über Äquatorial-
Afrika. Bayreuther Forum Ökologie, vol. 64. Bayreuth: BITÖK; 1999a. p. 105–18.
In: BITÖK, editor. Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie, vol. 64. Bayreuth: BITÖK; 1999b. p. 29–44.
p. 195–223.
2006b;11(1):41–66.
Phytocoenologia. 1995;25(5):467–560.
Bussmann RW, Beck E. Vegetation units of Mount Kenya Forest Reserve – comment. In: BITÖK,
Afrika. Bayreuther Forum Ökologie, vol. 64. Bayreuth: BITÖK; 1999. p. 17–28.
BITÖK, editor. Vegetationsökologische, ethnobotanische und –faunistische Beiträge über
Äquatorial-Afrika. Bayreuther Forum Ökologie, vol. 64. Bayreuth: BITÖK; 1999. p. 1–16.
Flora of tropical East Africa. Rotterdam: Balkema; 1970.
Dale IR, Greenway PJ. Kenya trees and shrubs. Buchanan: Nairobi; 1961.

Hedberg O. Vegetation belts of the East African mountains. Sven Bot Tidskr. 1951;45:140–99.
Mensch und Großwild. Jahrb Verein z Schutze der Bergwelt. 1992;57:207–26.
Speck H. Soils of the Mount Kenya Area, their formation, ecological, andagricultural significance.
In Winiger, M. ed. Mt. Kenya Area. Contributions to ecology and socio-economy. African
Studies Series. 1986;A1:47–72.
Wimbush SH. The african alpine bamboo. Empire Forestry J. 1945;24:33–9.
Olea capensis L.
Olea europaea subsp.
africana (Mill.) P.S. Greene
OLEACEAE

Grace N. Njoroge
Synonyms
Olea capensis L.: Maypea welwitschii Knobl.; Olea capensis subsp. enervis (Harv.)
I. Verd.; Olea capensis supsp. macrocarpa (C.H. Wright) I. Verd.; Olea enervis
Harv.; Olea guineensis Hutch. & C.A. Sm.; Olea hochstetteri Baker; Olea lancea
Lam.; Olea macrocarpa C.H. Wright; Olea perrieri A. Chev. ex H. Perrier; Olea
schliebenii Knobl.; Olea welwitschii (Knobl.) Gilg & Schellenb.; Ollea woodiana
Knobl.; Steganthus welwitschii (Knobl.) Knobl.
Olea europaea subsp. africana (Mill.) P.S. Greene: Olea africana Mill; Olea
chrysophylla Lam.
Local Names
Olea capensis: Kikuyu/Meru: Mucharagi, Mutharage, Mutharagi, Mutukuyu;

Elgon: Bumondet, Loliondet, Kiptakeriondu; Luhya: Muthukuyu, Omutukuyu;
Meru: Muukuru, Muriuntui; Nandi: Murguiywet; Ndorobo: Masaieta; Samburu:
R. W. Bussmann (*)
G. N. Njoroge

Lolio-ontoi; Kipsigis: Masaita; Okiek: Mosaita, Mosotik; Turkana: Eikwangat;

Iraqwi: Tlamno (Beentje 1994; Gachati 1989; Kokwaro 2009).
Olea europaea ssp. africana: Kikuyu: Mutamaiyu, Mutero; Swahili: Mzeituni,
Mzaituni, Ndorobo: Morien; Maa: Olorien; Chagga: Mlamuru (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: African olive
Botany and Ecology
Olea capensis L.: Tree 10–40 m high; bark pale gray, longitudinally fissured; slash
reportedly of various colors: orange-brown, cream or rich green; wood exceedingly
hard and durable. Young sterns glabrous, terete, bark gray, fairly smooth. Leaves
coriaceous; petiole 8–15 mm long, pale to conspicuously dark brown; lamina 4.5–9
(11) 1.2–4 cm, elliptic, ovate-elliptic or oblong-elliptic, with apex acute to
acuminate and tipped with a small hard point, and base cuspidate to acuminate;
upper surface glossy, with midrib impressed or level and the 5–6 pairs of lateral
nerves faintly raised, joining inconspicuously in loops; lower surface mat, and
paler, with prominent midrib and faintly raised lateral nerves; both surfaces punc-
tate with scattered circular scales just visible with hand lens, these sometimes
difficult to see on upper surface, lower surface never with metallic sheen; margin
slightly revolute, often somewhat undulate. Inflorescence a many-flowered panic-
ulate cyme, terminal and in axils of upper two leaf-pairs; bracts 0.3–1 mm long,
subulate. Calyx 1 mm long, cup-shaped, subentire with 4-toothed rim. Corolla
3 mm long, white, globose in bud and remaining thus for a prolonged period,
ultimately opening, the elliptic-acute lobes then reflexed; tube c. 1/3 as long as
lobes. Anthers 2 mm long. Gynoecium 1.5 mm long. Fruit up to 1.7 cm long (dry),
ellipsoid, with very thick woody endocarp. The species is widespread in Africa
from Guinea and Sierra Leone east to Ethiopia and Kenya, and south to
South Africa and Swaziland. It also occurs in Comoros and Madagascar, and on
Socotra Island (Yemen) (Turrill 1952). The species is relatively fast growing
(Bussmann 1999). A very characteristic component of semi-deciduous broad-
leaved Afromontane forests dominated by Cassipourea malosana (Bussmann
2002a, b, c, 2004, 2006a, b) (Figs. 1, 2, 3, 4, and 5).
Olea europaea subsp. africana (Mill.) P.S. Greene: An evergreen shrub or tree,
which grows up to 15 m tall. The leaves are borne in opposite pairs, are evergreen, 3–
9 cm long, elliptic, and silvery in appearance. The flowers are borne in axillary
clusters, with a four-lobed calyx, and a four-lobed corolla. The two stamens (male
parts) project beyond the mouth of the flower. The fruit has a hard endocarp (the
olive stone), which is surrounded by a fleshy, edible mesocarp. The wild African
olive occurs in Central, East, and southern Africa and in the Indian Ocean Islands. It
is also found in Arabia and from south-western Asia to China (Turrill 1952). The
species is relatively fast growing (Bussmann 1999). A very characteristic component
of dry Afromontane forests dominated by Juniperus procera (Bussmann 2002a, b, c,
2004, 2006a, b) (Figs. 6, 7, 8, and 9)
Olea capensis L. . . . 779
Fig. 1 Olea cf. capensis

(Oleaceae), inflorescence,

(Oleaceae), flowers, Bale
Olea capensis: In East Africa, bark decoctions are used as an emetic and anthel-
mintic, and to treat malaria, venereal diseases, and female sterility; bark ash is
applied as a dressing to wounds. In Swaziland, bark decoctions are taken to treat
peptic ulcers, and in South Africa, the bark is used for skin lightening. In southern
Africa, root powder is applied to fractures and joint swellings, and leaf infusions are
used to treat infections of the respiratory tract and pains. The boiled bark serves for
stomach problems and as anthelmintic (Bussmann 2006a, b).
Olea europaea ssp. africana: The leaves have been used for a long time to clean
wounds. Olive leaves are applied to lower blood pressure and to help improve the

(Oleaceae), Bale Mountains
Ethiopia. (Photo
R.W. Bussmann)

(Oleaceae), fruit, Bale
function of the circulatory system. Root decoction is used to treat malaria (Beentje
1994). The bark is soaked in water overnight and the infusion used to treat tape-
worms, and as general anthelmintic. The decoction is also drunk and used as steam
bath for rashes. The leaf decoction remedies liver problems (Delbanco et al. 2017;

(Oleaceae), bark cut, Bale
Fig. 6 Olea europaea subsp.

europaea (Oleaceae), leaves,
garden, Chicani, Bolivia.
Kokwaro 2009; Yineger et al. 2008). They are also taken as a mild diuretic and may
be used to treat conditions such as cystitis. Having some ability to lower blood sugar
levels, the leaves have been taken to treat diabetes. The oil is traditionally taken with
lemon juice in teaspoonful doses to treat gallstones. The leaf powder is applied for

europaea (Oleaceae), flowers,
garden, Chicani, Bolivia

europaea (Oleaceae), flowers,
ear problems (Lulekal et al. 2008; Teklehaymanot et al. 2007; Teklehaymanot 2009).
The boiled bark serves for stomach problems and as anthelmintic (Bussmann 2006a,
b; Nankaya et al. 2019, 2020), and also for malaria, colds, respiratory problems, and
flu (Muthee et al. 2011; Nankaya et al. 2020; Njoroge and Bussmann 2006a). It is
also used for anaplasmosis and to expel a retained placenta after birth (Muthee et al.
2011). Sometimes used for the treatment of diarrhea and as toothbrush (Gafna et al.
2017).

europaea (Oleaceae), habit,
Local Food Uses
Olea capensis: The oily fruits are edible and are used in southern Africa in the
preparation of beer and lemonade (Beentje 1994; Bussmann et al. 2011).
Olea europaea ssp. africana: The edible oil extracted from the mesocarp (pulp) of
the fruit and commonly used as a cooking and salad oil and in the preservation of
various foods.
Olea capensis: The heartwood, often traded as “ironwood,” is in high demand for
flooring, carpentry, and paneling, and is widely used for house and bridge construc-
tion, counter and tabletops, railway sleepers, tool handles, and wagon parts. It
produces beautiful furniture, turnery, and sliced veneer, and is often used by African
artists. It is suitable for interior trim, sporting goods, toys, novelties, and agricultural
implements (Beentje 1994). In South Africa, it has been used traditionally to make
assegais. It is also used as firewood and for charcoal production. The foliage serves
as fodder, especially during the dry season. The flowers produce nectar for honey-
bees. In South Africa, Olea capensis has been used as stock for grafting olive
cultivars from the Mediterranean region. The tree is considered sacred by the Maasai
people and is commonly used in ceremonies. The smoke is used to sterilize and give
fragrance to milk containers (Tian et al. 2017). Excellent timber for construction
(Bussmann et al. 2011) and as firewood (Kiefer and Bussmann 2004, 2008). The
leaves are eaten by livestock. Used for construction and fences. The wood makes
excellent rungus (war clubs), spear handles and sticks, and also excellent firewood
Olea europaea ssp. africana: The wood is valuable, hard, and fairly durable, but it is
rarely available in large sizes. It is used for turnery and furniture, and is much
appreciated for handicrafts; in larger sizes, it is also used for flooring and railway
Fig. 10 Olea europaea

subsp. africana (Oleaceae),
war club (rungus), made from
Olea wood, Sekenani camp,
R.W. Bussmann &
sleepers (Beentje 1994). The Maasai people of East Africa use it to make clubs and
for poles for houses. It makes excellent fuelwood and charcoal (Beentje 1994). The
bark decoction is used as enema to expel the placenta in cows (Kokwaro 2009).
Olive trees are planted for ornamental purposes, as firebreaks, and to control soil
erosion. The smoke is used to sterilize and give fragrance to milk containers (Tian
et al. 2017), and for fragrance in general (Mekonnen et al. 2015). The leaves are
eaten by livestock and used to treat endoparasites in cattle (Njoroge and Bussmann
2006b). Used for construction and fences. The wood makes excellent rungus (war
clubs), spear handles and sticks, and also excellent firewood (Bussmann 2006a, b;
Kiefer and Bussmann 2004, 2008) (Fig. 10).
References
p. 195–223.
2006b;111:41–66.
the forest expansion in east African montane forest ecosystems – a modeling approach. Lyonia.
2004;62:161–9.
10.1186/1746-4269-4-10.
pastoral Maasai girls in southern Kenya. J Ethnobiol Ethnomed. 2017;13:2. https://doi.org/
10.1186/s13002-016-0131-x.
Turrill WB. Flora of tropical East Africa: Oleaceae. London: Crown Agents for the Colonies; 1952.
31 pp. ISBN 978-1-84246-306-2.
10.1186/1746-4269-4-11.
Osyris lanceolata Hochst. & Steud.
SANTALACEAE

Grace N. Njoroge
Synonyms
Osyris lanceolata Hochst. & Steud.: Osyris abyssinica Hochst. ex A. Rich.; Osyris
densifolia Peter; Osyris divaricata Pilg.; Osyris laeta Peter; Osyris nepalensis Griff.;
Osyris oblanceolata Peter; Osyris parvifolia Baker; Osyris pendula Balf. f.; Osyris
quadripartita Salzm. ex Decne.; Osyris quadripartita var. canariensis Kämmer;
Osyris rigidissima Engl.; Osyris tenuifolia Engl.; Osyris urundiensis De Wild.;
Osyris wightiana J. Graham
Local Names
Osyris lanceolata: Kikuyu: Muthithi; Bena: Dunula; Irangi: Idululantumbi;

Kamba: Kithawa; Maa: Lassessi, Olosiyet; Marakwet: Jemokabyl; Kipsigis:
Kepurwet; Taita: Kijulu; Hehe: Muvamba-la-Fiduna; Ndorobo: Losesiai (Beentje
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Osyris lanceolata Hochst. & Steud.: Shrub or small tree, 1.5–9( 14) m tall, all
parts glabrous, bark coarsely furrowed, slash bright crimson, branches somewhat
flattened and sometimes rather pendent. Leaves usually alternate, elliptic or elliptic-
oblong, rarely obovate, 1.5–6.5 0.7–4 cm, abruptly apiculate, base cuneate, veins
immersed, only midvein raised beneath and running back down stem in a narrow
ridge, petiole 1–3 mm long, articulated to a small cushion. Flowers either hermaph-
rodite or male, plants androdioecious, hermaphrodite flowers in axils of upper
leaves, peduncles solitary, usually 1-flowered, occasionally flowers in 2–3 flowered
dichasia, male flowers both axillary and terminal, often panicled, each peduncle
usually terminating in an umbellate cluster of flowers, rarely flowers solitary or in a
2–3 flowered dichasium, peduncles 4–28 mm long, bracts and bracteoles linear-
lanceolate, 1–3 mm long. Perianth yellowish green, leathery, hermaphrodite flowers:
tube 0.5–0.6 mm long, obscured inside by disk, lobes 3( 4), spreading, ovate-
deltate, 1.5–2 1.8–2.2 mm, tips slightly hooded, male flowers similar. Stamens
3( 4), filaments 0.5–0.7 mm long, anthers 0.5 mm long. Ovary 2–2.8 mm long,
ovules 3( 4) in hermaphrodite flowers, ovules and placenta aborted in male
flowers, style 0.8–1 mm long, thick, cylindric, stigma in hermaphrodite flowers
normally 4- rarely 3-lobed, lobes ellipsoid, prominent, in male flowers both style
and stigma aborted or rudimentary. Fruit ellipsoid, 5–6.5 mm in diameter when dry,
epicarp thin, fleshy, red when ripe. Upland dry evergreen forest and mist forest,
with associated bushland and grassland, extending down rivers and from there
marginally into deciduous woodland, (50–)900–2700 m, widespread in Africa
from Algeria to Ethiopia and south to South Africa, in Europe (Iberian peninsula
and Balearic Islands), Asia (India to China), and Socotra (Polhill 2005). The
species is a characteristic component of the undergrowth of fry Afromontane
forests (Bussmann 2002) (Fig. 1).
Osyris lanceolata: Pounded leaves in cold water infusion are used for diarrhea.
Boiled roots (soup) used for gonorrhea, fever, and diarrhea. The bark decoction is
used to treat stomach problems in children (Maasai) (Kokwaro 2009). Bark is used to
remedy problems in pregnancy and as strengthening soup (Bussmann 2006). Used as
strengthening tea (Bussmann et al. 2006). The smoke of the burning bark is also used
for gynecological problems (Mohagehghzadeh and Faridi 2006). Used as anti-
malarial (Njoroge and Bussmann 2006a) and for neck problems (Njoroge and
Bussmann 2006b). Sometimes applied for ringworm, impotence, and fatigue
(Muthee et al. 2011). Osyris quadripartita is regarded as anti-bacterial (Geyid
et al. 2005), and also used to treat leishmaniasis (Giday et al. 2007; Mekonnen
et al. 2015), and for toothache (Yineger et al. 2008).
Osyris lanceolata Hochst. & Steud. 789
Fig. 1 Osyris lanceolata

(Santalaceae). Bale
Odo-Bulu forests, Ethiopia.
Local Food Uses
Osyris lanceolata: The fruits are edible (Beentje 1994).
Osyris lanceolata: Cold water leaf infusion for goats with stomach problems
(Kokwaro 2009). The roots yield a red dye, and root fibers are used to make baskets.
The wood can substitute sandalwood (Beentje 1994). Eaten by livestock, smoking
bark put close to swollen udder in cows (Bussmann 2006).
References
Geyid A, Abebe D, Debella A, Makonnen Z, Aberra F, Teka F, Kebede T, Urga K, Yersaw K,
Biza T, Haile Mariam B, Guta M. Screening of some medicinal plants of Ethiopia for their anti-
Polhill RM. Flora of tropical East Africa: Santalaceae. Kew: Royal Botanic Gardens; 2005.
2008;26:132–53.
Pavonia patens (Andrews) Chiov.
Pavonia urens Cav.
MALVACEAE

Grace N. Njoroge
Synonyms
Pavonia patens (Andrews) Chiov.: Pavonia burchellii (DC.) R.A. Dyer; Sida
patens Andrews
Pavonia urens Cav.: Pavonia bojeri Baker; Pavonia schimperiana Hochst. ex
A. Rich.; Pavonia tomentosa Hochst.
Local Names
Pavonia patens: Maa: Osupukai (Bussmann et al. 2006).

Pavonia urens: Kikuyu: Murera-Njau, Murwamba; Pare: Ishare; Kipsigis:
Modosiet; Maa: Olmeswa, Osubukioi-Orok; Marakwet: Matus; Samburu: Sulube
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Pavonia patens (Andrews) Chiov.: Erect to spreading, occasionally decumbent to

subscandent suffrutex or shrub up to about 1 m tall, biennial or perennial, varying
from sparsely and sometimes minutely stellate- or glandular-pubescent to stellate-
tomentose, sometimes also with longer patent hairs or fairly densely pilose; stems
herbaceous or wiry, terete, slender, reddish or yellowish or greenish, ultimately
glabrous and covered with a thin dark-gray to brown dark purple or black thin
smooth or finely fissured bark. Leaf-lamina 2–8 1–7 cm, usually ovate-pentagonal
in outline, often shallowly 3–5-lobed, with a usually deep and narrow basal sinus
(the basal lobes sometimes overlapping) and subacute to acute lobes, margin as a
rule coarsely and somewhat irregularly crenate to crenate-serrate, both surfaces
usually stellate-pubescent but the paler lower surface more densely so or occasion-
ally tomentose; petiole up to about 8 cm long, usually slender and hairy like the
stems; stipules up to about 5 mm long, setaceous, pubescent, usually caducous.
Flowers white, pale yellow, cream or orange, with or without a reddish center,
solitary in the upper leaf-axils; pedicels up to about 5 cm long, rarely longer, slender,
terete, pubescent, articulated in the upper 10–15 mm Epicalyx rotate; bracts S (6),
7–11 mm long, oblong to ovate-lanceolate or sub-rhomboid, acuminate to
acuminate-caudate, usually obtuse, 5-nerved, with or without long simple hairs in
addition to a stellate pubescence, usually ciliate, green and herbaceous but
accrescent in fruit and ultimately light brown. Calyx campanulate-cupulate, paler
and thinner in texture than the epicalyx, variously pubescent, usually divided a little
beyond the middle; lobes about 5 mm long, broadly ovate to somewhat rhombic,
acute or shortly acuminate, 5-nerved. Petals 10–15 mm long, often sparsely and
finely stellate-pubescent outside towards the apex. Staminal tube usually glabrous.
Mericarps about 5 4 3 mm, yellow or light brown when ripe, obliquely
triquetrous-obovoid, muticous, dorsally keeled and with warty protuberances or
muricate, laterally with 4–5 ridges, the latter and the protuberances often crowned
with sparse stellate hairs (at least when immature). Seeds about 3–5 2 mm,
obliquely obovoid-pyriform, somewhat angular, glabrous (Verdcourt and Mwachala
2009).
Pavonia urens Cav.: Erect suffrutex or shrub, 1–3 m tall, usually densely pubescent
on stems and leaves, sometimes also with longer soft hairs and/or with coarse harsh
and slightly pungent hairs (the latter causing a slight irritation of the skin) but
indumentum varying from pubescent to densely to sparsely tomentose-velutinous.
Leaf-lamina 3–20 cm in diameter, suborbicular-cordate in outline, the lower surface
usually more densely pubescent than the usually more or less scabrid upper surface,
usually 3–5-lobed, lobes triangular, acute and coarsely serrate-dentate, petiole
0.4–7 cm long, usually fairly uniform in length in one specimen, stipules about
5 mm long (occasionally longer), filiform, usually caducous. Flowers pale pink to
mauve or rather deep mauve-red, in subsessile clusters or in pedunculate
few-flowered inflorescences, occasionally solitary, in the upper leaf-axils of the
main branches and also sometimes on axillary short shoots, peduncle up to 5 cm
long, pedicels usually very short but up to 7 cm long in some flowers. Epicalyx of
Pavonia patens (Andrews) Chiov. . . . 793
Fig. 1 Pavonia patens

(Malvaceae), Sekenani,
R.W. Bussmann)
6–8 bracts, nearly as long as the calyx, bracts free, linear, pubescent and ciliate.
Calyx 8–10 mm long, cupuliform, densely pubescent and ciliate, glabrescent in fruit,
lobed to about the middle, lobes usually narrowly triangular, acute to acuminate.
Petals up to 2 cm long. Staminal tube glabrous. Mericarps about 5 mm long
(excluding the 3 apical protuberances or awns which are retrorsely spinose-hispid
and (1)5(7) mm long), dorsally reticulate with slightly raised ridges, somewhat
pubescent. Seeds 4 2 mm, reniform, faintly longitudinally striate (Verdcourt and
Mwachala 2009). Often found in the forest-bushland transition zone of Afromontane
forests, but also in the undergrowth of Phoenix reclinata dominated associations
(Bussmann 2002; Bytebier and Bussmann 2000) (Fig. 1).
Pavonia urens: A root decoction is used for intestinal problems (Kokwaro 2009).
The leaf paste is applied to boils (Rabearivony et al. 2015). The root powder is taken
to cure impotence (Teklehaymanot et al. 2007). Pavonia arabica leaf juice is applied
to snakebites (Taklehaymanot and Giday 2010).
Pavonia urens: Stems are used to make ropes (Beentje 1994; Bussmann et al. 2011).
Browsed by livestock (Bussmann et al. 2011).
Pavonia patens is used for the same purpose (Bussmann 2006; Bussmann et al.
2006, 2011). Browsed by livestock (Bussmann et al. 2011).
References
nationalities and peoples regional state, Ethiopia. J Ethnopharmacol. 2010;130:76–84.
Verdcourt B, Mwachala G. Flora of tropical East Africa: Malvaceae. Kew: Royal Botanic Gardens;
2009. 170 pp. ISBN 978-1-84246-189-1 [Fl. Trop. E. Africa, Malv.].
Phoenix reclinata Jacq.
ARECACEAE

Grace N. Njoroge
Synonyms
Phoenix reclinata Jacq.: Fulchironia senegalensis Lesch.; Phoenix abyssinica

Drude; Phoenix baoulensis A. Chev.; Phoenix comorensis Becc.; Phoenix
djalonensis A. Chev.; Phoenix dybowskii A. Chev.; Phoenix equinoxialis Bojer;
Phoenix leonensis Lodd. ex Kunth; Phoenix reclinata var. comorensis (Becc.) Jum.
& H. Perrier; Phoenix reclinata var. madagascariensis Becc.; Phoenix spinosa
Schumach. & Thonn.
Local Names
Phoenix reclinata: Kikuyu: Mukindu; Swahili: Mkindu; Boni: Gonyoorriya;

Borana: Meti; Digo: Mchindu; Ilwana: Gedo; Kamba: Makindu; Kipsigis:
Sosiyot; Luo: Othith; Maa: Ol-tukai; Oromo: Konchor; Rutoro: Mukindu;
Sanya: Itikindu; Somali: Ajol; Taveta: Mhongana; Taita: Kigangathei; Turkana:
R. W. Bussmann (*)
G. N. Njoroge

nakadoki (Beentje 1994; Gachati 1989; Kokwaro 2009); English: African date
palm, Wild date palm, Senegal date palm, Swamp date palm, False date palm
Botany and Ecology
Phoenix reclinata Jacq.: Clustering, very rarely solitary palm, often forming dense
thickets with trunks ultimately 10–15 m tall, more or less 15 cm diameter, occasion-
ally flowering while still acaulescent. Trunk dull brown, free of leaf-sheaths below,
with persistent leaf-sheaths above, in uppermost 1–2 m irregularly marked with
oblique leaf-scars and bearing persistent leaf-bases, injured stem exuding a clear
yellowish gum. Crown of about 25 green leaves or more, dead leaves frequently
rather long-persistent. Leaf to 2.5 m long, bright shiny green, not glaucous, arcuate,
true petiole about 15 cm, with coarse red-brown sacking-like sheath, apparent petiole
about 50 cm long by 2.5 cm wide, armed with fanned irregularly arranged
acanthophylls to 6 cm long by 4 mm wide, leaflets about 120 on each side of the
rachis, arranged very stiffly and regularly above, grouped below in 2’s–4’s, fanned,
to 25 cm long by 2 cm wide, pointed, occasionally sharply so, when old splitting
along the main vein, when young bearing caducous white indumentum on the lower
surface. Inflorescences with prophyll 20–50 cm long by 7–10 cm wide, often long
persisting, frequently splitting longitudinally into 2 halves, bright orangey brown at
anthesis, fading to dull gray-brown, covered with flocculent caducous gray-brown
indumentum when young. Peduncle of male inflorescence not greatly elongating,
sometimes scarcely emerging from the bract, 10–30 cm long by about 1.5 cm wide,
greatly compressed, rachillae 40–70 in number, arranged in groups and partial
spirals, to 15 cm long, usually much less, 2.5 mm in diameter near the base, tapering
to 1.5 mm distally. Female inflorescence emerging from the bract and often greatly
elongating after anthesis, with the fruiting rachillae pendulous, rachillae about 40–
60, 15–40 cm long, bearing up to 40 greenish flowers, singly or in small groups.
Male flowers creamy white, rapidly turning brown, musty scented, calyx about 1 mm
high, petals about 6–7 mm long, acute, fleshy, somewhat dentate towards the tip, the
tip slightly reflexed at anthesis, stamens 6, slightly shorter than the petals, pale
brown. Female flowers greenish, rounded, about 2 mm. in diameter, calyx about
1.5 mm high, petals rounded about 2 mm by 2 mm, carpels 3, about 2 mm tall, the
stigmas reflexed, just emerging from the tightly imbricating petals. Male flowers
creamy white, rapidly turning brown, musty scented, calyx about 1 mm high, petals
about 6–7 mm long, acute, fleshy, somewhat dentate towards the tip, the tip slightly
reflexed at anthesis, stamens 6, slightly shorter than the petals, pale brown. Peduncle
of male inflorescence not greatly elongating, sometimes scarcely emerging from the
bract, 10–30 cm long by about 1.5 cm wide, greatly compressed, rachillae 40–70,
arranged in groups and partial spirals, to 15 cm long, usually much less, 2.5 mm in
diameter near the base, tapering to 1.5 mm. distally. Female flowers greenish,
rounded, about 2 mm in diameter, calyx about 1.5 mm high, petals rounded about
2 mm by 2 mm, carpels 3, about 2 mm tall, the stigmas reflexed, just emerging from
the tightly imbricating petals. Female inflorescence emerging from the bract and
Phoenix reclinata Jacq. 797
Fig. 1 Phoenix reclinata (Arecaceae), young specimen, Mt. Kenya National Park, Chogoria,
often greatly elongating after anthesis, with the fruiting rachillae pendulous, rachillae
about 40–60, 15–40 cm long, bearing up to 40 greenish flowers, singly or in small
groups. Fruit almost always developed from 1 carpel only, very rarely all 3 develop-
ing, calyx in fruit to 2 mm high, petals in fruit to 5 mm high by 8 mm wide, fruit 1.3–
1.7 cm long by 0.9–1.3 cm wide, varying from pale yellow to orange or dull red,
epicarp smooth, mesocarp 1–2 mm thick, dry or moist and sweet. Seed 1–1.2 cm
long by 0.6–0.8 cm wide, deeply grooved along one side, embryo lateral. In the
lowlands, tending to grow along watercourses, in high rainfall areas and in the
mountains, growing also on open rocky hillsides, cliffs and even in rain forest, but
here, apparently confined to areas where the canopy is sparse, sea level to 3000 m.
Distributed throughout sub-Saharan Africa, from Senegal and Gambia eastwards to
Somalia and southwards to South Africa and Madagascar. It also occurs in Egypt,
Saudi Arabia, and Yemen, and is widely planted as an ornamental in tropical and
subtropical regions, also outside tropical Africa. (Dransfield 1986). The species
occurs as companion in wet Afromontane forests along creeks and rivers, and also
along small ponds and watercourses in dry savannah like forests. Sometimes Phoe-
nix reclinata forms dense populations especially on steep slopes, in areas disturbed
by fire, e.g., in Mathews Range, Kenya (Bussmann 2002, 2006) (Figs. 1, 2, 3, and 4).
Phoenix reclinata: In traditional medicine in Senegal, root macerations are consid-

ered astringent and taken for the treatment of stomachache and diarrhea, a decoction
of the apical bud is used in baths and draughts against tiredness, and preparations of
the leaflets are externally applied for the treatment of eye diseases. In Côte d’Ivoire,
Fig. 2 Phoenix reclinata (Arecaceae), mature specimen, Mt. Kenya National Park, Chogoria,
the fruit is used for the treatment of female sterility. In Uganda, the pounded roots
mixed with banana juice or beer are taken against gonorrhea and impotence, and
fresh leaves are chewed and swallowed against abdominal pain. In Tanzania, a root
decoction is drunk against epilepsy. The leaves can be roasted, pounded, and mixed
with water to remedy stomachache and flu (Bekalo et al. 2009; Kokwaro 2009).
Local Food Uses
Phoenix reclinata: The fruits are edible (Beentje 1994).
Phoenix reclinata: The leaflets or strips of the leaflets of young, unexpanded leaves
are widely used for making mats, bags, baskets, hats, sieves, rope, string, fishing
nets, traps, parasols, and ornaments. Strips from mature leaves are used for weaving
hats. The split petiole and rachis of mature leaves are used in coarse weaving, e.g., of
baskets, mats and fish traps, and for tying. The whole rachis is used for making huts
and fish kraals. The Maasai use the rachis for cleaning gourds. The leaves are used
for thatching and as fans (Beentje 1994). Hand brooms are made from the stem by
pounding its end until the fibers separate. Dried inflorescences are also used as
brooms. The trunk, although often bent, is much used in the construction of huts,
houses, fences, bridges, landing-stages, traps, and hives. In Ghana, the wood is made
Phoenix reclinata Jacq. 799
Fig. 3 Phoenix reclinata

(Arecaceae), young specimen,
Laikipia Plateau, Kenya.
into drums. The wood is also used as fuelwood and is suitable for charcoal produc-
tion. The base of the stem is sometimes eaten. The palm is tapped for sap to be
fermented into a palm wine. The apical bud is eaten as a vegetable (“palm cabbage”),
which is said to be slightly bitter. The ripe fruits are eaten, sometimes after having
been immersed in boiling water for a moment. In Sierra Leone, the kernels are
parched and ground into flour for consumption. Roasted seeds are used as a coffee
substitute. The roots yield an edible gum and contain tannins. In Kenya and
Tanzania, a brown dye is obtained from the root. The species is widely planted as
an ornamental and locally for shade and amenity. The leaves and petioles are used for
dune fixation. In various countries, the leaves have ceremonial uses. In Senegal, they
are made into loincloths to be worn at circumcision ceremonies. In Ethiopia, the
leaves are used on Palm Sunday in the Orthodox Church and as decorations for
weddings. Between Palm Sunday and Easter, children use the young soft leaflets for
making finger rings, and woven into floormats and baskets, especially for weddings,
and to store milk and butter (Bussmann et al. 2011; Luizza et al. 2013).
Fig. 4 Phoenix reclinata (Arecaceae), mature specimen, Laikipia Plateau, Kenya. (Photo
R.W. Bussmann and N.Y. Paniagua-Zambrana)
References
doi.org/10.1186/1746-4269-5-26.
2006;111:41–66.
Dransfield J. Flora of Tropical East Africa: Palmae. Rotterdam: Balkema; 1986. 55 pp. ISBN
90-6191-329-2.
Res Appl. 2013;11:315–39.
Physalis peruviana L.
SOLANACEAE

Grace N. Njoroge
Synonyms
Physalis peruviana L.: Alkekengi pubescens Moench; Boberella peruviana (L.)

E.H.L. Krause; Boberella pubescens (L.) E.H.L. Krause in Sturm; Physalis
chenopodifolia Lam.; Physalis edulis Sims; Physalis escilenta Salisb.; Physalis
latifolia Lam.; Physalis peruviana var. latifolia (Lam.) Dunal; Physalis pubescens L.;
Physalis tomentosa Medik.
Local Names
Physalis peruviana: Kikuyu: Kinathi; Luhya: Shiyengo Shisatsa (Gachati 1989;

Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Physalis peruviana L.: Herbs perennial, 45–90 cm tall. Stems erect, sparingly
branched, densely pubescent. Petiole 2–5 cm, leaf blade broadly ovate to cordate,
6–15 4–10 cm, densely pubescent, base cordate, margin entire or with a few
indistinct teeth, apex short acuminate. Pedicel ca. 1.5 cm. Calyx broadly campanu-
late, 7–9 mm. Corolla yellow, spotted in throat, 1.2–1.5 1.2–2 cm. Filaments and
anthers blue-purple, anthers 3–4 mm long. Fruiting calyx green, ovoid, with 5–10
weak angles, 2.5–4 cm, pubescent. Berry yellow, 1–1.5 cm in diameter. Seeds
yellow, about 2 mm in diameter Flowering summer, fruiting autumn (Wu et al.
1994–2013). In Africa, mostly regarded as noxious weed (Bussmann et al. 2011)
(Figs. 1, 2, and 3).
Physalis peruviana: A leaf decoction is used against malaria (Kokwaro 2009). The
leaf decoction is used to treat asthma (Njoroge and Bussmann 2006). Leaves and
stem are employed to remedy postpartum pain (Njoroge and Bussmann 2009). The
species is also used against typhoid and as anthelmintic (Njoroge et al. 2004). In
Madagascar, it is often sold in local markets to treat rheumatism, urinary tract
problems, syphilis, stomach ulcer, and hepatitis (Randriamiharisoa et al. 2015).
Bladder cherry fruits have diuretic properties, and fruit oil accelerates healing of
wounds. Extracts from the fruit are used by homeopaths. Tea from dried leaves and
fruits is useful in case of hypertension. The plant contains a wide variety of
physalins, which have antibacterial and leishmanicidal activities. The fruits contain
vitamin C, the seeds fatty oils. An infusion and decoction of fresh and dried fruits is
applied as diuretic. The syrup from the fruits is used as anti-inflammatory and
emollient. An ointment from fruits is used externally to relieve inflammatory
Fig. 1 Physalis alkekengi

(Solanaceae). Bakuriani,
Georgia. (Photo:
R. Bussmann)
Physalis peruviana L. 803

(Solanaceae). Immature fruits.
Bakuriani, Georgia. (Photo:
R. Bussmann)
processes of the skin, and a decoction of fruits is ingested to treat syphilis. The fruits
are used as antimalarial. In other regions, the fruits serve as emetic (Fayvush et al.
2017). The smoke of burnt Physalis species is used to treat skin diseases
Local Food Uses
Physalis peruviana: Fruits can be used as a food (pickled and as a dried fruit, after
frost and fresh). The leaves are sometimes eaten in phkhali (a vegetable and walnut
spread) (Fayvush et al. 2017). Also eaten raw (Lulekal et al. 2008).
Physalis peruviana: The fruits are used for dyeing silk in yellow and orange tones as
well as for dyeing white oil of buffalo cow in yellow color (Fayvush et al. 2017).

(Solanaceae). Mature fruits.
References
Fayvush G, Aleksanyan A, Batsatsashvili K, Mehdiyeva N, Kikvidze Z, Khutsishvili M, Maisaia I,
Sikharulidze S, Tchelidze D, Alizade V, Paniagua Zambrana NY, Bussmann RW. Physalis
alkekengi L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer Interna-
tional Publishing; 2017. https://doi.org/10.1007/978-3-319-49412-8_81.
4269-4-10.
2009;82:262–9.
Physalis peruviana L. 805
org/10.1186/s13002-015-0046.
Phytolacca dodecandra L’Hér.
PHYTOLACCACEAE

Grace N. Njoroge
Synonyms
Phytolacca dodecandra L’Hér.: Phytolacca abyssinica Hoffm.; Phytolacca

dodecandra var. brevipedicellata H. Walter; Pircunia abyssinica Moq.
Local Names
Phytolacca dodecandra: Kikuyu: Muhoko; Kipsigis: Batkanet; Maa: Oldinagaras;

Marakwet: Kipsugotit; Nandi: Kobot Kanet; Samburu: Jebetit; Turkana:
Ejemait; Luhya: Libokho; Luo: Mahjoho, Oooho, Owoho; Tukuyu: Mbopola;
Sebei: Sikotit; Iraqw: Tloxi (Beentje 1994; Gachati 1989; Kokwaro 2009);
English: Endod, soap berry, African soap berry
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Phytolacca dodecandra L’Hér.: A semi-succulent straggling or scrambling shrub up

to 4 m, sometimes climbing up to 6–9( 18) m, glabrous or less commonly crisped
pubescent on the vegetative parts. Leaf-blades ovate to elliptic, 6–15 cm long, 3–8.5
( 10) cm wide, narrowed to a prominently and ultimately recurved mucronate tip,
cuneate, rounded or cordate to the shortly decurrent unequal-sided base, glossy, with
translucent margins (at least when dry), crystals visible as short raised lines and dots
on the younger leaves, primary lateral nerves 4–8 on either side, petioles mostly
1.5–4 cm long. Racemes shortly pedunculate, about 15–30( 40) cm long, sub-
densely very many-flowered, usually crisped pubescent except often towards the
base, bracts subulate to linear-lanceolate, 1–2 mm long, pedicels 1.5–5( 10) mm
long, longest in long-staminate flowers, with small bracteoles near the middle or
above. Flowers dimorphic, scented. Long-staminate flowers white or cream, stamens
(10–)12–15, filaments 3–4( 5) mm long, anthers oblong-elliptic, 0.8–1.2 mm long,
carpels half as long as the filaments and only very rarely developing. Short-
staminate flowers yellowish-green, stamens 8–15, with 1–1.5 mm long filaments and
rather smaller anthers, carpels (4–)5( 8), obliquely flattened-ovoid, united only at
the base. Sepals triangular-lanceolate to oblong or ovate-oblong, bluntly pointed,
2–3 mm long, slightly fleshy, pubescent at least along the margins, becoming
reflexed. Berry bluntly star-shaped in outline, the free part of the swollen carpels
spreading, with the style-bases at the points, becoming more rounded, juicy and
ripening orange or red, 5–8 mm across. Seeds lenticular, 2.5–3( 4) mm across,
papillate and striate towards the hilum, glossy black. Occurring in a wide range of
forest, woodland, bushland, thicket and grassland communities, commonly riparian,
generally perhaps on old forest land, limited only by extremes of altitude and
rainfall, 500–2400 m. Native to sub-Saharan Africa and Madagascar, introduced in
Asia and tropical America. As weed in drier Afromontane forests (Bussmann 2002)
(Figs. 1, 2, 3, 4, 5, 6, 7, and 8).
Fig. 1 Phytolacca icosandra

(Phytolaccaceae),
Leymebamba, Peru. (Photo
R.W. Bussmann and
Phytolacca dodecandra L’Hér. 809
Fig. 2 Phytolacca americana

(Phytolaccaceae), Zugdidi,
Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and
Phytolacca dodecandra: In Central and East Africa and Madagascar, various parts
of Phytolacca dodecandra are used as a medicine, despite the toxicity of the plant.
An extract of the roots, leaves, fruits, and seeds is taken in small quantities as a

Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and
purgative for humans and animals, as an anthelmintic, and to treat edema and
intestinal problems such as diarrhea and abdominal pain, as a laxative, emetic,
sudorific, and diuretic. Also used for venereal diseases, especially syphilis (Bekalo
et al. 2009; Kokwaro 2009). The leaf sap, and sometimes the crushed roots and
fruits, are applied to wounds and skin ailments such as ringworm, scabies, eczema,
psoriasis, leprosy, boils, and vitiligo; the sap is cicatrizing and hemostatic and causes
Fig. 6 Phytolacca
cf. dodecandra
(Phytolaccaceae), Bale
Fig. 7 Phytolacca
cf. dodecandra
a burning sensation on the skin. In DR Congo, the fresh, dried, or powdered leaves
are used for the same purposes. An infusion of the fruit or roots is taken orally, and
the young leaves and shoots are chewed to induce abortion. A large lump of butter is
taken to stop further contractions. An infusion of the fruit or the root decoction is
also widely taken to treat venereal diseases, bilharzia, rabies, malaria, sore throat and
other respiratory problems, rheumatic pain, jaundice, as well as anthrax and leeches
in animals. In southern Nigeria, the leaf decoction is given to newborn babies as a
gentle laxative. In Central Africa, the mashed leaves are eaten with banana as a
stimulant and tonic, especially after childbirth. In Congo, the leaf sap is used as eye
drops to cure conjunctivitis and river blindness. In Rwanda, leaf sap is used as ear
drops to treat otitis. In DR Congo, the boiled leaves with chicken are given to
children with asthma and tuberculosis. In East Africa, ground leaves are applied to
Fig. 8 Phytolacca
cf. dodecandra
tumors, the root decoction is also drunk to cause vomiting as treatment of enlarged
glands. In Tanzania, macerated leaves or root bark are used to treat epilepsy. In
Madagascar, a decoction of the aerial parts is applied to treat hemorrhoids. In
South Africa, the root infusion is taken to treat infertility. Used for “Gara Gelcha”
and “Chachabsa” in Ethiopia (Yineger et al. 2007, 2008). Also ingested to treat
hepatitis and jaundice (Teklehaymanot 2009), as abortive and against rabies
(Teklehaymanot et al. 2007). To treat ascariasis, gonorrhea, malaria, rabies, sore
throat, rheumatic pain, jaundice, syphilis, pruritus, eczema, and vitiligo (Giday et al.
2007, 2009, 2010; Luizza et al. 2013; Tadeg et al. 2005), itchy skin, and wound
treatment (Giday et al. 2007), as well as to treat scabies (Gedif and Hahn 2003). The
roots are chewed for stomach problems, and the twigs are used as toothbrush
Local Food Uses
Phytolacca dodecandra: There is considerable difference in opinion about the

edibility of the leaves. In Côte d’Ivoire and southern Nigeria, a sauce or soup is
prepared from the young shoots and young leaves, and in DR Congo, the leaves are
cooked as a vegetable. In eastern and southern Africa, the whole plant is considered
poisonous, and it is said to have caused accidental death of people eating the leaves
as a vegetable. In Gabon, the fruits are eaten. In East Africa, the plant is regarded as
highly toxic (Beentje 1994).
Phytolacca dodecandra: In Uganda and DR Congo, certain cattle tribes use the plant
to increase milk production of cows. The fruits, when dried, powdered, and mixed
with water, yield a foaming detergent, traditionally used in Ethiopia, Somalia, and
Uganda for washing clothes, and also to wash the body. Soap has also been made
from the ashes of burnt plants in southern Nigeria, or from the fresh leaves in
Angola. In Ethiopia and Zimbabwe, unripe fruits are widely applied to control
bilharzia-transmitting snails, as they contain molluscicidal saponins. It is a cheaper
and less toxic product than synthetic molluscicides. In East Africa, the pounded plant
is also used as a poison to catch fish. In West Africa and Ethiopia, the leaves are
considered edible for cattle and goats, whereas in most of eastern and southern
Africa, they are considered poisonous. In East Africa, the stems are used as ties in the
construction of huts and fences. The fruits yield a red dye and the leaves a yellow
dye. In Zimbabwe, the most common use of the fruits is to color the floor in houses.
In Ethiopia, Phytolacca dodecandra is planted as a hedge, and the leaf or fruit extract
is sometimes added to drinks and foods as a stimulant, and to curdle milk. The plant
is usually not used as firewood, as the smoke is believed to reduce the male sexual
ability. A decoction is used as purgative for livestock. Also as washing to remove
lice, and in cows as infusion to remove a retained placenta after birth (Kokwaro
2009). The leaf juice is used as acaricide (Wanzala 2017). Used as fishing poison
(Neuwinger 2004). Smoke as anti-aphrodisiac (men) (Mohageghzadeh and Faridi
2006), mosquito repellent (Karunamoorthi et al. 2009), and plant used to make
unspecified household tools (Mekonnen et al. 2015). The fruits are used to wash
clothes (Bussmann et al. 2011; Luizza et al. 2013). The extract is used as mollusci-
cide (against bilharzia) (Bussmann et al. 2011).
References
doi.org/10.1186/1746-4269-5-26.
Res Appl. 2013;11:315–39.
2005;100:168–75.
10.1186/1746-4269-4-11.
Pinus patula Schltdl. & Cham.
Pinus radiata D. Don.
PINACEAE

Grace N. Njoroge
Synonyms
Pinus patula Schltdl. & Cham.: Pinus patula subsp. tecunumanii (F. Schwerdtf. ex
Eguiluz & J.P. Perry) Styles
Pinus radiata D. Don.: Pinus insignis Douglas ex Loudon
Local Names
Pinus patula: Kikuyu: Mucinda-Nugu (Gachati 1989).

Pinus radiata: Kikuyu: Mucinda-Nugu (Gachati 1989).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Pinus patula Schltdl. & Cham.: Tree that reaches up to 40 m in height and a
diameter of 1.20 m. Conical trunk, straight and without sneakers. It has whorled
branches, which begin to form from the base. The bark on young trees and twigs is
thin and scaly, reddish brown. In mature trees, it is thick fissured and dark grayish
brown. Leaves in groups of 3 and sometimes 4, rarely 5 in some fascicles; about
15–30 cm long acicular, thin, sharp, vertically drooping, bright green, with finely
serrated edges. Flowers on catkins that form long conical cones, 7–9 cm to 12 cm,
sessile, somewhat curved, oblique and pointed and usually grouped from 3 to
6. Originally from the region between Tamaulipas and Oaxaca in Mexico. It grows
in plant formations: humid and very humid low montane forest. In its natural
distribution, it is part of the cloud forests and associated with the species: Pinus
ayacahuite, Pinus michoacana, Pinus lumholtzzi, and Pinus leiophylla. Common
plantation tree in Afromontane forests and contributing to their destruction
(Bussmann 1996, 2006) (Figs. 1, 2, 3, and 4).
Pinus radiata D. Don.: Trees to 30 m; trunk to 0.9 m diameter, contorted to straight;
crown broadly conic, becoming rounded to flattened. Bark gray, deeply V-furrowed,
furrow bases red, ridges irregularly elongate-rectangular, their flattened surfaces
scaly. Branches level to downcurved or ascending, poorly self-pruning; twigs slen-
der, red-brown, sometimes glaucous, aging gray, rough. Buds ovoid to ovoid-
cylindric, red-brown, about 1.5 cm, resinous. Leaves (2–3) in a fascicle,
spreading-ascending, persisting 3–4 years, (8–)9–15( 20) cm, 1.3–1.8( 2) mm,
Fig. 1 Pinus kochiana (Pinaceae). Bakuriani, Georgia. (Photo: R. Bussmann)

Pinus patula Schltdl. & Cham. . . . 817
Fig. 2 Pinus kochiana

(Pinaceae). Bakuriani,
Georgia. (Photo:
R. Bussmann)

(Pinaceae). Kazbegi, Georgia.
(Photo: N. Paniagua-
Zambrana)

(Pinaceae). Male cones
collected and dried.
N. Paniagua-Zambrana)
straight, slightly twisted, deep yellow-green, all surfaces with fine stomatal lines,
margins serrulate, apex conic-subulate; sheath (1–)1.5–2 cm, base persistent. Pollen
cones ellipsoid-cylindric, 10–15 mm, orange-brown. Seed cones maturing in 2 years,
shedding seeds soon thereafter, but often serotinous and persistent 6–20 years,
solitary to whorled, spreading to recurved, curved, very asymmetric, ovoid before
opening, broadly ovoid when open, 7–14 cm, pale red-brown and lustrous, scales
rigid, stalks to 1 cm; apophyses toward outer cone base increasingly mammillate,
those on inward cone side and middle and apex of cone more level; umbo central,
mostly depressed, with small central boss or occasionally with slender, deciduous
prickle. Seeds compressed-ellipsoid; body ca. 6 mm, dark brown; wing 20–30 mm.
Common plantation tree in Afromontane forests and contributing to their destruction
(Bussmann 1996, 2006).
Pinus patula needles are used to treat diarrhea (Njoroge and Bussmann 2006).
Pine roots are also used to treat tuberculosis. Pollen is eaten dried to help
bronchitis and inhaled for sinusitis (Batsatsashvili et al. 2017; Bussmann et al.
2014, 2016a, b, c, 2017a, b, 2018; Bussmann 2017). Pinus roxburghii is used
widely for asthma and bronchitis, and the resin also for sores (Bhat et al. 2013;
Joshi et al. 2010). The plan serves also as nerve tonic and antiseptic, for skin ulcers,
and as hemostatic (Kunwar et al. 2009, 2010a, b, 2012, 2013), and boils (Kunwar
et al. 2013). Sometimes the resin is used to treat snakebites (Ur-Rahman et al. 2018)
and also serves for toothache and diarrhea (Muhammad et al. 2019). The resin is
used to treat cracked toes (Joshi et al. 2010). Similar uses are reported for Pinus
tinctoria (Kunwar et al. 2010a, b). Pinus gerardiana fruits are used for headache
(Wali et al. 2019), joint health (Sher et al. 2016), and skin problems (Ali et al. 2019).
Pinus wallichiana is used to treat heart attacks, wounds, stomach and urinary
problems, asthma and bronchial congestion (Ullah et al. 2019; Wali et al. 2019), as
well as skin problems (Ali et al. 2019) (Figs. 5, 6, and 7).

(Pinaceae). Pollen for sale.

(Pinaceae). Syrup made from
flowers. Bakuriani, Georgia.
Zambrana)

(Pinaceae). Syrup. Tusheti,
Georgia. (Photo:
R. Bussmann)

(Pinaceae). Chewing-gum
made from resin and inner
bark. Bakuriani, Georgia.
Zambrana)
Fig. 9 Pinus radiata (Pinaceae). Resin extraction, Madagascar. (Photo: R. Bussmann)
Fig. 10 Pinus radiata

(Pinaceae). Resin extraction,
Madagascar. (Photo:
R. Bussmann)
Fig. 11 Pinus radiata

(Pinaceae). Resin extraction,
Madagascar. (Photo:
R. Bussmann)
Local Food Uses
Leaves and young cones are sometimes eaten (Batsatsashvili et al. 2017; Bussmann et al.
2014, 2016a, b, c, 2017a, b, 2018; Bussmann 2017; Liu and Bussmann 2020) (Fig. 8).
Pinus patula is often planted as live fence (Mekonnen et al. 2015).

Pinus gerardiana is used as firewood, the cone as detergent, for furniture, and the
gum for hair removal. The species has high market value (Singh et al. 2019; Wali et
al. 2019) (Figs. 9, 10, and 11).
References
org/10.32859/era.18.24.1-19.
Tchelidze D, Aleksanyan A, Alizade V, Paniagua Zambrana NY, Bussmann RW. Pinus kochiana
Klotzsch ex K. Koch. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer
International Publishing; 2017. https://doi.org/10.1007/978-3-319-49412-8_51.
2006;111:41–66.
Bussmann RW ed. Ethnobotany of the Caucasus. Springer International Publishing International
Publishing: Cham 2017;XXVII, 746p. ISBN 978-3-319-49411-1
2014;12:237–313.
Samtskhe-Javakheti, Tusheti, Svaneti, and Racha-Lechkhumi, republic of Georgia (Sakartvelo),
(Republic of Georgia), Caucasus. Med Aromat Plants. 2016b;5:5.
Khutsishvili M, Batsatsashvili K, Hart RE. Medicinal and food plants of Tusheti, Khevsureti and
Pshavi, Sakartvelo (Republic of Georgia), Caucasus. Acta Soc Bot Pol. 2016c;86(2):3517.
(Republic of Georgia), Caucasus. Indian J Tradit Knowl. 2017a;16(1):25–34.
gia), Caucasus. Indian J Tradit Knowl. 2017b;16(1):7–24.
Sakartvelo (republic of Georgia), Caucasus. Indian J Tradit Knowl. 2018;17(1):7–33.
Biodiversität, Naturausstattung im Himalaya, vol III. Erfurt; 2009. Universität Erfurt, p. 475–89.
Kunwar RM, Burlakoti C, Chowdhary CL, Bussmann RW. Medicinal plants in Far-west Nepal:
their indigenous uses and pharmacological validity. Med Aromat Plant Sci Biotechnol. 2010a;4
(special issue 1):28–42.
pharmacological appraisal. J Ethnobiol Ethnomed. 2010b;6:35.
species in Far-west Nepal – a valuable resource being wasted. J Mt Sci. 2012;9:589–600.
and management in Far-west Nepal. J Ethnobiol Ethnomed. 2013;9:24.
Liu B, Bussmann RW. Pinus sylvestris L. var. mongolica Litv. In: Batsatsashvili K Kikvidze Z,
Bussmann RW, editors. Ethnobotany of mountain regions Central Asia and Altai. Springer
International Publishing; 2020. https://doi.org/10.1007/978-3-319-77087-1_103-1.

https://doi.org/10.32859/era.18.14.1-11.
the Southern and Tribal regions of Khyber Pakhtunkhwa province, Pakistan. Ethnobot Res Appl
2019;188. https://doi.org/10.32859/era.18.8.1-20.
Wali S, Ahmad Jan H, Bussmann RW. Quantitative ethnomedicinal study of indigenous medicinal
plants used for digestive disorders of Laspur Valley, Chitral, Northern Pakistan. Ethnobot Res
Appl. 2019;18:32. https://doi.org/10.32859/era.18.32.1-18.
Piper capense L.
PIPERACEAE

Grace N. Njoroge
Synonyms
Piper capense L.: Piper bequaertii De Willd.; Piper emirnense Baker; Piper
humboldtii C. DC.
Local Names
Piper capense: Kikuyu: Muruya, Murungu, Suguya; Kipsigis: Laraba; Rutoro:

Mulembat (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Piper capense L.: A shrub, erect or more or less scandent. Leaves ovate to elliptic, or
rounded ovate, 5–18 cm long, 2.5–13 cm wide, distinctly acuminate at the apex,
cuneate to cordate at the base, glabrous above, glabrous to pubescent on nerves
R. W. Bussmann (*)
G. N. Njoroge

beneath or in variety more or less hairy all over, palmately 3–9( 11)-nerved, the
3 median nerves reaching the apex, petiole 0.5–5( 8.3) cm long, glabrous to hairy,
stipule-like outgrowths lanceolate, 1–1.6 cm long, thin, deciduous. Weakly erect
aromatic shrub or subshrub or more or less herbaceous or sometimes a straggling
liane 1–5 m. long or tall, sometimes several-stemmed from the base, stems glabrous
or hairy, swollen at the nodes, up to 3 cm wide (in life), rootstock tuberous. Stamens
2–3. Flowers hermaphrodite or female, either both on one spike or on separate
spikes, spikes creamy white, solitary, leaf-opposed, terminal and pseudolateral but
in reality terminal but overtopped by sympodial growth, 0.7–10 cm long, peduncle
1–5.2 cm long, glabrous or hairy, bracts more or less round, 1–1.5 mm wide,
pubescent below. Fruit white or green, rounded ovoid, 2–4 mm long, compressed,
obtuse, seeds brown, shiny (Verdcourt 1996). Common in the undergrowth of
evergreen Afromontane forests (Bussmann 2002) (Figs. 1, 2, 3, and 4).
Fig. 1 Piper aduncum

(Piperaceae), Leymebamba,
Piper capense L. 827


Fig. 4 Piper aequale

Piper capense: The seeds are used to treat cough. The root decoction serves as
anthelmintic, and the leaf infusion is used for diarrhea (Bekalo et al. 2009; Kokwaro
2009). Used to treat common cold (Njoroge and Bussmann 2006).
In Asia, Piper nigrum is used to treat goiter, Piper longum is a commonly used
plant in ayurvedic medicine (Reang et al. 2016). Piper nigrum also serves for
jaundice, typhoid, body pain, pneumonia, diabetes, cough, gastroenteritis, dysentery,
stomach disorder to improve appetite, and to treat dog bites (Raj et al. 2018). In
Madagascar, the species is used for digestive disorders and as poultice for joint pain
(Razafindraibe et al. 2013). Piper borbonense is used in Madagascar for digestive
system problems (Rakotoarivelo et al. 2015; Randrianarivony et al. 2016). Piper
sp. is used to treat gonorrhea (Rabearivony et al. 2015). Piper angustifolium is used
to increase vigor (Ali et al. 2019).
The most widespread use of Piper species occurs in South America, where the
genus has its major distribution.
Piper acutifolium is used for uterus, uterus prolapse, cold, flu, internal cold,
kidney infection, prostate, and bad air (Bussmann et al. 2016). Also used for colds,
cough, wounds, for bone pain, and rheumatism (Quiroga et al. 2012).
Piper aduncum stems and leaves are, e.g., used to treat hemorrhage, sores,
indigestion, dysentery, and as astringent (Bussmann et al. 2018). The whole fresh
plant is used to treat liver and wounds (Béjar et al. 2001; Bussmann and Sharon
2006a, 2007a). Leaves, fresh or dried, are used to treat cold, fungus, cough, wounds,
bronchitis, chills, tuberculosis, and for stopping a hemorrhage, and has antibacterial
properties (Bussmann and Sharon 2006b, 2007b, 2015a, b; Monigatti et al. 2013).
Piper aequale leaves and stems, fresh or dried, are used to treat liver, hepatitis,
and infection in the body (Bussmann and Sharon 2006b, 2007b, 2015a, b).
Piper capense L. 829
References
org/10.32859/era.18.24.1-19.
Medicinales del Sur Ecuatoriano, 340p. San Diego: Latino Herbal Press; 2001.
doi.org/10.1186/1746-4269-5-26.
Syst Geogr Plants. 2002;712:959–74.
9789962-2-2.
Bussmann RW, Sharon D. Plants of the four winds – The magic and medicinal flora of Peru. Plantas
978-0-9789962-3-9.
medicinal flora of northern Peru. St. Louis: William L. Brown Center, MBG; 2015a. ISBN
978-0-9960231-2-2.
9960231-3-9.
Bussmann RW, Paniagua-Zambrana NY, Romero C, Hart RE. Astonishing diversity – the medicinal
plant markets of Bogotá, Colombia. J Ethnobiol Ethnomed. 2018;14(1):43. https://doi.org/
10.1186/s13002-018-0241-8.
(2):450–64.
Quiroga R, Meneses L, Bussmann RW. Medicinal ethnobotany in Huacareta (Chiquisaca, Bolivia).
J Ethnobiol Ethnomed. 2012;8:29.
018-0208-9.

gascar Conserv Dev. 2016;11(2):44–51.
2013;9:73.
Aromac Plants. 2016;5:2.
Verdcourt B. Flora of tropical East Africa: Piperaceae. Rotterdam: Balkema; 1996.
Pistacia aethiopica Kokwaro
ANACARDIACEAE

Grace N. Njoroge
Local Names
Pistacia aethiopica: Kikuyu: Muheheti, Muhehete, Mucherere; Kamba: Musaa;

Kipsigis: Kibirirgorokiet, Chepkorokwet; Maa: Ol-Daangudwa; Samburu:
Olongonorok, Iltorel, Lasamarai; Tugen: Tulda (Gachati 1989; Beentje 1994).
Botany and Ecology
Pistacia aethiopica Kokwaro: Glabrous evergreen spreading tree 5–15 m high or

more, rarely a shrub, often multi-stemmed; bole up to 0.6 m in diameter; bark brown-
black; twigs and leaves turpentine-scented when crushed; sometimes with reddish
crenate leaf-galls or “witches’ broom” stem-galls up to 30 cm long. Leaves up to
10 cm long, parinnate, with the lower leaflets subopposite to alternate, one of the
terminal pair occasionally aborting thus making the leaf appear imparipinnate; rachis
narrowly winged; leaflets (2–)3–9 pairs, usually 3–4 paired in the Flora area oblong
to oblong-lanceolate or ovate, 1–5 cm long, 0.5–1.5( 2) cm broad, sessile, usually
R. W. Bussmann (*)
G. N. Njoroge

entire, apex obtuse to emarginate, base cuneate, coriaceous, conspicuously reddish

when young. Flowers yellowish or purplish; bracts ovate, about 2 mm long; brac-
teoles about 5, 0.7–1 mm long, sometimes ciliate. Inflorescences compact spikes or
racemes 1–5 cm long. Stamens 4–6. Drupe obliquely globose, 4–5 mm in diameter,
apiculate, green turning red, with mango-like smell when crushed (Kokwaro 1986).
Common in the undergrowth of dry, Juniperus procera dominated Afromontane
forests (Bussmann 2002a, b, 2006).
Pistacia aethiopica: Twigs are used as toothbrush. The root infusion is drunk as tea
(Beentje 1994). The leaves are boiled to treat common cold (Njoroge and Bussmann
2006). Pistacia integerrima is used to treat gout (Ahmad Jan et al. 2019). Pistacia
terebinthus smoke is used to cure eye diseases (Mohagheghzadeh and Faridi 2006).
The fruits of Pistacia atlantica are digestive (Muhammad et al. 2019), and the fruits
of Pistacia khinjuk are used for jaundice (Sher et al. 2016). The same species is used
to treat cough, asthma, gum used to remove phlegm, and the gum smoke for eye
problems (Wali et al. 2019).
Local Food Uses
Pistacia aethiopica: The seeds are eaten.
Pistacia aethiopica: The wood serves as poles, and the trunk yields excellent mastic
(Beentje 1994). The roots of Pistacia mutica yield a dye used for wool (Bussmann
et al. 2016, Mehdiyeva et al. 2017). Pistacia lentiscus smoke is used to purify the air
(Mohagheghzadeh and Faridi 2006). Pistacia khinjuk gum is used to mend broken
pots and as glue, the wood is used as firewood and often sold, and the leaves serve as
forage (Wali et al. 2019).
References
medicine research, prospects and limitations in Pakistan: a literature review. Acta Ecol Sin 2019.
https://doi.org/10.1016/j.chnaes.2019.12.005.
Pistacia aethiopica Kokwaro 833
2006;111:41–66.
Caucasus. J Ethnobiol Ethnomed. 2016;12:43. https://doi.org/10.1186/s13002-016-0110-2.
Kokwaro JO. Flora of tropical East Africa: Anacardiaceae. Rotterdam: Balkema; 1986.
Mehdiyeva N, Alizade V, Paniagua Zambrana NY, Bussmann RW. Pistacia mutica Fisch. &
C.A. Mey. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer Interna-
tional Publishing; 2017. https://doi.org/10.1007/978-3-319-49412-8_118.
appraisal of Kurram agency tribal area Pakistan. Indian J Tradit Knowl. 2019;184:631–47.
Ethnobot Res Appl. 2019;18:35. https://doi.org/10.32859/era.18.35.1-30.
Podocarpus milanjianus Rendle
Afrocarpus falcatus (Thunb.) C.N. Page
Afrocarpus gracilior (Pilg.) C.N. Page
PODOCARPACEAE

Synonyms
Afrocarpus falcatus (Thunb.) C.N. Page: Decussocarpus falcatus (Thunb.) de

Laub.; Nageia falcata var. falcatus (Pilg.) Silba; Podocarpus falcatus (Thunb.)
R. Br. ex Mirb.
Afrocarpus gracilior (Pilg.) C.N. Page: Afrocarpus falcatus subsp. gracilior (Pilg.)
Silba; Decussocarpus gracilior (Pilg.) de Laub.; Nageia falcata var. gracilior (Pilg.)
Silba; Podocarpus gracilior Pilg.
Podocarpus milanjianus Rendle: Afrocarpus falcatus subsp. milanjianus (Rendle)
Silba; Podocarpus ulugurensis Pilg.
R. W. Bussmann (*)
G. N. Njoroge
J. Grimshaw

Local Names
Afrocarpus falcatus: Kikuyu: Muthengera; Elgeyo/Marakwet/Tugen: Benet,

Kamba: Muvengea; Kipsigis/Sebei: Saptet; Maa: Olbiribiri; Meru: Mubiribiri,
Samburu/Somali: Dibiribiri; Turkana: Lotimat; English: Podo, east African yel-
lowwood (Beentje 1994; Gachati 1989; Kokwaro 2009).
Afrocarpus gracilior: Kikuyu: Muthengera; Chagga: Mvavavi (Beentje 1994;
Podocarpus milanjianus: Chagga: Mvavavi; Kikuyu: Muthengera; Kipsigis/
Nandi: Saptet, Lumbwa: Sadebet; Maa: Olpiripiri; Marakwet: Sosaita, Serti;
Meru: Mubiribiri; Samburu: Biribiriti; Sebei: Sitatet; Somali: Biribiriti; Taita:
Msembangala; Tugen: Septa; English: East African yellowwood, Podo (Beentje
Botany and Ecology
Afrocarpus falcatus (Thunb.) C.N. Page: Trees 10–25(–60) m tall and up to

210 cm diameter, often with a clear bole of more than 20 m. Bark gray, often
purplish, smooth at first, later flaking in rectangular to rounded plates. Twigs
round or quadrangular, ridged by decurrent leaf bases. Terminal buds about 1 mm
diameter, outer bud scales narrowly triangular, 2–2.5 mm long and 1 mm wide.
Juvenile leaves opposite, linear-lanceolate, up to 12 cm 6 mm. Adult leaves
spirally inserted, linear-lanceolate, (1–)2–4(–4.5) cm (1.2–)2–4(–6) mm, midrib
on lower surface and very slightly raised on upper surface, stomata present on both
surfaces, arranged in 14–20 distinct longitudinal lines on either side of midrib.
Pollen cones in groups of 1–4, subsessile, catkin-like, 5–13 2–3.5 mm, brownish,
microsporophylls broadly triangular-trullate, 0.6–0.8 mm long and 0.8–1.4 mm
wide, each bearing two pollen sacs 0.6–0.7 mm long and about 0.3–0.4 mm in
diameter. Seed cones solitary on scaly or leafy branches 7–27 mm long and 1.5–
2.5 mm in diameter, only the terminal scale fertile, epimatium verrucose, globose to
obovoid, 12–18 mm long, glaucous to grayish green, ripening to a yellow- or light
red-brown. Seed entirely enclosed by epimatium, globose, 10–12(–14) mm diameter
with a hard-smooth seed coat 1–2 mm thick. It closely resembles A. gracilior, which
has slightly bigger leaves (mostly 3–6 cm long), and slightly larger (to 18 mm long),
oval cones with smooth skin, while A. falcatus has spherical cones with verrucose
skin (like an orange) (Melville 1958). Very hard to germinate, and almost impossible
to store seeds (Bussmann and Lange 2000). Very slow growing species (Bussmann
1999).
Afrocarpus gracilior (Pilg.) C.N. Page: Trees to 40 m tall and 250 cm diameter,
forming a broad, domed crown in mature trees. Bark first smooth, becoming platy
and flaky in larger trees. Branchlets are more or less quadrangular within dense foliar
units. Leaves spirally arranged, straight or falcate, to 18 cm long in juvenile plants,
otherwise 3–6 cm long and 2–4 mm wide, gray-green, apex acute. Seed cones
Podocarpus milanjianus Rendle Afrocarpus falcatus (Thunb.). . . 837
solitary on branchlets, at maturity, a single seed covered by a fleshy epimatium that

ripens from green to yellow or orange, oblong, pear-shaped or spherical, 15–20 mm
long, the seed has a hard, smooth coat. In appearance it closely resembles A. falcatus,
which has slightly smaller leaves (mostly 2–4 cm long), and slightly smaller
(to 15 mm long), spherical cones with verrucose skin (like an orange). It occurs at
elevations of about 2100–2700 m (Melville 1958) (Figs. 1, 2, and 3).
Podocarpus milanjianus Rendle: Tree up to 35 m tall with thin flaking reddish-
brown bark. Leaves 2–15 cm 5–12 mm, shortly petiolate, spreading, strap-shaped,
margin long-attenuate in the upper half, stomata on the under surface only, juvenile
leaves similar but often longer and slightly broader. Male cones solitary or paired, up
to 5 cm long, pinkish. Male cones solitary or paired, up to 5 cm long, pinkish.
Female strobili solitary, fertile scales 1 or 2, seed at maturity 1 cm long, sub-globose,
drupe-like, green to purple, surmounting a fleshy, red to purple, glaucous receptacle,
testa thin and brittle, enclosed in a thin, very resinous integument. Female strobili
solitary, fertile scales 1 or 2, seed at maturity 1 cm long, sub-globose, drupe-like,
Fig. 1 Afrocarpus gracilior

(Podocarpaceae), Oromiya
Province, Ethiopia. (Photo
J. Grimshaw)
Fig. 2 Afrocarpus gracilior

(Podocarpaceae), Oromiya
Province, Ethiopia. (Photo
J. Grimshaw)
green to purple, surmounting a fleshy, red to purple, glaucous receptacle, testa thin
and brittle, enclosed in a thin, very resinous integument. Podocarpus milanjanus
occurs in mountain forest from southern Sudan through eastern Central Africa and
East Africa to Zimbabwe, Mozambique, Swaziland and eastern and southern
South Africa, where it descends to sea level. It is also found in the highlands of
south-eastern Nigeria and western Cameroon and has been recorded from an insel-
berg in Equatorial Guinea. It is occasionally planted as ornamental tree within its
natural area of distribution (Melville 1958). Germinates well both in light and dark
conditions, but hard to store (Bussmann and Lange 2000). Dominant component in
many Afromontane forests (Bussmann 2002a, b, 2006a, b), and fast growing
(Bussmann 1999) (Figs. 4, 5, 6, 7, 8, and 9).
Fig. 3 Afrocarpus gracilior (Podocarpaceae), leaves, Oromiya Province, Ethiopia. (Photo

J. Grimshaw)
Fig. 4 Podocarpus milanjanus (Podocarpaceae), open Podocarpus – Bamboo forest, Mt. Kenya
National Park, Kenya. (Photo R.W. Bussmann)
Fig. 5 Podocarpus
milanjanus (Podocarpaceae),
leaves, Mt. Kenya National
Park, Kenya. (Photo
R.W. Bussmann)
Fig. 6 Podocarpus
female cones, Mt. Kenya
R.W. Bussmann)
Fig. 7 Podocarpus
leaves, Mt. Kenya National
Park, Kenya. (Photo
R.W. Bussmann)
Fig. 8 Podocarpus
female branch, Mt. Kenya
R.W. Bussmann)
Fig. 9 Podocarpus
male cone, Mt. Kenya
R.W. Bussmann)
Afrocarpus falcatus: Bark infusion used for stomach problems (Kokwaro 2009).
Root decoction taken orally or applied to skin for cancer (Teklehaymanot 2009).
Afrocarpus gracilior: Bark infusion used for stomach problems (Kokwaro 2009).
The leaf juice is used against vomiting (Teklehaymanot et al. 2007).
Podocarpus milanjanus: In Kenya a bark infusion is taken to treat stomachache
(Beentje 1994). In Uganda a leaf decoction is taken as vermifuge.
Local Food Uses
Podocarpus milanjanus: People occasionally eat the fleshy receptacles at the base
of the seeds (Beentje 1994).
Afrocarpus falcatus: Good timber tree (Beentje 1994).

Afrocarpus gracilior: Good timber tree (Beentje 1994).
Podocarpus milanjanus: The wood, often traded as “podo,” is highly valued for
furniture and ship building, for example, for masts and planks. It is also used for
poles, paneling, boxes, veneer, and plywood. It is popular for making butchers’
blocks because it is fairly hard, without scent, and does not chip easily. It is suitable
for construction, flooring, joinery, interior trim, vehicle bodies, railway sleepers,
toys, novelties, agricultural implements, musical instruments, coffins, food con-
tainers, vats, carving, pattern making, matches, turnery, hardboard, and particle
board. It is also used as fuelwood, and it is considered a high-quality pulpwood. In
the Mau region in Kenya the bark is used as waterproof cover for beehives (Beentje
1994). Used for construction and firewood (Bussmann et al. 2006). The tree is
planted as ornamental and roadside tree, having a characteristic appearance with
its conical shape and bright green glossy leaves, and also as shade tree in coffee,
cocoa and banana plantations. It is the national tree of South Africa.
References
reference to those of Mount Kenya. J East Afr Nat Hist. 1999;881-2:69–78.
Appendices 1–7. www.naturekenya.org/JournalEANH.htm
2006b;111:41–66.
Nat Hist. 2000;891-2:101–11.
the Maasai of Sekenani Valley, Maasai Mara, Kenya. Journal of Ethnobiology and Ethnomed-
icine 2006;2:22.
Melville R. Flora of Tropical East Africa: Gymnospermeae. London: Crown Agents for Overseas
Governments & Administrations; 1958. 16 pp. ISBN 978-1-84246-268-3
Portulaca oleracea L.
PORTULACACEAE

Grace N. Njoroge
Synonyms
Portulaca oleracea L.: Portulaca consanguines Schltdl.; Portulaca intermedia Link;

Portulaca marginata Kunth; Portulaca neglecta Mack. & Bush; Portulaca
officinarum Crantz; Portulaca oleracea subsp. granulatostellulata (Poelln.) Danin &
H.G. Baker; Portulaca oleracea subsp. nicaraguensis Danin & H.G. baker; Portulaca
oleracea subsp. nitida Danin & H.G. Baker; Portulaca oleracea subsp. sylvestris
Thell.; Portulaca oleracea var. granulatostellulata Poelln.; Portulaca oleracea var.
macrantha Eggers; Portulaca oleracea var. micrantha Eggers; Portulaca oleracea
var. opposita Poelln.; Portulaca oleracea var. parvifolia (Haw.) Grieseb.; Portulaca
olitoria Pall.; Portulaca parvifolia Haw.; Portulaca pilosa L.; Portulaca pusilla
Kunth; Portulaca retusa Engelm.; Portulaca sativa Haw.; Portulaca suffruticosa
Thwaites
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Portulaca oleracea: Kikuyu: Gatumia; Rufiji: Mkoga; Ndorobo: Loirabirab

(Gachati 1989; Kokwaro 2009); English: Purslane
Botany and Ecology
Portulaca oleracea L.: Annual. The stem (7)10–30 cm long, glabrous, fleshy,
prostrate, mostly appressed to the soil or ascending, branched from base. The leaves
are alternate, sessile, cuneately narrowing towards the base, fleshy. The flowers are
solitary or in clusters of 2 or 3, in the stem crotches or in the leaf axils. The sepals are
herbaceous, obtusely keeled. The petals (4–6) are free or connate at base, united with
the calyx, obovate, and yellow. The stigmas are linear. The stamens 8–15, free or
adnate at base to petals. The capsule is ovoid or globular, 5–8 mm long. The plant
can be found in sand deposits on riverbanks, fields, weed-infested places around
villages, and gardens. In ruderal habitats, near roads and cultivated fields. Up to
middle mountain belt, on an elevation up to 1200 m. Flowering June–July, fruits
from July to September (Figs. 1, 2, and 3).
Phytochemistry
Portulaca oleracea: Alkaloids, glycosides, saponins, catecholamines (noradrena-

line, dopamine), oxalic acids, coumarins, flavonoids, cardiac and anthraquinone
glycosides, proteins, fatty acids.
Fig. 1 Portulaca oleracea

(Portulacaceae), Tusheti,
Georgia. (Photo
R.W. Bussmann and
Portulaca oleracea L. 847

Georgia. (Photo
R.W. Bussmann and

Georgia. (Photo
R.W. Bussmann and
Portulaca oleracea: The roots are used to treat snakebites, either fresh or pounded
and soaked in water. Also applied to wounds and burns (Kokwaro 2009). In Kenya,
the plant serves to treat typhoid (Njoroge et al. 2004). The whole plant is used to treat
urinary infections and burns; leaves are used to treat fever and as vermifuge
(Bussmann et al. 2018a, b). The whole fresh plant is used to treat fever and liver
problems (Béjar et al. 2001; Bussmann and Sharon 2006a, 2007a; Paniagua-
Zambrana et al. 2020). The whole fresh plant is used to treat liver, kidneys infections,
hepatitis, inflammation of the liver, and cleansing of the liver (Bussmann and Sharon
2006b, 2007b, 2015a, b; Bussmann and Glenn 2010a, b; Paniagua-Zambrana et al.
2020). Often ingredient in herb mixtures (Bussmann et al. 2010). In Pakistan,
Portulaca is used to treat liver infections, as anthelmintic, for jaundice, asthma, and
as diuretic (Umair et al. 2019; Yeşil et al. 2019). In India, it is used to remedy
jaundice, liver, and kidney problems (Verma et al. 2007).
Local Food Uses
Portulaca oleracea: Leaves and shoots are cooked and eaten as vegetable (Kokwaro
2009). Leaves are used as food (Bussmann et al. 2018). The species is widely
consumed in Eurasia, especially the Caucasus (Batsatsashvili et al. 2017; Bussmann
2017; Bussmann et al. 2016, 2018; Yeşil et al. 2019). Also eaten in Ethiopia, together
with Portulaca quadrifida (Balemie and Kebebew 2006; Bussmann et al. 2011).
Portulaca oleracea: Sometimes eaten by livestock (Bussmann et al. 2011).
References
Tchelidze D, Aleksanyan A, Alizade V, Paniagua Zambrana NY, Bussmann RW. Portulaca
oleracea L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer Interna-
tional Publishing; 2017.
Medicinales del Sur Ecuatoriano. San Diego: Latino Herbal Press; 2001, 340p.
2017. XXVII, 746p. ISBN 978-3-319-49411-1.
Bussmann RW, Glenn A. Peruvian medicinal plants for the treatment of liver and gallbladder
ailments. Arnaldoa. 2010a;17(2):243–54.
Bussmann RW, Glenn A. Medicinal plants used in Northern Peru for reproductive problems and
female health. J Ethnobiol Ethnomed. 2010b;6:30.
9789962-2-2.
978-0-9789962-3-9.
medicinal flora of Northern Peru. St. Louis: William L. Brown Center, MBG; 2015a. ISBN
978-0-9960231-2-2.
Portulaca oleracea L. 849
9960231-3-9.
Bussmann RW, Glenn A, Meyer K, Rothrock A, Townesmith A. Herbal mixtures in traditional
medicine in Northern Peru. J Ethnobiol Ethnomed. 2010;6:10.
(Sakartvelo), Caucasus. J Ethnobiol Ethnomed. 2016;12:43. https://doi.org/10.1186/s13002-
016-0110-2.
Sakartvelo (Republic of Georgia), Caucasus. Indian J Tradit Knowl. 2018a;17(1):7–33.
Bussmann RW, Paniagua-Zambrana NY, Romero C, Hart RE. Astonishing diversity – the medicinal
plant markets of Bogotá, Colombia. J Ethnobiol Ethnomed. 2018b;14(1):43. https://doi.org/
10.1186/s13002-018-0241-8.
Paniagua Zambrana NY, Bussmann RW, Romero C. Portulaca oleracea L. In: Paniagua Zambrana
NY, Bussmann RW, editors. Ethnobotany of mountain regions – ethnobotany of the Andes.
Cham: Springer International Publishing; 2020. https://doi.org/10.1007/978-3-319-77093-
2_21-1.
10.1186/s13002-019-0285-4.
Yeşil Y, Çelik M, Yılmaz B. Wild edible plants in Yeşilli (Mardin-Turkey), a multicultural area.
J Ethnobiol Ethnomed. 2019;15:52. https://doi.org/10.1186/s13002-019-0327-y.
Pouteria adolfi-friederici (Engl.) A. Meeuse
Pouteria altissima (A. Chev.) Baehni
SAPOTACEAE

Grace N. Njoroge
Synonyms
Pouteria adolfi-friederici (Engl.) A. Meeuse: Aningeria adolfi-friedericii (Engl.)

Robyns & Gilbert; Sideroxylon adolfi-friedericii Engl.
Pouteria altissima (A. Chev.) Baehni: Aningeria altissima (A. Chev.) Aubrév. &
Pellegr.; Hormogyne altissima A.Chev.; Hormogyne gabonensis A.Chev.; Pouteria
giordani Chiov.; Rhamnoluma altissima (A. Chev.) Baehni; Sideroxylon altissimum
(A. Chev.) Hutch. & Dalziel; Sideroxylon gabonense (A. Chev.) Lecomte ex Pellegr.
Local Names
Pouteria adolfi-friederici: Kikuyu: Muna; Kipsigis: Cheptatet; Marakwet: Seite;

Meru: Mutunguru; Sebei: Lulyo, Luliondet; Tugen: Ngecheberet, Nejebbe;
Ndorobo: Kipworbet, Nkilejo (Beentje 1994; Gachati 1989; Kokwaro 2009);
English: Aningeria
Pouteria altissima: English: Aningeria
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Pouteria adolfi-friederici (Engl.) A. Meeuse: Tall tree, height up to 50 m, with long

straight fluted bole and buttressed base. Young shoots with dense ferrugineous
pubescence; older branches blackish-brown and subglabrous. Petioles 1–2 cm long,
twisted, with ferrugineous or grayish-brown pubescence. Leaf-lamina elliptic to
oblong or obovate-elliptic, 4–21.5 cm long, 2–8.5 cm wide, pellucid dots visible
only in thin-textured sapling or youngest shade leaves, apex acute or rounded, base
narrowly to broadly cuneate and sometimes decurrent with petiole, margin not or
strongly inrolled; lower surface with varying density of ferrugineous pubescence,
wearing away or stripping and becoming glabrous and with hairs on midrib and
nerves only; lateral nerves 10–25 each side. Flowers fascicled in current or fallen leaf
axils; pedicels 5–10 mm long, densely pubescent. Sepals 4 or 5, ovate to more or less
oblong, up to 6 mm long, 3 mm wide, pubescent externally with hairs of variable
density and size. Corolla cream; tube up to 6.5 mm long; lobes ovate or rounded,
up to 2 mm long. Free part of filaments up to 1.5 mm long. Staminodes subulate or
sometimes expanded and petaloid, up to 1.5 mm long. Style up to 6.5 mm long.
Fruits greenish, narrowly ellipsoid, up to 4 cm long, apex with short about 1 cm long
beak, pubescent or puberulous with short ferrugineous hairs. Seed ovoid to narrowly
ellipsoid, up to 3 cm long; testa shiny brown; scar lateral, pale, elliptic. Pouteria
adolfi-friedericii occurs from eastern DR Congo, southern Sudan, and south-western
Ethiopia south to eastern Zimbabwe (Hemsley 1968). Pouteria adolfi-friederici, in
vegetation science much better known under its old name Aningeria adolfi-
friederici, is a characteristic species of the upper tree layer of wet, Ocotea
usambarensis and Syzygium guineense dominated Afromontane forests (Bussmann
2002a, b, 2006). The species germinates well, but only under light conditions
(Bussmann and Lange 2000) (Figs. 1, 2, and 3).
Pouteria altissima (A. Chev.) Baehni: Tall tree, up to 50 m, with deeply fluted bole
and spreading crown; bark of trunk pale creamy gray, irregularly and shallowly
Fig. 1 Pouteria adolfi-

friederici (Sapotaceae), Bale
Pouteria adolfiriederici (Engl.) A. Meeuse. . . 853

friederici (Sapotaceae), Bale

friederici (Sapotaceae), bark
cut, Bale Mountains National
fissured. Young stems yellowish puberulous, soon glabrescent, older twigs dark gray
with wrinkled surface. Leaves membranous, pellucid-punctate, not confined to
branch ends. Lamina 4–16 cm 2·5–6 cm, ovate or obovate-oblong, apex rounded
or more often cuspidate with blunt to emarginate apiculum, base rounded or
cuspidate; petioles up to 1 cm long. Lateral nerves 11–23 on each side. Leaves

completely glabrous or with midrib and lateral nerves of lower surface lanate-
puberulous. Pedicels up to 1 cm long. Calyx c. 4 mm long, sepals elliptic or oblong.
Corolla 5–6 mm long, greenish-yellow. Anthers 1 mm long. Style up to 4 mm long.
Fruit up to 2 cm in diameter and red when ripe, obovoid to subglobose. Seed up to
1.5 cm long. Pouteria altissima occurs from Guinea east to south-western Ethiopia,
western Kenya, and north-western Tanzania, and south to northern Zambia
(Hemsley 1968).
Pouteria adolfi-friederici: In Ethiopia, the fruits are used as a traditional taenidial

drug, usually as a paste in barley porridge. The bark decoction is used for stomach-
ache (Kokwaro 2009). In Ethiopia, the species is used for its antibacterial properties
(Desta 1995; Lulekal et al. 2008).
Local Food Uses
Pouteria adolfi-friederici: The sweet fruit pulp is eaten raw. The seed oil can be
eaten (Beentje 1994).
Pouteria adolfi-friederici: In East Africa, the wood is valued for furniture. It is also
suitable for light construction, light flooring, interior trim, joinery, cabinet work,
boats, vehicle bodies, boxes and crates, veneer and plywood, and pulpwood. It is
used as firewood and for charcoal production (Beentje 1994). The tree is used as a
shade tree in plantations. Often used as firewood (Bussmann et al. 2011).
Pouteria altissima: The wood (trade names: aningré, aniégré, anigré, asanfena,
asanfona, osan, mukangu) is especially recommended for high-quality sliced and
peeled veneer. In West Africa, it is also used for light carpentry, interior joinery, high-
class furniture, and molding. In East Africa, it is considered suitable for the same
purposes and additionally for light construction, vehicle bodies, musical instru-
ments, boxes and crates, railway sleepers, toys and novelties, turnery, and pulpwood
for paper production. It is also used as firewood and for the production of charcoal.
In Ethiopia and Uganda, Pouteria altissima is used as a shade tree in coffee, banana,
and cocoa plantations, and it is considered useful as a bee plant. It is sometimes
planted as roadside tree.
Pouteria adolfiriederici (Engl.) A. Meeuse. . . 855
References
Bussmann RW. Islands in the desert–forest vegetation of Kenya’s smaller mountains and highland
2006;111:41–66.
Nat Hist. 2000;89(1–2):101–11.
Hemsley JH. Flora of Tropical East Africa: Sapotaceae. London: Crown Agents for Overseas
Governments & Administrations; 1968. 78 pp. ISBN: 978-1-84246-313-0.
Angetu District, Southeastern Ethiopia. J Ethnobiol Ethnomed. 2008;4:10. https://doi.org/
10.1186/1746-4269-4-10.
Prunus africana (Hook. f.) Kalkman
ROSACEAE

Grace N. Njoroge
Synonyms
Prunus africana (Hook. f.) Kalkman: Pygeum africanum Hook. f.; Pygeum
crassifolium Hauman
Local Names
Prunus africana: Kikuyu: Muiri; Matengo: Migambo; Chagga: Mkonde-Konde;

Vinza: Mfubia; Luganda: Ntasesa; Maa: Ol-Koijuk; Kamba: Mutimailu; Kisii:
Kiburabura; Luhya: Mwiritsa; Elgeyo/Kipsigis/Ndorobo: Tenduet; Marakwet:
Tendwet; Meru: Mweria; Samburuz: Lemalan; Sebei: Amaatet, Oromoti;
Watende: Murugutu; English: Red stinkwood (Beentje 1994; Gachati 1989;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Prunus africana (Hook. f.) Kalkman: A much-branched evergreen tree 10 m (up to

25) tall, or rarely a shrub 3–5 m tall, entirely glabrous except for the flowers. Bark
dark brown to gray, rugged. Leaves petiolate; lamina (4) 6–11 (15) cm 2–4 (5·5)
cm, elliptic, lanceolate-elliptic or oblong-lanceolate, glabrous, coriaceous or sub-
coriaceous, the apex obtuse to subacuminate, the base broadly cuneate to rarely
rounded, margins coarsely crenate-serrate to subentire with dark glandular dots in
the incisions (the most proximal gland on one or both margins of the leaf sometimes
conspicuous); petiole 1–2 cm long, channelled, often reddish; stipules linear, 1.5–
2.0 cm long, caducous. Racemes usually solitary, 2–5 (8) cm long, 7–15 flowered,
arising from the axils of scales at the base of lateral shoots which may also produce
leaves in their upper part; peduncle usually 5–10 mm long; pedicels (3) 5–7 (10) mm
long; bracts triangular, small, soon caducous. Perianth (4)5(6)-merous, sometimes
irregularly so. Calyx-tube more or less cyathiform, (3.0) 3.5–4.0 (5.0) mm in
diameter at the mouth, rather fleshy, glabrous outside, usually hairy inside; calyx-
lobes 1–1.5 mm long, triangular, glabrous except for the ciliate apex. Petals up to
2 mm long, creamy-white, elliptic to oblong, reflexed, hairy abaxially especially
towards their margins and sometimes adaxially also. Stamens 25–35; filaments
1.5 mm long, glabrous; anthers didymous, 0.5–1.0 mm long. Ovary ovoid, sparsely
longhaired; style 1.5 mm long, sparsely haired; stigma peltate, slightly 2- or 3-lobed.
Fruit 5–8 mm 8–12 mm, transversely ellipsoid, broader than long, slightly
didymous and thus appearing as if 2-locular, dry, usually glabrous, red to purplish-
brown (Graham 1960). Common in the drier parts of Afromontane forests
(Bussmann 2002a, b, 2006a, b; Bytebier and Bussmann 2000; Navarro-Cerillo
et al. 2008). Heavily overharvested for medicinal purposes and wood trade
(Bussmann 2004; Hamilton 2004; Onyango et al. 2009; Stewart 2003) (Figs. 1, 2,
3, and 4).
Fig. 1 Prunus africana

(Rosaceae), Bale Mountains
National Park, Demaro forest.
Prunus africana (Hook. f.) Kalkman 859


Prunus africana: The leaves are steamed in water and the vapors inhaled for fever.
The leaf infusion is used to improve the appetite and for malaria (Beentje 1994;
Bussmann 2006a, b; Kokwaro 2009; Njoroge and Bussmann 2006), also to treat
toothache and ear infections (Giday et al. 2009), as well as skin itches and pimples
(Njoroge and Bussmann 2007), back pain in women, and HIV/AIDS (Njoroge and
Bussmann 2009). The species serves top treat prostate problems and urinary infec-
tions (Muthee et al. 2011). It is used to increase appetite, treat stomachache,
infections, and wounds (Nankaya et al. 2019, 2020).

Prunus africana: A bark extract is used as purgative for cattle (Kokwaro 2009).
Timber for construction and furniture (Beentje 1994). The bark is used to hang
beehives (Bussmann et al. 2011) and sometimes as firewood (Kiefer and Bussmann
2004, 2008).
References
Mountain Forests: The role of natural and anthropogenic disturbance. Lyonia 2004;61:98–111.
Prunus africana (Hook. f.) Kalkman 861
2006b;111:41–66.
doi.org/10.1186/1746-4269-5-34.
Hamilton AC. Medicinal plants, conservation and livelihoods. Biodivers Conserv. 2004;13:1477–
517.
2004;62:161–9.
Navarro-Cerillo RM, Clemente M, Padrón EVA, Hernandez-Bermejo E, García-Ferrer A, Kasimis
N. Forest structure in harvested sites of Afromontane forest of Prunus africana (Hook. f.)
Kalkm., in Bioko (Equatoiriual Guinea). Afr J Ecol. 2008;46(4):620–30.
Knowl. 2009;82:262–9.
Ethnomed. 2006;2:8.
Onyango JC, Nyunja RAO, Bussmann RW. Conservation of biodiversity in the East African
Tropical Forest. Lyonia. 2004;72:151–7.
Stewart KM. The African cherry (Prunus africana): can lessons be learned from an over-exploited
medicinal tree? J Ethnopharmacol. 2003;89:3–13.
Psidium guajava L.
MYRTACEAE

Grace N. Njoroge
Synonyms
Psidium guajava L.: Guajava pyrifera (L.) Kuntze; Myrtus guajava (L.) Kuntze;
Myrtus guajava var. pyrifera (L.) Kuntze; Pisdium guajava var. cujavillum (Burman)
Krug & Urb.; Pisdium guajava L. var. guajava; Pisdium guava Grieseb.; Psidium
guayava Raddio; Psidium igatemyense Barb. Rodr.; Psidium igatemyensis Barb.
Rodr.; Psidium pomiferum L.; Psidium pumiferum L.; Psidium pumilum Vahl;
Psidium pumilum var. guadalupense DC.; Psidium pyriferum L.
Local Names
Psidium guajava: Kikuyu: Mubera; Luo: Mapera (Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Psidium guajava L.: Trees, to 13 m tall. Bark gray, smooth, peeling in strips.
Branchlets angular, pubescent. Petiole ca. 5 mm; leaf blade oblong to elliptic,
6–12 cm 3.5–6 cm, leathery, abaxially pubescent, adaxially slightly rough,
secondary veins 12–15 on each side of midvein and usually impressed, reticulate
veins obvious, base rounded, apex acute to obtuse. Flowers solitary or 2 or 3 in
cymes. Hypanthium campanulate, ca. 5 mm, pubescent. Calyx cap nearly
rounded, 7–8 mm, irregularly opening. Petals white, 1–1.4 cm. Stamens 6–
9 mm. Ovary adnate to hypanthium. Style as long as stamens. Berry globose,
ovoid, or pyriform, 3–8 cm, with persistent calyx lobes at apex; flesh white or
yellow; placenta reddish, well developed, fleshy. Seeds many. Flowering summer
(Wu et al. 1994–2013). Psidium guajava is a very aggressive and invasive pioneer
species found in all kind of disturbed habitats, including lava flows (Fonge et al.
2005) (Figs. 1, 2, 3, 4, and 5).
Fig. 1 Psidium guajava

(Myrtaceae), garden, Chicani,
Bolivia. (Photo: R. W.
Bussmann and N. Y.
Paniagua-Zambrana)
Psidium guajava L. 865

(Myrtaceae), flowers, garden,
Chicani, Bolivia. (Photo:
R. W. Bussmann and N. Y.
Paniagua-Zambrana)

(Myrtaceae), young fruits,
Psidium guajava: The leaf decoction is drunk to treat jaundice (Kokwaro 2009), and
for common cold and asthma (Njoroge and Bussmann 2006a), diarrhea and dysen-
tery, cough, malaria, and stomachache (Randrianarivony et al. 2017; Rakotoarivelo
et al. 2015; Razafindraibe et al. 2013). Some antibiotic activity has been reported
(Malik et al. 2015). The leaves are also boiled to treat diarrhea, and for cancer and
liver problems (Bussmann and Sharon 2006). Tender shoots are taken directly either
by chewing or paste for the treatment of diarrhea, dysentery, piles, and vomiting
(Debbarma et al. 2017). The fruits are used as laxative, for colic, astringent, and for

(Myrtaceae), young fruits,

(Myrtaceae), fruits, Chicani,
Bolivia. (Photo: R. W.
Bussmann and N. Y.
Paniagua-Zambrana)
constipation (Kunwar et al. 2010, 2015). Young shoots are used to treat dysentery,
diarrhea, and stomach pain (Raj et al. 2018).
Psidium cattleyanum leaves are also used for diarrhea and stomach problems
(Rabearivony et al. 2015; Rakotoarivelo et al. 2015).
Local Food Uses
Psidium guajava: Fruits are edible.

Psidium guajava L. 867
Psidium guajava: The leaves are boiled, and the decoction is used to treat diarrhea in
cattle (Njoroge and Bussmann 2006b).
References
Fonge BA, Yinda GS, Focho DA, Fongod AGN, Bussmann RW. Vegetation and soil status on an
80-year old lava flow of Mt. Cameroon, West Africa. Lyonia. 2005;81:17–39.
Kunwar RM, Burlakoti C, Chowdhary CL, Bussmann RW. Medicinal plants in farwest Nepal: their
indigenous uses and pharmacological validity. Med Aromat Plant Sci Biotechnol. 2010;4(1):28–42.
medicines in farwest Nepal. J Ethnopharmacol. 2015;163:210–9.
018-0208-9.
2017; https://doi.org/10.1186/s13002-017-0147-x.
2013;9:73.
Psychotria orophila E.A.A. Petit
Psychotria sp.
RUBIACEAE

Grace N. Njoroge
Synonyms
Psychotria orophila E.A.A. Petit: Grumilea elliottii K. Schum & K. Krause
Local Names
Psychotria orophila: Kikuyu: Mukomakoma; Maa: Olobarat (Beentje 1994;

Botany and Ecology
Psychotria orophila E.A.A. Petit: Shrub or small tree 1.5–10 m tall, with glabrous
stems. Leaf-blades elliptic or oblong-elliptic, 7–18 cm long, 3.5–7 (–10) cm wide,
acute, obtuse or slightly acuminate at the apex, cuneate at the base, glabrous above,
R. W. Bussmann (*)
G. N. Njoroge

pubescent beneath on the midnerve and sometimes on the lateral nerves as well, or
entirely glabrous beneath save for the hairy domatia; nodules absent; petiole 0.5–
3 cm long, glabrous or pubescent; stipules ovate or ovate-elliptic, 1–2 (–2.5)
cm. long, shortly bifid for 1–2 mm or entire, glabrous save for a few marginal
hairs, deciduous. Flowers heterostylous, 5-merous, in much-branched panicles,
lowest branches usually 4; peduncles 2–7 (–15) cm long, glabrous or slightly
pubescent; secondary branches 0.3–3.5 cm long, ferruginous pubescent; pedi-
cels obsolete or 1–1.5 mm long in fruit; main bracts lanceolate with sheathing
lobed bases, 5 mm long, ciliate; other bracts and bracteoles smaller, ferruginous
pubescent. Calyx-tube obconic, 1 mm long, glabrous; limb cupuliform, 1–1.5 mm
long, glabrous, lobes very short or obsolete, margined with ferruginous hairs.
Corolla white or salmon-tinged above in bud, white when developed; tube 2.5–
5 mm long; lobes oblong-elliptic, 1.8–4 mm long, 0.8 mm wide, thickened at the
apices. Stamens with filaments 1.5 mm long in short-styled flowers and 0.5 mm long
in long-styled flowers. Style 2.5 mm long in short-styled flowers and 3.5–7 mm long
in long-styled flowers; stigma-lobes 0.5–1 mm long. Drupes red, with 2 pyrenes,
ellipsoid, 6–7 mm long, 4–5.5 mm wide; pyrenes semi-ellipsoid, 6 mm long, 4.5 mm
wide, 2 mm thick; dorsal face 5-grooved. Seeds blackish red, semiglobose, 4 mm
long, 3.5 mm wide, 1.5 mm thick; ventral face plane, dorsal face 5-grooved; albumen
ruminate. In evergreen forest; 1650–2670 (–3400) m (Verdcourt 1976; Bridson and
Verdcourt 1988, 1991; Bussmann 2002a, b, 2006a).
In DR Congo, Psychotria is used medicinally against intestinal diseases and worms.

In eastern Africa, the rhizomes are used to treat snakebites; fresh rhizomes are
chewed and the juice swallowed, or they may be pounded and the infusion used
both orally and externally on the wound (Beentje 1994). The leaves of Psychotria
kirkii and Psychotria orophila are boiled to treat malaria and boiled for strengthen-
ing soup (Bussmann 2006b).
Local Food Uses
Psychotria sp.: In Malawi, the leaves are cooked with ashes to prepare a slimy
vegetable called “thelele,” which is particularly liked by older people.
The leaves of Psychotria kirkii and Psychotria orophila are browsed by livestock
the wood serves for fence construction and firewood; the young branches make good
arrows (Bussmann 2006b).
Psychotria orophila E.A.A. Petit. . . 871
References
Bridson DM, Verdcourt B. Flora of Tropical East Africa: Rubiaceae (part 2). Rotterdam: Balkema;
1988.
Bridson DM, Verdcourt B. Flora of Tropical East Africa: Rubiaceae (part 3). Rotterdam: Balkema;
1991.
2006a;111:41–66.
Verdcourt B. Flora of Tropical East Africa: Rubiaceae (part 1). London: Crown Agents for Overseas
Pterolobium stellatum (Forssk.) Brenan
FABACEAE

Grace N. Njoroge
Local Names
Pterolobium stellatum: Kikuyu: Mutanda-Mbogo, Mutanga-Ruri; Kipsigis/

Ndorobo: Kipkososit; Luo: Osirimadongo; Maa: Engisar ngadiun; Nandi: Korkos;
Tugen: Kisibiri, Kormande; Meru: Kibai (Beentje 1994; Gachati 1989; Kokwaro
2009).
Botany and Ecology
Pterolobium stellatum (Forssk.) Brenan: Scrambling or climbing shrub up to 15 m

high, seldom semierect. Stems about densely pubescent at least when young, armed
with reflexed prickles up to 6 mm long, in pairs at the nodes and often with scattered
ones between the nodes. Leaves about densely puberulous when young, sometimes
becoming glabrescent with age: petiole 1.5–3.6 cm long, rhachis 5–15 (18) cm long
(in our area), armed on the lower side with paired reflexed prickles at the insertions
R. W. Bussmann (*)
G. N. Njoroge

of the pinnae and occasionally with single prickles in between the pinnae, and on the
upper side often with straight ascending prickles singly at the insertions of the
pinnae, pinnae 5–13 pairs, rhachillae 2–6 cm long, leaflets 7–15 pairs per pinna,
(4) 6–10 (12) mm long, 2–4 mm wide, narrowly oblong or elliptic-oblong, with the
terminal ones about obovate, rounded to slightly emarginate apically, lower surface
puberulous to pubescent or glabrous, upper surface glabrous or sometimes
puberulous. Racemes 5–18 cm long, terminal and axillary, aggregated into panicles
up to 35 cm long, puberulous to shortly pubescent, bracts subulate, rapidly decidu-
ous. Flowers sweetly scented, pale yellowish-white, on pubescent pedicels 3–6 mm
long. Sepals 2–3 mm long, 0.75–1.4 mm wide, greenish, pubescent basally at least,
Scrambling or climbing shrub up to 15 m high, seldom semierect. Stems-about
densely pubescent at least when young, armed with reflexed prickles up to 6 mm
long, in pairs at the nodes and often with scattered ones between the nodes. Leaves
about densely puberulous when young, sometimes becoming glabrescent with age:
petiole 1.5–3.6 cm long, rhachis 5–15 (18) cm long (in our area), armed on the lower
side with paired reflexed prickles at the insertions of the pinnae and occasionally
with single prickles in between the pinnae, and on the upper side often with straight
ascending prickles singly at the insertions of the pinnae, pinnae 5–13 pairs,
rhachillae 2–6 cm long, leaflets 7–15 pairs/pinna, (4) 6–10 (12) mm long, 2–4 mm
wide, narrowly oblong or elliptic-oblong, with the terminal ones about obovate,
rounded to slightly emarginate apically, lower surface puberulous to pubescent or
glabrous, upper surface glabrous or sometimes puberulous. Racemes 5–18 cm long,
terminal and axillary, aggregated into panicles up to 35 cm long, puberulous to
shortly pubescent, bracts subulate, rapidly deciduous. Flowers sweetly scented, pale
yellowish-white, on pubescent pedicels 3–6 mm long. Sepals 2–3 mm long, 0.75–
1.4 mm wide, greenish, pubescent basally at least, usually about reflexed in flower.
Petals about the same size as the sepals, oblanceolate-oblong. Stamens all exserted,
4–6 mm long. Ovary densely pubescent. Pods red to scarlet when young but
becoming brown with age, 3–5 cm long (including the wing), wing 0.9–1.6 cm
wide, pubescent, puberulous or at maturity about glabrescent. Seeds olive, 9–11 mm
long, 5–6.5 mm wide, ellipsoid (Brenan 1959). Common in heavily browsed,
disturbed, Afromontane forest glades (Bussmann 2002) (Figs. 1, 2, 3, and 4).
Pterolobium stellatum: Root decoction used against stomachache (Maasai) (Beentje

1994). The roots are also used to treat snakebites. Leaves in decoction serve for
tuberculosis and chest illnesses (Bekalo et al. 2009; Kokwaro 2009). The root is
boiled to treat diarrhea (Flatie et al. 2009), and to treat ear, nose, and throat diseases
(Njoroge and Bussmann 2006). The plant is also used to treat back pain in women
(Njoroge 2009).
Pterolobium stellatum (Forssk.) Brenan 875
Fig. 1 Pterolobium stellatum

(Fabaceae), flowering
specimen, Mt. Kenya National
Park, disturbed lower

(Fabaceae), branches,
disturbed lower Afromontane
forest, Kenya. (Photo: R. W.
Bussmann)

(Fabaceae), branches,
disturbed lower Afromontane
forest, Kenya. (Photo: R. W.
Bussmann)

(Fabaceae), fruits, Mt. Kenya
National Park, disturbed lower
Pterolobium stellatum: Browsed by wildlife (Bussmann 2006).
References
doi.org/10.1186/1746-4269-5-26.
Pterolobium stellatum (Forssk.) Brenan 877
Brenan JPM. Flora of Tropical East Africa: Leguminosae, subfamily Mimosoideae. London: Crown
Agents for Overseas Governments & Administrations; 1959. 173 pp.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Njoroge GN. Ethnotherapeutic management of Sexually Transmitted Diseases (STDs) and repro-
ductive health conditions in Central Province, Kenya. Indian J Tradit Knowl. 2009;82:262–9.
Rapanea melanophloeos (L.) Mez.
PRIMULACEAE

Synonyms
Rapanea melanophloeos (L.) Mez.: Ardisia erythroxyloides Thouars ex Roem. &

Schult; Myrsine madagascariensis A. DC.; Myrsine melanophloeos (L.) R. Br.;
Myrsine simensis Hochst. ex A. DC.; Rapanea bovinii Mez; Rapanea comorensis
Mez; Rapanea erythroxyloides (Thouars ex Roem. & Schult.) Mex; Rapanea
seychellarum Mez; Rapanea simensis (Hochst. ex A. DC.) Mez; Rapanea thomensis
Exell
R. W. Bussmann (*)
G. N. Njoroge
J. Grimshaw

Local Names
Rapanea melanophloeos: Kikuyu: Mugaita; Ndorobo: Karambario, Kwarabariet;

Pokot: Sirtuot; Chagga: Kirjasi; Nandi: Kititet; Senga: Mpaja; Meru: Mugteta,
Muthitha; Taita: Mungala; Elgeyo/Nandi/Marakwet/Sebei: Sitotua, Situtua;
Kipsigis/Marakwet: Sitiotwet; Marakwet: Karabar; Kipsigis: Kipogong’yot
Botany and Ecology
Rapanea melanophloeos (L.) Mez.: Tree 3–20 m tall with trunk up to 0.6 m
diameter and twigs rather thick and soft, the very young parts sometimes brown-
tomentose but soon glabrous. Leaves petiolate, glabrous, oblong or oblong-
lanceolate, 6–12 cm long, 1.5–4 cm broad, tapering into petiole, apex subacute,
Fig. 1 Rapanea
melanophloeos (Primulaceae),
Naro Moru, Kenya. (Photo:
R. W. Bussmann)
Rapanea melanophloeos (L.) Mez. 881
Fig. 2 Rapanea
R. W. Bussmann)
rounded or sometimes emarginate, rather thick and veins mostly sunken, margins
slightly recurved when dry, minutely and sometimes obscurely gland-dotted on
both surfaces, petiole 5–15 mm long, slightly grooved along upper surface and
sometimes with few hairs along margin in young stage. Inflorescence in few-
several-flowered fascicles from the axils of the lower leaves or from above leaf
scars or on very abbreviated twigs, bracts minute, broadly ovate, pedicels stout,
3–5 mm long, angular, glabrous, functionally male, female or bisexual. Calyx
5-parted with segments ovate-lanceolate, minutely ciliate. Corolla 3–4 mm long,
united into a tube about 1 mm long, lobes oblong or elliptic-lanceolate, minutely
papillose along the inner margin, dark spotted. Stamens inserted within the corolla
tube, sessile or subsessile, anthers oblong, 1.5 mm long, minutely papillose on the
obtuse apex. Ovary with very short style and thick stigma. Drupe globose,
4–5 mm diameter (Taylor 1958). The species is a common component of dry
Afromontane forests (Bussmann 2002a, b, 2006a; Bytebier and Bussmann 2000)
(Figs. 1, 2, 3, 4, and 5).
Fig. 3 Rapanea
R. W. Bussmann)
Fig. 4 Rapanea
R. W. Bussmann)
Rapanea melanophloeos (L.) Mez. 883
Fig. 5 Rapanea
relict of cleared forest, about
3500 m, Bale Mountains
National Park, Bale
Mountains buffer zone above
Goba, Oromiya Province,
Ethiopia. (Photo:
J. Grimshaw)
Rapanea melanophloeos: Fruits eaten raw, or pounded with gruel, as anthelminthic.

Bark and roots also used in decoction for that purpose. Contains glycosides, thus to
be used with care (Beentje 1994; Kokwaro 2009). Sometimes the species is used as
anthelminthic (Njoroge and Bussmann 2006; Muthee et al. 2011) and also to treat
heartburn and gonorrhea (Muthee et al. 2011).
Rapanea melanophloeos: Seeds pounded and mixed with water as anthelminthic for
livestock (Kokwaro 2009). Hard timber, but not durable in the ground (Beentje
1994). Used as firewood (Bussmann 2006b). The hard wood is also used to make
charcoal, and the leaves are used for wrapping bread for baking (Luizza et al. 2013).
References
2006a;111:41–66.
Res Appl. 2013;11:315–39.
Taylor P. Flora of Tropical East Africa: Primulaceae. London: Crown Agents for Overseas Gov-
ernments & Administrations; 1958.
Rhamnus prinoides L’Hér.
Rhamnus staddo A. Rich.
RHAMNACEAE

Grace N. Njoroge
Synonyms
Rhamnus prinoides L’Hér.: Alaternus prinoides (L’Hér.) Raf.; Rhamnus pauciflora

Hoschst. ex A. Rich.; Ziziphus lucida Moench
Local Names
Rhamnus prinoides: Kikuyu: Mukarakinga; Chagga: Mfunganungu, Mshimbamba;

Kamba: Zambizi; Kipsigis: Konisityet; Kisii: Omungura; Maa: Ol-Konjii,
Ol-Kokola, Ol-Konyel; Marakwet: Konisit; Meru: Kwasisitiet; Samburu: Ikenyeli;
Sebei: Muapuarua; Tugen: Mizisitwa, Kukukua (Beentje 1994; Gachati 1989;
Kokwaro 2009).
Rhamnus staddo: Kikuyu: Mubura; Borana: Gale; Kamba: Muthunthi; Kipsigis:
Kusisitiet, Ketarwet; Kisii: Omonmanengo; Luo: Amurukia, Rahande; Maa:
R. W. Bussmann (*)
G. N. Njoroge

Ol-Kokola; Marakwet: Kipser; Meru: Mukuruu, Ngakula; Ndorobo: Korsosiat,

Arapsoi; Samburu: Ikokokai, Kokilai (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Rhamnus prinoides L’Hér.: Shrubs or trees up to 8 m tall, the branches sometimes

scrambling or sprawling. Year-old branches blackish, lenticellate, glabrate, youngest
twigs usually minutely pubescent with dark spreading hairs. Leaf-blades elliptic-ovate
to ovate-elliptic or narrowly so, (1–) 2–10 cm long, (8–) 14–40 mm wide, rounded to
cuneate at base, acuminate, firm, lustrous, nearly glabrous except for minute hairs on
the major nerves beneath, glandular-serrulate, on each side of midrib with 4–6 second-
ary nerves which are impressed above, prominulent beneath, petioles (3–) 5–10 mm
long. Stipules subulate, 3–5 mm long, minutely pubescent, caducous. Flowers rarely
solitary, usually in fascicles of 2–3 (–8), 5-merous, pedicels filiform, minutely pubes-
cent, 3–10 mm long in flower, 7–15 mm in fruit. Cup minutely pubescent externally,
2 mm wide. Sepals about 2 mm long. Petals spatulate, about 1 mm long, or usually
absent in East African populations. Fruit nearly globose, 5–6 mm thick, reddish. Forest,
evergreen bushland and thicket, 700–3700 m (Johnston 1972). Common in regenera-
tion stages of Juniperus procera dominated Afromontane forests (Bussmann 2002).
Sometimes semidomesticated (Balemie and Kebebew 2006) (Figs. 1, 2, 3, 4, and 5).
Fig. 1 Rhamnus prinoides

(Rhamnaceae), Mount Kenya
Bussmann)
Rhamnus prinoides L’Hér. . . . 887

Bussmann)

Bussmann)

Bussmann)

Bussmann)
Rhamnus staddo A. Rich.: Deciduous shrubs or small trees, 0.5–7 m tall, sometimes
scandent. Bark smooth, gray, slash cream, year-old branches dark brown-gray,
slender, glabrate, youngest twigs densely to less commonly sparsely hispidulous
with gray hairs 0.1–0.2 mm long, short shoots (many-noded twigs) often produced
late in the growing season, elongated twigs occasionally thorn-tipped. Flowers borne
singly at the nodes, often appearing at the nodes of the short-shoots of the previous
season and thus in somewhat fascicle-like aggregations, greenish or sometimes said
to be cream color or orange, fragrant, pedicels usually glabrous, 1–2 (–4) mm long in
flower, in fruit about 2 (–4.5) mm long. Seed about 4–5 mm long, pale brownish,
smooth, with a ventral groove. Sepals 4, deltoid, 1–1.7 mm long, glabrous or rarely
hispidulous on back. Petals 4, linear to oblanceolate, about 0.5–0.8 mm long, not
clasping, caducous. Stamens 4, said to be brownish when fresh. Fruit nearly globose,
about 5–5.5 mm long, reddish or scarlet when ripe, with 3 stones or by abortion
fewer, each stone on drying eventually capable of colubrinoid dehiscence, revealing
an orange or brick red inner lining. Leaf-blades obovate or narrowly obovate or
broadly oblanceolate, rarely elliptic, (1–) 2–3 (–4.4) cm long, (5–) 6–15 (–20) mm
wide, cuneate, blunt to acute and mucronate, dark olive green or brown-green
(at least when dry), glabrous or hispidulous on the major nerves beneath and
occasionally above or rarely hispidulous on both surfaces with gray hairs 0.1–
0.2 mm long, obscurely glandular-serrulate or occasionally with the teeth reduced
to vanishingly small size, with 5–8 pairs of obscure secondary nerves, petioles (1–)
Fig. 6 Rhamnus staddo

Bussmann)
Fig. 7 Rhamnus staddo

Bussmann)
2–5 (–8) mm long, usually at first hispidulous and later glabrescent. Stipules filiform
to subulate, 1–3 mm long, caducous to subpersistent. Cup glabrous or rarely
hispidulous (Johnston 1972). Common in regeneration stages of Juniperus procera
dominated Afromontane forests (Bussmann 2002) (Figs. 6 and 7).
Rhamnus prinoides: Root is used for indigestion (Meru), gonorrhea and rheuma-
tism (Maasai), and malaria (Kipsigis) (Beentje 1994; Kokwaro 2009). Also used as
diuretic and to increase fertility in women (Kokwaro 2009). Used as antimalarial
(Njoroge and Bussmann 2006a), and to treat cough, chest pain, and colds (Njoroge
and Bussmann 2006b). Used to improve backache, general body health, malaria, and
stomachache, as well as arthritis and rheumatism (Muthee et al. 2011; Nankaya et al.
2019, 2020). Sometimes used to treat prostate problems and brucellosis (Muthee
et al. 2011). In Ethiopia, it is also used to treat hepatitis (Yineger et al. 2007) and
eczema (Teklehaymanot and Giday 2007). The species has shown antibacterial
properties (Desta 1995) and is used to treat snakebites (Bussmann 2006).
Rhamnus staddo: Root is used for malaria and anaplasmosis (Meru), as well as
venereal diseases (Maasai) (Beentje 1994; Kokwaro 2009). To treat gonorrhea,
diabetes and endometritis (Muthee et al. 2011), and common colds (Yineger et al.
2008). Traditionally used to treat polio and tuberculosis (Bussmann 2006).
Rhamnus prinoides: Used in ceremonies (Samburu) (Kokwaro 2009) and the

fermentation of honey wine (tej) (Bussmann et al. 2011; Luizza et al. 2013;
Teklehaymanot 2009). The wood serves to make tools (Mekonnen et al. 2015) and
for construction (Bussmann et al. 2011).
References
Johnston MC. Flora of Tropical East Africa: Rhamnaceae. London: Crown Agents for Overseas
Res Appl. 2013;11:315–39.
1746-4269-3-12.
2008;26:132–53.
Rhus natalensis Bernh. ex C. Krauss
Rhus ruspolii Engl.
Rhus tenuinervis Engl.
Rhus vulgaris Meikle
ANACARDIACEAE

Grace N. Njoroge
Synonyms
Rhus natalensis Bernh. ex C. Krauss: Rhus macrocarpa Engl.; Searsia natalensis

Bernh. ex C.Krauss) F.A. Barkley; Toxicodendron macrocarpum (Engl.) Kuntze;
Toxicodendron natalense (Bernh. ex C.Krauss) Kuntze
Rhus ruspolii Engl.: Rhus bequaertii Robyns & Lawalrée; Searsia ruspolii (Engl.)
Moffett.
Rhus tenuinervis Engl.: Rhus buluwayensis Diels; Rhus commiphoroides Engl. &
Gilg; Rhus kwebensis N.E. Br.; Searsia tenuinervis (Engl.) Moffett; Toxicodendron
tenuinerve (Engl.) Kuntze
Rhus vulgaris Meikle: Rhus pyroides Burch. var. pyroides; Searsia pyroides
(Burch.) Moffett; Toxicodendron pyroides (Burch.) Kuntze
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Rhus natalensis: Kikuyu: Muthigiu, Muthigio; Luhya: Busangula; Meru:

Mirimathua; Digo: Mngongo, Mugwa-Nyahi; Hehe: Musagara; Zinza: Msense;
Toro: Msheshe; Luo: Sangla; Swahili: Mlishangwe; Borana: Dabobiss; Boni:
Idamudu; Gabbra: Dabobbessa; Kamba: Mutheu, Kitheu; Kipsigis: Suriet,
Sirondet; Maa: Olmisigiyoi; Marakwet: Seria; Pokot: Siryewo; Samburu:
Limisigyioi; Tugen: Sirwo, Siryande; Turkana: Ekadetewa; Nandi: Monjororioyat,
Siriat; Somali: Ilka-Adeis; Taita: Kitariki (Beentje 1994; Gachati 1989; Kokwaro
2009).
Rhus ruspolii: Kamba: Kitheu (Beentje 1994).
Rhus tenuinervis: Kamba: Kitheu; Maa: Olruisigiyoi (Kokwaro 2009).
Rhus vulgaris: Luo: Awayo, Sangala-Modongo; Haya: Msagara; Kikuyu:
Muthigio; Shambaa: Mtuntano; Chagga: Nyungu; Maa: Olmisigiyioi,
Ol-Munyushi; Kamba: Kitheu, Mutheu; Kipsigis: Monchororiat, Siriat; Meru:
Murimuthu; Sebei: Njowaruwa (Beentje 1994; Kokwaro 2009).
Botany and Ecology
Rhus natalensis Bernh. ex C. Krauss: Shrub 2–3 m high or a small tree up to 8 m;

bark of the branchlets grayish or whitish and those of the old branches dull gray,
lenticellate and rough; branchlets, especially the flowering ones sometimes densely
tomentose. Petiole 1.5–3.5 ( 4) cm long, convex below, longitudinally grooved
above, glabrous or pubescent; median leaflet obovate, oblong or elliptic, 2.5–9 cm
long, 1–3.5 cm wide, obtuse or rounded and sometimes emarginate at the apex
(rarely acute), somewhat cuneate at the base; lateral leaflets ovate, obovate or
oblong-obovate, (1.8–) 2.5–7 cm long, 1–3.5 cm wide, rounded and sometimes
emarginate at the apex, somewhat cuneate at the base, sessile; all leaflets slightly
or conspicuously discolorous (greenish gray on both surfaces or dull green or
brownish above and greenish gray beneath), entire or undulate-crenate along the
margin, papyraceous or subcoriaceous; midrib slightly raised on the upper surface
and prominent beneath; lateral nerves slightly raised on both surfaces but more so on
the upper surface; reticulation scarcely or not at all visible. Panicles up to 12 cm
long, lax, generally shorter than the leaves, the axis and branches somewhat pilose;
pedicels 1–2 mm long. Calyx-segments ovate, 0.3–0.5 mm long. Petals oblong,
1–1.5 mm long, whitish or greenish. Staminodes present in female flowers. Disk
shallowly 5-lobulate. Ovary subglobose; styles 0.5–0.7 mm long, reflexed; stigmas
subcapitate. Drupe oblong-reniform, 5–6 mm in diameter, glabrous. Deciduous and
evergreen bushland and woodland, riverine associations, forest edges, often on well-
drained slopes; 1–3000 m (Kokwaro 1986). Very common in the undergrowth of
drier Afromontane forests (Bussmann 2002a, b) (Figs. 1 and 2).
Rhus ruspolii Engl.: Shrub or small tree 2–7 m high; branchlets somewhat cylindric,
ferrugineus tomentose to glabrate and lenticillate. Petiole 1.8–4 cm long, sub-
cylindric, flattened or longitudinally grooved above, ferrugineus tomentellous to
Rhus natalensis Bernh. ex C. Krauss. . . 895
Fig. 1 Rhus cf. natalensis

(Anacardiaceae), Mt. Kenya
Bussmann)
glabrate; leaflets oblong-obovate or elliptic, 5.5–18 cm long, 3–11 cm broad, the

median usually slightly larger than the lateral ones, irregularly rounded to obtuse or
acute at the apex, rather asymmetric and obtuse or cuneate (particularly the laterals)
to narrowly cuneate (especially the median), halfway broadly crenate from the apex
and entire towards the base, papyraceous or subcoriaceous, ferrugineus pubescent,
discolorous especially the upper surface which also turns glabrate; midrib and the
lateral nerves prominently raised beneath and conspicuous above; reticulation ini-
tially concealed by the indumentum but clear in the older leaves. Panicles 10–28 cm
long, terminal and in the upper leaf-axils, pyramidal, ferrugineus tomentellous or
pubescent, the terminal ones usually with bract-like leaves at the base; flowers rather
glomerate, brownish white; male with pedicel 0.5–2 mm long, pubescent. Calyx-
lobes oblong-ovate, 0.5–0.7 mm long, pubescent. Petals elliptic or oblong-ovate, 1–
1.5 mm long, 0.6–0.9 mm broad, puberulent on the exterior. Disk saucer-shaped and
crenate. Ovary globular, 0.5–1 mm in diameter; styles 3, 0.3–0.4 mm long. Drupes
globose 3–4 mm in diameter, glabrous and shiny, pinkish-red. Upland evergreen
bushland, forest edges, riverine associations, 1200–2450 m (Kokwaro 1986). Very
common in the undergrowth of drier Afromontane forests (Bussmann 2002a, b).
Fig. 2 Rhus cf. natalensis

(Anacardiaceae), Mt. Kenya
Bussmann)
Rhus tenuinervis Engl.: A much-branched sometimes thorny shrub or small tree up

to 8 m high; branches with dull grayish rough lenticellate bark, glabrous or
glabrescent; branchlets somewhat densely spreading pilose with slender hairs, yel-
lowish. Petiole 1–4 cm long, convex below, somewhat longitudinally grooved
above, pubescent or pilose; median leaflet ovate, obovate, subcircular, ovate-oblong
or elliptic, 2.5–11.5 cm long, 1.5–5 (–7) cm broad, obtuse to rounded or truncate at
the apex, subacute or acute to cuneate at the base, often much contracted and
subpetiolate; lateral leaflets oblong-ovate to subcircular, 1.5–6 cm long, 1–4 cm
broad, cuneate at the base; all leaflets discolorous (glaucous or dull green above,
paler and generally yellowish below), papyraceous to subcoriaceous, somewhat
spreading pubescent on both surfaces, relatively glabrescent or almost glabrous,
the upper 2/3 of the margin coarsely crenate or bicrenate, the lower portion entire,
rarely the leaflets entire; midrib and lateral nerves prominent beneath; venation
scarcely visible. Panicles up to 15 (–25) cm long, axillary and terminal, lax, the
axis and branches pilose; pedicels 1–2 mm long, articulated below the calyx. Calyx-
segments ovate, 0.4–0.6 mm long, obtuse. Petals 1.2–1.5 mm long, obtuse. Disk
5-lobulate and patelliform. Ovary globose; styles 0.4 mm long. Drupe com-
pressed, subquadrangular or subcircular, 5–7 mm in diameter, pruinose, brownish.
Wooded grassland and bushland, especially on rocky slopes, hardly in miombo;
900–1950 m (Kokwaro 1986). Very common in the undergrowth of drier
Afromontane forests (Bussmann 2002a, b).
Rhus vulgaris Meikle: Shrub or small tree up to 7 m high, often thorny; branchlets
brownish and spreading hairy; old branches dark grayish. Petiole 1–2.5 cm long,
slender, terete, pubescent, somewhat longitudinally grooved above; median leaflet
oblanceolate to elliptic, 3–8 cm long, 1–2.3 cm broad, acute at the apex, cuneate at
the base; lateral leaflets normally 2/3 or 3/4 the size of the median one, elliptic; all
glaucous (pale greenish and slightly darker above when dry), entire or some with
dentate margin, submembranous, appressed pilose or sometimes densely sericeous,
occasionally nearly glabrous; midrib and lateral nerves slender, raised on both
surfaces and more prominent beneath; reticulation very close and visible on both
surfaces. Panicles 2–5 (–7) cm long, terminal and axillary, pubescent; bracts 1 mm
long; flowers. Calyx segments 0.5–8 mm long. Petals oblong, 1.2–1.5 mm long.
Fig. 3 Rhus cf. vulgaris

(Anacardiaceae), Bale
Drupe subglobose, slightly compressed when dried, 3–4 mm in diameter (Kokwaro

1986). Very common in the undergrowth of drier Afromontane forests (Bussmann
2002a, b) (Figs. 3, 4, 5, and 6).



Rhus natalensis: Roots are pounded in water and the extract is drunk for flu,
abdominal pain and gonorrhea, hookworm, and diarrhea. Leaves are used to treat
cough, and the steam from boiled leaves is inhaled for colds. The leaf decoction is
drunk for stomachache, especially in children (Kokwaro 2009). Used to remedy
stomachache (Bussmann 2006). The fruits are eaten to prevent chest problems, and
the twigs are used as toothbrush (Bussmann et al. 2006). The plant serves also as
antimalarial (Njoroge and Bussmann 2006), and to treat back pains, diarrhea,
anthrax, postpartum pains, and toothache (Njoroge et al. 2004).
Rhus tenuinervis: Young shoots and leaves are chewed for heartburn (Kokwaro
2009).
Rhus vulgaris: Boiled fruits are used for diarrhea. Boiled stems are applied to
wounds. Powdered roots are used for gonorrhea and the leaves for hemorrhoids
(Kokwaro 2009). The fruit decoction is used for diarrhea (Beentje 1994). The plant
serves also as antimalarial (Njoroge et al. 2004; Njoroge and Bussmann 2006).
Rhus ruspolii: Used to remedy stomachache (Bussmann 2006) and to treat wounds
Local Food Uses
Rhus natalensis: The fruit is eaten (Beentje 1994; Bussmann 2006).

Rhus tenuinervis: The fruit is eaten (Beentje 1994).
Rhus vulgaris: The fruit is eaten (Beentje 1994).
Rhus ruspolii: The fruit is edible (Bussmann 2006).
Rhus coriaria is used as spice (Yeşil et al. 2019).
Rhus natalensis: A bark infusion is given to livestock with East Coast Fever
(Kokwaro 2009). The leaves are browsed by livestock (Bussmann 2006). Burnt
for its good smell and used as firewood (Bussmann et al. 2006). Given top livestock
to treat endometritis and foot and mouth disease (Muthee et al. 2011). Produces
excellent charcoal (Tian 2017).
Rhus vulgaris: A root decoction is used for diarrhea in livestock (Kokwaro 2009).
Rhus ruspolii: The leaves are browsed by livestock (Bussmann 2006). Rhus
glutinosa is used as fuelwood (Mekonnen et al. 2015).
References
Kokwaro JO. Flora of Tropical East Africa: Anacardiaceae. Rotterdam: Balkema; 1986.
Ethnomed. 2006;2:8.
agro-ecosystems. Lyonia. 2004;72:71–87.
10.1186/s13002-016-0131-x.
Yeşil Y, Çelik M, Yılmaz B. Wild edible plants in Yeşilli (Mardin-Turkey), a multicultural area.
J Ethnobiol Ethnomed. 2019;15:52. https://doi.org/10.1186/s13002-019-0327-y.
Ricinus communis L.
EUPHORBIACEAE

Grace N. Njoroge
Synonyms
Ricinus communis L.: Ricinus africanus Mill.; Ricinus angulatus Thunb.; Ricinus
armatus Andrews; Ricinus atropurpureaus Pax & K. Hoffm.; Ricinus badius Rchb.;
Ricinus cambodgensis Benary; Ricinus digitatus Norhona; Ricinus europaeus
T. Nees; Ricinus giganteus Pax & K. Hoffm.; Ricinus glaucus Hoffmanns.; Ricinus
hybridus Besser; Ricinus inermis Mill.; Ricinus japonicus Thunb.; Ricinus krappa
Steud.; Ricinus laevis DC.; Ricinus leucocrpus Bertol.; Ricinus lividus Jacq.; Ricinus
macrocarpus Popova; Ricinus macrophllus Bertol.; Ricinus medicus Forssk.;
Ricinus medius J. F. Gmel.; Ricinus megalospermus Delile; Ricinus messeniacus
Heldr.; Ricinus metallicus Pax & K. Hoffm.; Ricinus microcarpus Popova; Ricinus
minor Mill.; Ricinus nanus Bald.; Ricinus obermannii Groenl.; Ricinus peltatus
Norhona; Ricinus perennis Steud.; Ricinus persicus Popova; Ricinus purpurascens
Bertol.; Ricinus ruber Miq.; Ricinus rugosus Mill.; Ricinus rutilans Müll. Arg.;
Ricinus sanguineus Groenl.; Ricinus scaber Bertol. ex Moris; Ricinus speciosus
Burm. f.; Ricinus spectabilis Blume; Ricinus tunisensis Desf.; Ricinus undulatus
R. W. Bussmann (*)
G. N. Njoroge

Besser; Ricinus urens Mill.; Ricinus viridis Willd.; Ricinus vulgaris Mill; Ricinus
zanzibarinus Popova
Local Names
Ricinus communis: Kikuyu: Mwariki; Swahili: Mbarika, Mbono mdogo, Mbono,

Mnyonyo; Borana: Kobor; Ilwana: Mobonu; Kamba: Kivunu; Kipsigis: Menuet;
Luhya: Mubonenone, Ibono; Maa: Oldule; Marakwet: Manwa; Meru: Mwariki;
Oromo: Koboo; Pokot: Pondon; Rendile: Fololo; Somali: Gitkalat; Acholi: Coga
Macon; Taita: Mbonu; Luo: Odagwa; Luganda: Nsogasoga (Beentje 1994;
Gachati 1989; Kokwaro 2009), English: Castor bean
Botany and Ecology
Ricinus communis L.: An erect, single-stemmed but often much-branched bushy or

tree-like glabrous, more or less glaucous annual or perennial herb commonly up to
5 m, but occasionally attaining 10 m. Stems up to 12 cm in diameter at the base,
hollow, herbaceous but often becoming woody, pale bluish-green or reddish tinged.
Young shoots often pruinose and purplish tinged. Petioles (4–) 10–30 cm long,
usually smooth, rarely sparingly beset with small cylindrical papillae, petiolar glands
discoid to turbinate, dark green, blades up to 1 m across, (5–) 7–9 (–12) lobed,
usually more than halfway but occasionally less, the median lobe usually 7–30 cm
long and 2–8 cm wide, but occasionally up to 75 cm long and 20 cm wide, the lateral
lobes being proportionately progressively smaller, lobes ovate-lanceolate, lanceo-
late, oblong or linear-oblong, acutely acuminate, coarsely and sometimes somewhat
irregularly glandular-serrate, firmly membranaceous to thinly chartaceous, lateral
nerves (15–) 20–25 on the median lobe, progressively fewer on the lateral lobes, not
prominent above, slightly so beneath, craspedodromous, dark olive-green above,
paler beneath, midribs and nerves often reddish beneath. Stipular sheath ovate,
1.3–2.7 cm long, often reddish or purplish, soon caducous, leaving a about circular
scar. Inflorescence 10–30 cm long, bracts lanceolate, about 1 cm long, green, reddish
tinged, bracteoles smaller than the bracts, but otherwise similar. Male flowers:
pedicels 0.5–1.7 cm long, calyx-lobes ovate, 5–8 mm long, (2–) 3–5 mm wide,
acute, pale green, often tinged reddish or purplish, stamens 7–8 mm long, anthers
0.5 mm long, pale yellow. Female flowers: pedicels 0.5–1 cm long, extending to
2–4.5 cm in fruit, sepals narrowly lanceolate, about 5 mm long, acutely acuminate,
often reddish or purplish tinged, ovary trilobate-subglobose, 2 mm long and wide,
styles 2–5 mm long, about one-third bifid to almost completely bipartite, usually red,
rarely yellowish. Female flowers: pedicels 0.5–1 cm long, extending to 2–4.5 cm in
fruit, sepals narrowly lanceolate, about 5 mm long, acutely acuminate, often reddish
or purplish tinged, ovary trilobate-subglobose, 2 mm long and wide, styles 2–5 mm
long, about one-third bifid to almost completely bipartite, usually red, rarely yel-
lowish. Fruit strongly trilobate, 1–1.8 cm long, 1–1.5 cm diameter, sparingly, evenly
Ricinus communis L. 905
or densely beset with narrowly cylindric fleshy processes 3–5 mm long, each tipped
with a sharp curved bristle, bluish green, often tinged reddish or purplish. Seeds
7–12 mm long, 5–8 mm wide, 4–6 mm deep, smooth, shiny, grayish, silvery or
beige, usually variously streaked, mottled and flecked with olive-brown, chestnut or
dark grayish brown, caruncle depressed-conic, 1–2 mm long, 2–3 mm across.
Widely grown and readily naturalized in areas of moderate to good rainfall in
frost-free areas, poisonous to stock, sea level 1900 (–2590) m. Ricinus communis
is indigenous to north-eastern tropical Africa. It was already grown for its oil in
Egypt some 6000 years ago and spread through the Mediterranean, the Middle East,
and India at an early date. It is now widely cultivated in most drier areas of the
tropics and subtropics and in many temperate areas with a hot summer. It naturalizes
easily and grows in many areas as a ruderal plant (Figs. 1, 2, and 3).
Fig. 1 Ricinus communis

(Euphorbiaceae), Loja,
Ecuador. (Photo: R. W.
Bussmann and N. Y.
Paniagua-Zambrana)

Bussmann and N. Y.
Paniagua-Zambrana)

Bussmann and N. Y.
Paniagua-Zambrana)
Ricinus communis: In medicine, castor oil is used primarily as a purgative. It is

commonly referred to in South Africa as “blue bottle” because of the characteristic
blue bottle in which it was traditionally packed and sold. It was much feared by
children because of the unpleasant taste. The oil is now sometimes given as a
sweetened aromatized emulsion or as capsules. It stimulates peristalsis by irritating
the intestinal mucosa but causes little griping. It is also applied as an emollient in the
treatment of sores and as a solvent for antibiotic eyedrops. Neutral sulfated castor
bean oil can replace soap in certain cases of contact dermatitis. Castor bean oil has
been used as an abortifacient and is given orally, alone or with quinine sulfate, to
induce labor in pregnancy at term. In Kenya, the Kikuyu use the root decoction to
expel the placenta (Beentje 1994; Kokwaro 2009). Ricinoleic acid prepared from the
oil is a component of contraceptive creams and jellies. The roots are also chewed to
remedy infections with guneaworm (Dracunculus medinensis), and the decoction is
used to stimulate appetite and for stomach problems. Juice extracted from stem and
leaves serves for treatment of ulcers, stomachache, and diarrhea. Fresh leaves are
sometimes applied to wounds. Pounded seeds, although toxic, are used to stop after-
birth bleeding, as purgative, and the seed oil for earache. The flowers mixed with
Commiphora africana leaves are used to smoothen the skin (Meru) (Kokwaro 2009).
In the Bale region of Ethiopia, the seed oil is used for wounds (Desta 1995; Luizza
et al. 2013). The species is also used to treat cough (Njoroge and Bussmann 2006a),
skin rashes (Njoroge and Bussmann 2007), and placenta problems (Njoroge 2009). In
Madagascar, it is employed for asthenia, hemorrhoids, wounds, intestinal parasites,
cold, as galactagogue, purgative, laxative, anthelminthic, for, headache, rheumatism,
dental cavities, wounds, and fevers (Randriamiharisoa et al. 2015). Used in Ethiopia to
treat rabies (Yineger et al. 2008). Also used to treat gonorrhea (Lulekal et al. 2008;
Wondimu et al. 2007), for flu, headache, stomach pain and chest pain (Teklehaymanot
2009; Teklehaymanot and Giday 2010), and toothache (Teklehaymanot and Giday
2007). In Madagascar, it is used to treat breast infections in lactating women and cough
in infants (Randrianarivony et al. 2016). For placenta problems, skin rashes and
urinary problems (Njoroge et al. 2004). As purgative for stomachache and diarrhea
(Giday et al. 2009; Nankaya et al. 2019). The species has some antifertility effects
(Gebrie et al. 2005). Seeds used to treat gallstones, alcoholism and as laxative
(Bussmann et al. 2016). For scars and swellings (Quiroga et al. 2012). As poultice
to treat sprains and fractures (Béjar et al. 2001; Bussmann and Sharon 2006a, 2007a).
The fresh leaves are used to treat constipation (Bussmann and Sharon 2006b, 2007b,
2015a, b). The seed oil is also used for hematomas and wounds, especially in
veterinary medicine (Monigatti et al. 2013). In India, Ricinus is used to treat swellings
and rheumatism (Debbarma et al. 2017), as well as cough, headache, jaundice, dental
problems, joint pain, wounds, stomach problems, indigestion, skin disorders, boils and
burns, and serves as anthelminthic (Flatie et al. 2009; Raj et al. 2018). In Nepal, the
seeds serve for constipation and the roots as analgesic (Kunwar et al. 2009, 2010,
2015). Applied externally for lumpy skin disease in livestock (Kioko et al. 2015).
Ricinus communis: Although they are somewhat toxic, mature leaves are occasion-
ally used as a fodder, but care must be taken to avoid the more toxic young leaves.
Castor bean is commonly grown as an ornamental. The press cake of castor bean
seeds is poisonous and allergenic and is mainly used as fertilizer or as fuel. Methods
to detoxify the press cake and make it suitable as an animal feed have been
developed, but even after treatment some toxicity may remain, horses are particu-
larly sensitive to it. Another product extracted from the press cake is a lipase used in
the industrial processing of fats. About 95% of castor seed is used for the expression
of oil, which consists mainly of triglycerides of ricinoleic acid, is non-drying and
non-edible. Traditionally, it is used for illumination. As a lamp oil, it is believed to
give a cooler and brighter light than other vegetable and mineral oils, burn more
steadily and produce very little soot. It is now only used in rural areas and even there
often mixed with or as a substitute for kerosene. Currently, castor bean oil is
primarily used as a high-quality lubricant and a versatile raw material in the chemical
industry. It has long been used as a lubricant in carts and Persian wheels. It is
characterized by a high lubricity, high viscosity remaining constant over a wide
range of temperatures, and insolubility in aliphatic petrochemical fuels and solvents,

making it suitable for equipment operating under extreme conditions such as in
arctic zones and in aviation. Another specialized use of castor bean oil is in crumb-
rubber manufacturing, where it prevents rubber crumbs from coagulating. Highly
purified, food-grade castor oil is used as an anti-stick agent for candy molds and as a
lubricant for machinery in industrial food processing. Castor bean oil is further
employed as a plasticizer in the coating industry, as a disperser for dyes, and as
filler in cosmetics such as lipsticks, nail varnishes, and shampoos. Saponification of
castor oil yields a clear, transparent soap. Washing jute fiber with the soap gives it a
shiny silky appearance. The soap has poor detergent qualities but is easily water
soluble. Kenyan herders use the leaves for theileriosis in cattle (Njoroge and
Bussmann 2006b). The seed oil is used for greasing baking pans, to soften leather,
and the leaves are browsed by cattle (Bekalo et al. 2009; Bussmann et al. 2011;
Luizza et al. 2013). Used as acaricide (Wanzala 2017). Used to make unspecified
household tools (Mekonen et al. 2015).
References
Medicinales del Sur Ecuatoriano. San Diego: Latino Herbal Press; 2001. p. 340.
doi.org/10.1186/1746-4269-5-26.
longevidad – La flora medicinal de Vilcabamba. Honolulu: Arogya; 2007a. ISBN: 978-0-
9789962-2-2.
de los cuatro vientos – La flora mágica y medicinal del Perú. Honolulu: Arogya; 2007b. ISBN:
978-0-9789962-3-9.
medicinal flora of Northern Peru. St. Louis: William L. Brown Center, MBG; 2015a. ISBN:
978-0-9960231-2-2.
medicinal del Norte de Peru. St. Louis: William L. Brown Center, MBG; 2015b. ISBN: 978-0-
9960231-3-9.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Gebrie E, Makonnen E, Debella A, Zerihun L. Phytochemical screening and pharmacological
evaluations for the antifertility effect of the methanolic root extract of Rumex steudelii.
macology of medicinal plants in far-west Nepal. Ethnobot Res Appl. 2009;7:5–28.
indigenous uses and pharmacological validity. Med Aromatic Plant Sci Biotechn. 2010;1(4
special issue):28–42.
medicines in farwest Nepal. J Ethnopharmacol. 2015;163:210–9.
Res Appl. 2013;11:315–39.
10.1186/1746-4269-4-10.
(2):450–64.
Njoroge GN. Ethnotherapeutic management of Sexually Transmitted Diseases (STDs) and repro-
ductive health conditions in Central Province, Kenya. Indian J Tradit Knowl. 2009;82:262–9.
ment of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol.
2006b;108:332–9.
Quiroga R, Meneses L, Bussmann RW. Medicinal ethnobotany in Huacareta (Chiquisaca, Bolivia).
J Ethnobiol Ethnomed. 2012;8:29.
018-0208-9.
org/10.1186/s13002-015-0046.

1746-4269-3-12.
semipastoralist people in lower Omo River Valley, Debub Omo Zone, Southern Nations,
https://doi.org/10.1155/2017/8647919. Article ID: 8647919, 33 pages.
10.1186/1746-4269-4-11.
Rubus apetalus Poir.
Rubus rosifolius Sm.
Rubus volkensii Engl.
ROSACEAE

Grace N. Njoroge
Synonyms
Rubus apetalus Poir.: Rubus adolfi-friederici Engl.; Rubus adolfi-friederici var.

rubistylis Gust.; Rubus assaortinus Chiov.; Rubus borbonicus Pers.; Rubus
chiovendae Gust.; Rubus ecklonii Focke; Rubus exsuccus Steud. ex A. Rich.;
Rubus interjungens Gust.; Rubus madagascaricus Gust.; Rubus petalabigens
Gust.; Rubus petitianus A. Rich.; Rubus pinnatiformis Gust.; Rubus quartinianus
A. Rich.
Rubus rosifolius Sm.: Rubus comintanus Blanco; Rubus commersonii Poir.; Rubus
coronarius (Sims) Sweet; Rubus jamaicensis Blanco; Rubus pinnatus Willd; Rubus
rosifolius var. coronarius Sims; Rubus rosifolius var. wuyishanensis Z. X. Yu; Rubus
sinensis Sims
Rubus volkensii Engl.: Rubus chiesae Chiov.; Rubus mauensis Engl.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Rubus apetalus: Kikuyu: Mutare; Pare: Mtelia (Beentje 1994; Gachati 1989;
Kokwaro 2009).
Rubus rosifolius: Kikuyu: Mutare; Kipsigis: Tagaimamiet; Maa: Engainagut;
Tugen: Mowonwo (Beentje 1994; Gachati 1989; Kokwaro 2009).
Rubus volkensii: Kikuyu: Mutare-Kigombe, Mutare; Kipsigis: Tagaimamiet; Maa:
Engaiyaguji (Beentje 1994, Gachati 1989, Kokwaro 2009).
Botany and Ecology
Rubus apetalus Poir.: A scrambling shrub. Flowering branches densely and softly
white- or yellowish-pilose, sometimes villous, the hairs largely spreading,
prickles slender, hooked or straightly deflexed, up to 4–5 mm long. Leaves
imparipinnate (2– (3–) jugate) or trifoliolate, or the uppermost simple, leaflets rather
variable in shape, commonly lanceolate, oblong-lanceolate, or oblong-elliptic,
6.5–9 cm 4–6 cm, rather obtuse or the upper ones usually very acute, basally
rounded to subcordate, sharply but shallowly serrate or biserrate with teeth 1 (–2)
mm deep, dark green and variably hairy above (often rather evenly but thinly covered
with appressed hairs), paler beneath, and softly pilose or more usually gray- or
greenish- or whitish-tomentose, sometimes thickly covered with soft, whitish or
fulvous hairs, terminal petiolules 1–2 (–3) cm long, the lateral ones up to 7 mm
long. Inflorescence paniculate, narrowly to broadly pyramidal, up to 21 9 cm, the
basal branchlets spreading at a wide angle or horizontally, flowers rather congested,
axis, branchlets and pedicels whitish-tomentose interspersed with variably numerous,
fulvous or pale fulvous, stiff hairs. Petals absent or very small. Calyx 5–8 mm long,
deeply divided into lanceolate, clasping lobes 4–7 mm long with a protracted apex,
hairy as the pedicels. Carpels varying from glabrous to densely hairy (and sometimes
with a dense apical tuft). Fruit black. Edges of upland rainforest, upland bushlands,
thickets, roadsides, 1275–2100 m (Graham 1960). Common in Afromontane forests
(Bussmann 2002, 2006a, b).
Rubus rosifolius Sm.: A handsome scrambling shrub up to 2.5 m tall, with erect or
arching stems covered although scarcely densely so with long spreading white hairs
interspersed with scattered or locally numerous amber-green glands. Prickles
scattered, small, up to 3 mm long, straight or abruptly decurved. Leaves
imparipinnate (or the uppermost simple), 3–5 jugate (5–7 (–11) jugate on the
turions), leaflets up to 7.5–9 cm 3.5 cm, ovate to oblong-lanceolate, long
drawn-out to an acute apex and with a shortly cuneate, rounded or truncate
base, jaggedly doubly serrated, the primary serratures saw-like, up to 5 mm deep,
covered above and below with rather long whitish hairs but usually more sparsely so
Rubus apetalus Poir. . . . 913
and paler green beneath. Inflorescence consisting of axillary and terminal flowers,
borne singly on pedicels 2–4 cm long. Flowers (2–) 2.5–3 (–4) cm in diameter when
fully open. Calyx deeply divided, lobes lanceolate or ovate-lanceolate with a caudate
apex, 8–20 mm long, connate for 3–4 mm at the base, exceeding the petals. Petals
white turning to pink, broadly ovate to subcircular. Carpels not more than 1 mm
long, produced “in hundreds.” Receptacle elongating, becoming ellipsoid-
cylindrical, up to 2 cm long when fully mature. Ripe fruit scarlet, glistening, edible.
Edges of upland and lowland rainforests and plantations, secondary bushland,
abandoned cultivations, etc., 900–1450 m. A native of eastern Asia, introduced
into Africa, and quite commonly naturalized (Graham 1960). Common in
Afromontane forests (Bussmann 2002, 2006a, b).
Rubus volkensii Engl.: A handsome, viscid shrub, up to 4 m tall. Stems brownish-
red, densely covered with stipitate and sessile glands intermingled with loose white
hairs and scattered prickles. Leaves imparipinnate, or trifoliolate within the inflo-
rescence with the terminal leaflets petiolulate, the lateral ones sessile, leaflets
ovate, ovate-elliptic or ovate-lanceolate, up to 10.5 6.5 cm, often long-
acuminate, the base rounded to cordate, sharply serrate or biserrate, with very
acute teeth, green on both surfaces, pilose and glandular on both surfaces but more
so beneath, the young leaves softly villous, rachis hairy, glandular and prickly as
the stems. Inflorescence loosely paniculate, terminal and axillary, scarcely or not
exceeding the leaves. Peduncles densely stipitate-glandular. Pedicels densely
stipitate-glandular. Flowers large, 2.5–3 cm diameter, apparently somewhat
drooping. Calyx densely stipitate-glandular on the outer side, the calyx sometimes
with basal pricklets, divided almost to the base into ovate-lanceolate, caudate-
acuminate lobes 2–2.5 cm long. Petals yellow or palely so, sometimes white,
broadly ovate, 1.5–2 cm long, obtuse. Carpels pubescent. Ripe fruit viscid, edible
but rather acid, apparently yellow or orange becoming bright red or brownish-red
at maturity. Edges and clearings in upland rainforest and moist bamboo-thicket,
2100–3450 m (Graham 1960). Common in Afromontane forests (Bussmann 2002,
2006a, b) (Figs. 1, 2, 3, 4, and 5).
Rubus apetalus: Fruits boiled with sugar are used for anemia, and the leaf infusion is
used to treat diabetes (Kokwaro 2009). Leaves are boiled to treat tonsillitis (Njoroge
and Bussmann 2006).
Rubus pinnatus: The leaves are boiled to treat cough (Luizza et al. 2013).
Rubus steudneri: Medicinal uses in Ethiopia (Lulekal et al. 2008). The leaves are
boiled for respiratory problems (Yineger et al. 2008). The roots are used for
stomachache with diarrhea (Giday et al. 2009).
Fig. 1 Rubus pinnatus

R. W. Bussmann)
Local Food Uses
Rubus apetalus: Fruits are eaten (Beentje 1994; Bussmann 2006a, b; Bussmann
et al. 2011).
Rubus volkensii: Fruits are eaten (Beentje 1994).
Rubus steudneri: Fruits are eaten (Bussmann et al. 2011; Luizza et al. 2013).
Rubus adolfi-friederici: Fruits are eaten (Bussmann et al. 2011).
Fig. 2 Rubus pinnatus

R. W. Bussmann)
Rubus steudneri: Browsed by cattle (Bussmann et al. 2011).

Rubus volkensii: Leaves are boiled to expel evil spirits (Yineger et al. 2008).
Fig. 3 Rubus volkensii

R. W. Bussmann)
Fig. 4 Rubus steudneri

R. W. Bussmann)
Fig. 5 Rubus steudneri

R. W. Bussmann)
References
2006b;111:41–66.
Governments & Administrations; 1960. p. 61. ISBN: 978-1-84246-310-9.
Res Appl. 2013;11:315–39.
10.1186/1746-4269-4-10.
2008;26:132–53.
Rumex abyssinicus Jacq.
Rumex usambarensis (Dammer) Dammer
POLYGONACEAE

Grace N. Njoroge
Synonyms
Rumex abyssinicus Jacq.: Rumex bequaertii var. quarrel (De Wild.) Robyns;
Rumex schimperi Meisn.
Rumex usambarensis (Dammer) Dammer: Acetosa trinervia Rech. f.) Á. Löve &
B. M. Kapoor; Acetosa usambarensis (Dammer) Degraeve; Rumex trinervius
Rech. f.
Local Names
Rumex abyssinicus: Kikuyu: Mugagatio; Swahili: Mchachu, Mchumvichumvi;

Kamba: Kinyonwe; Maa: Enkaiswishoi (Beentje 1994; Gachati 1989; Kokwaro
2009); English: Sorrel, Dock, Spanish rhubarb
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Rumex abyssinicus Jacq.: A large, very stout, perennial herb, up to 4 m tall. Nut
acutely trigonous, 2.25–3 mm 2 mm., shining, light brown or dark with lighter
angles. Stems glabrous, green or reddish-green, up to 3 cm wide at the base. Leaves
petiolate, large, up to 30 20 cm, lush, glabrous or papillose, usually triangular-
hastate but varying to sagittate, scutate, sublinear, or more rarely ovate; the lobes
spreading, more rarely directed forward; apically acute to very obtuse; with a
variably deep and wide basal sinus; primary nerves palmately arranged. Inflores-
cence a large, much branched, leafless panicle, oblong or pyramidal in outline, up to
40 cm long, 25 cm or more broad. Inner tepals wing-like, accrescent, elongate-
rounded, pellucid, reticulately veined, green becoming reddish-brown or brown,
5.5 4.5 mm. (7 7 mm., fide Rech. f.), with a basal sinus up to 1 mm deep,
and bearing a small reflexed protuberance at the base Petioles long, up to 14 cm,
those of the lower leaves often exceeding the lamina. Outer tepals ovate, 1.5 mm
long, membranous and brown, later reflexing. Flowers borne in fascicles, on filiform
pedicels up to 5 mm long (Graham 1958). In Afromontane forest glades, often under
heavy wildlife browsing (Bussmann 1996) (Figs. 1, 2, 3, 4, 5, 6, and 7).
Rumex usambarensis (Dammer) Dammer: A shrub or straggling glabrous climber,
up to 3 m tall or more. Stems brown. Leaves astringent, petiolate, often clustered,
narrowly to broadly elliptic with a hastate base, a little narrowed above the basal
lobes, up to 5–9 cm long, apically acute; lobes reflexed, small, not exceeding
5 2 mm; leaves markedly trinervate (except in very narrow leaves), the lateral
nerves arising arcuately from the base of the midrib and remaining complementarily
parallel throughout most of their length. Petioles 1–4 cm long. Inflorescence a much-
branched, slender, leafless panicle. Flowers in fascicles, on filiform pedicels up to
5 mm long. Outer tepals 1.75–2 mm long, ovate, obtuse, later reflexing. Inner tepals
wing-like, 5–7 mm in diameter, pellucid, subequal, circular when mature, with a
squarish basal sinus 1–1.5 mm deep, red or reddish-brown, reticulately veined and
with a small wart-like reflexed protuberance at the base. Nut trigonous, ovoid,
2–2.5 mm 1 mm, brown, shining. Open mist-forest, also upland grassland,
bushland and exposed rocky slopes, 870–2400 m. Widespread in tropical Africa,
most commonly in the highlands, particularly in central and eastern Africa, and
Madagascar (Graham 1958).
Rumex abyssinica: The root paste is applied for snakebite. The root infusion is used
to remedy stomachache, indigestion, and flatulence (Kokwaro 2009).
Rumex usambarensis: The sap of the aerial parts is applied as a treatment for
pneumonia, stomachache, and cough in eastern Africa. A decoction of the whole
Rumex abyssinicus Jacq. . . . 921
Fig. 1 Rumex
sp. (Polygonaceae),
open forest glades, Kenya.
plant was originally used for smallpox and serves to treat scabies (Kokwaro 2009).
The fresh leaves are snuffed for sore throat, and the leaf infusion can be used for
fever. In Ethiopia, the plant is used to treat jaundice and related liver diseases,
scrofula, stomachache, neckache and low blood pressure, and as a wound dressing,
hemostatic, and depurative; the rhizome is used as a taeniafuge. In DR Congo, a leaf
compress is applied to areas of rheumatism, an infusion is taken as a purgative, and
root sap is applied against scabies. In Tanzania, the stem and rhizome are believed to
act as a galactagogue. The whole plant, fresh or dried, is ground up in Tanzania and
placed on sores and parts affected by scabies. An extract of the rhizome is taken to
control mild forms of diabetes in eastern Africa and, with water, to cure stomach-
ache. Pounded rhizomes and roots are applied on wounds and are also considered to
have purgative properties. The leaves of Rumex nepalensis are used for stomach-
ache (Giday et al. 2009).
Fig. 2 Rumex
sp. (Polygonaceae),
Local Food Uses
Rumex abyssinicus: The tender shoots and leaves of both species are edible and
widely used as a vegetable. They have an acid taste and are eaten fresh or cooked,
alone or together with other vegetables (Beentje 1994). In Tanzania, the stem is
chewed like sugarcane for its sweetness, and the leaves are eaten as an acidic snack
by herdsmen, farmers, and children. The seeds of Rumex bequaertii and Rumex
steudelii can be eaten (Bussmann 2006).
Fig. 3 Rumex cf. crispus

(Polygonaceae), Mt. Kenya
National Park, open forest
glades, Kenya. (Photo: R. W.
Bussmann)
Rumex abyssinicus: The rhizomes yield a yellow and red dye. The dye is used in
Ethiopia in butter as a condiment, to give it a rich yellow color, and as protection
against rancidness. The dye is also used to impart a red color to the feet and hands of
women. In Uganda, the plant is occasionally cultivated to obtain the red dye for
coloring wickerwork and mats of grass and raffia, and in Rwanda to obtain the
yellow dye. The plant is browsed by livestock. In Rwanda and Tanzania, crushed
plants are used to scour clean cooking pots blackened over the fire and to remove
grease. A leaf paste of Rumex abyssinicus can be used to treat mastitis in cows
(Kokwaro 2009).
Fig. 4 Rumex cf. crispus

(Polygonaceae), Mt. Kenya
National Park, open forest
glades, Kenya. (Photo: R. W.
Bussmann)
Fig. 5 Rumex
sp. (Polygonaceae),
Fig. 6 Rumex
sp. (Polygonaceae),
Fig. 7 Rumex abyssinicus.

(Polygonaceae), c. 3600 m,
edge of Bale Mountains,
Oromiya Province, Ethiopia.
(Photo: J. Grimshaw)
References
Graham RA. Flora of Tropical East Africa: Polygonaceae. London: Crown Agents for Overseas
Salvadora persica L.
SALVADORACEAE

Grace N. Njoroge
Local Names
Salvadora persica: Swahili: Msuake, Mswaki, Musuake; Borana: Adi, Huda;

Dassenach: Nyeteh; Gabbra: Adde; Turkana: Esekon; Taita: Kizungumoto;
Gogo: Mkunguni; Giriama: Mueza-Moyo, Mjunmgumoto; Oromo: Ade, Dalkath;
Pokomo: Muade; Pokot: Asiokonyon; Rendile: Hagay; Samburu: Sakotei; Sanya/
Somali: Ade, Adhei; Ilwana: Musuaki; Kamba: Mukayau; Maa: Oremit, Eremit;
English: Toothbrush tree (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Salvadora persica L.: Bushy, often sub-scandent, evergreen shrubs and treelets,
60 cm–5 m tall, branches long and often drooping, more or less terete, glabrous or
occasionally loosely pubescent, the older whitish. Leaves rather thick, minutely pellucid
dotted, verrucose, dark to bright green, oblong-elliptic, ovate-oblong to suborbicular,
2.5–5 cm long (sometimes up to 9 cm long) and 1–3.8 cm broad, slightly narrowed to
R. W. Bussmann (*)
G. N. Njoroge

the rounded, mucronate apex and shortly cuneate at the base, petiole 5–10 mm long,
patently spreading to arcuate-ascending. Inflorescence of terminal and lateral panicles
(the terminal usually overtopping the leaves), with long slender branches, the lateral
panicles in the axils of the upper leaves only, flowers small greenish-yellow, on short,
about 1 mm long, pedicels. Calyx campanulate, about 2 mm long, shortly 4-lobed,
glabrous or very minutely papillose and scaly, occasionally puberulous. Corolla greenish
yellow, about 3 mm long, tube short, about 1 mm long, lobes oblong, obtuse and
sometimes notched at the apex, cucullate in bud, eventually the upper half reflexing over
the rim of the calyx. Stamens 4, inserted in the sinuses on the tube of the corolla,
filament usually just over 1 mm long and anther just under 1 mm long. Ovary 1-celled
with one erect basal ovule, style very small with an inconspicuous stigma. Fruit a drupe,
white tinted rose and red when mature, globose, smooth, about 5 mm diameter, seed
solitary, cotyledons large, endosperm none (Verdcourt 1968).
Salvadora persica: The most widespread use of toothbrush tree is for chewing sticks or
natural toothbrushes. Small twigs (around 3–5 mm in diameter) are used and have both
physical and antimicrobial action, helping to control plaque and prevent tooth decay.
Roots and stems contain numerous active compounds, including salvadorine and
benzylisothiocyanate, which inhibit bacteria that cause tooth decay, and tannins,
which reduce plaque and gum disease. Leaves can be made into a mouthwash with
similar properties. Chewing sticks have been used for dental hygiene as far back as the
time of the Babylonians (around 7000 B.C.). Salvadora persica is mentioned in the
Qur’an and the Bible (as “mustard seed” or “pepper bush”). The leaves are used to treat
a range of ailments, including stomachache, rheumatism, and sores. The root decoction
is used to treat gonorrhea, spleen problems, malaria, stomachache, and is used as
anthelminthic, as well a chest problems. The decoction is also used as galactagogue.
Pounded roots are applied as poultice for boils. The root latex is applied to sores. Tea
from dried bark is used for colds and fever (Bussmann 2006; Kokwaro 2009). In
Ethiopia used for boils, chest pain, abscesses, tuberculosis, cough with blood, flu,
malaria, and cancerous swellings (Teklehaymanot and Giday 2010). Used to treat
tick-borne diseases (Kioko et al. 2015). Also employed for stomachache, malaria,
constipation, and as anthelminthic (Muthee et al. 2011; Nankaya et al. 2019, 2020). In
Madagascar, Salvadora angustifolia is used to remedy diarrhea, premature fontanel
closure, and for mothers during pregnancy (Randrianarivony et al. 2016a, b, 2017).
Salvadora oleoides is used for toothache, colic, and as tonic in Pakistan (Umair et al.
2019).
Local Food Uses
Salvadora persica: The fruit can be eaten fresh, cooked, dried and stored, or made
into a fermented drink. The leaves have a bitter, peppery taste and are eaten as a
green vegetable or made into a sauce (Beentje 1994; Bussmann 2006).
Salvadora persica L. 929
Salvadora persica: When pressed, seeds yield oil that is rich in lauric and myristic
acids and can replace coconut oil in production of soaps and detergents. Being tough
and resilient to harsh conditions, toothbrush tree is often planted as a windbreak and
can be used to improve saline or impoverished soils. Leaves are browsed by cattle,
sheep, goats, and camels, although they are said to make milk taste bad. The flowers
are a good source of nectar for honeybees. The root decoction is given as emetic to
cattle with anthrax (Kokwaro 2009). Eaten by livestock (Bussmann 2006). The
decoction is given to cattle to expel the placenta (Bussmann 2006). Used to treat
tick-borne diseases (Kioko et al. 2015). In livestock used to treat mange and scabies
(Wanzala 2017).
References
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Medicinal plants of the maasai of Kenya: a review.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016a. ISBN: 978-0-9960231-4-6.
2017. https://doi.org/10.1186/s13002-017-0147-x.
10.1186/s13002-019-0285-4.
Verdcourt B. Flora of Tropical East Africa: Salvadoraceae. London: Crown Agents for Overseas
https://doi.org/10.1155/2017/8647919. Article ID: 8647919, 33 pages.
Sambucus africana Engl.
CAPRIFOLIACEAE

Grace N. Njoroge
Synonyms
Sambucus ebulus subsp. africana (Engl.) Bolli
Local Names
Sambucus africana: Kikuyu: Makomboki (Gachati 1989).
Botany and Ecology
Sambucus africana Engl.: Shrubby herb, 1.2–4 m tall, the roots spreading horizon-
tally and producing stems at intervals. Stems mostly juicy but somewhat woody at
the base, at first covered with multicellular hairs but later glabrous. Leaves
imparipinnate, attaining 75 cm in length and 45 cm in width, (5–) 6–11-foliolate;
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Sambucus africana (Caprifoliaceae), Mt. Kenya National Park, Naro Moru route. (Photo:
R. W. Bussmann)
leaflets subsessile, occasionally apical one partly fused with a lateral, elliptic-oblong,
ovate or oblong-lanceolate, 4.5–18 cm long, 1.3–8 cm wide, acuminate, very
unequally rounded at the base, finely and sharply toothed (the basal teeth often
glandular), at first hairy, later glabrous above except for minute hairs on the main
nerves, bullate, sparsely pubescent on the main nerves beneath which are slightly
raised and reticulate; petioles 0.6–12 cm long; stipules conspicuous, leafy, toothed
(true nature uncertain); on the rachis near the leaflet bases there are often glandular
stipules. Inflorescences corymbose, 7–15 cm wide, ferruginous pubescent, sub-
tended by 1–3-foliolate glandular-toothed leaves; bracts and bracteoles linear,
small. Calyx pinkish or purplish; tube ribbed; lobes triangular, about 1 mm long.
Corolla white or creamy-white; tube 1.5 mm long; lobes elliptic, 4–6 mm long,
3–3.5 mm wide, induplicate-valvate in bud. Fruit oblong or ellipsoid, 4.2–7 mm
long, 2.5–4 mm wide, blue-black, 9–10-ribbed; fruiting pedicels purplish. Pyrenes
3–4, bony, pale reddish-brown, linear-oblong, 4–5.5 mm long, 1–3 mm wide,
somewhat trigonous in section. Upland grassland or evergreen bushland, clearings
in or on the edges of moist bamboo thicket and upland rain forest, or sometimes in
upland rain forest undergrowth; 1800–3150 m. In Afromontane forests especially in
the bamboo zone and in disturbed areas where animals rest (Bussmann 1996, 2002a,
b, 2004, 2006) (Fig. 1).
Sambucus africana is an important species in the regeneration cycle of East
African Bamboo. The dense forests dominated by the African Bamboo, Oldeania
africana (in the following in this chapter, we use the much better used synonym
Sinarundinaria alpina), which are found on various East African mountains, form a
clear-cut class (Sinarundinarietea alpinae, Bussmann 1994, 1997, 1999a, b, 2002;
Bussmann and Beck 1995, 1999; Bussmann and Lange 1999; Bytebier and
Bussmann 2000) of their own. Sinarundinaria alpina (Poaceae), Impatiens hoehnelii
Sambucus africana Engl. 933
(Balsamiaceae) and Pseudocarum eminii (Umbelliferae), as well as Selaginella

kraussiana (Selaginellaceae), Cyperus dereilema, Anthriscus sylvestris (Umbelliferae),
and Cardamine obliqua (Cruciferae) are the most characterizing species of this
forest type.
soils, namely humic Andosols. On Mount Kenya, the Sinarundinarietea alpinae are
and a stem diameter of up to 10 cm. New vegetative shoots are sprouting from old
4500–7000 culms produced/hectare require longer periods of favorable climate.
die, occurs in areas of about one to several 100 acres in more or less regular intervals
of at least 40 years (Wimbush 1945). On Mount Elgon, Clayton (1970) observed a
flowering cycle of only 15 years.
Agnew (1985) referring to a long-term knowledge of the vegetation of the
1. Pioneer – Regeneration from sections of old rhizomes and rhizome fragments and
occasionally from fallen culms, lasting about 3–5– years during which dense
ever been reported from stands of Sinarundinaria (Figs. 2 and 3).
2. Building – During this period, the growing bamboo exceeds the average plant
getting momentum. The pH of the soil starts decreasing (Figs. 4, 5, and 6).
3. Mature – No further increase in culm height takes place during this stage. The culms
show even distribution and the soil pH is decreasing further (Figs. 7, 8, and 9).
4. Flowering and Sambucus Stage – Within 2–3 years after mass flowering, the

route. (Photo: R. W.
Bussmann)

Bussmann)
bamboo therefore could be indicating the sites of frequent flowering (Figs. 10, 11,
and 12).
Kenya (Bussmann 1994; Bussmann and Beck 1995; Bussmann and Lange 1999),
Mt. Nyiro in Northern Kenya (Bussmann 2002; Bytebier and Bussmann 2000), and
the Harenna Escarpment in the Ethiopian Bale Mountains (Bussmann 1997). How-
ever, the vegetation showed considerable floristic differences on wet and drier sites
of bamboo forests, even if the areas compared were in the same stage of the cycle.
In the North of Mt. Kenya, young bamboo was apparently invading parts of
Podocarpus latifolius forests with open undergrowth as a new habitat, forming
dense, more or less extended thickets. To clarify whether this represented a new
colonization, very young sprouts were excavated in several places, especially when
forming outposts far from young bamboo thickets. All samples indicated that the
new shoots arose from old rhizome fragments. Therefore, the areas which seemed to

Park, Naro Moru route.
be newly colonized, proved to be regenerating old bamboo stands, where

Sinarundinaria alpina had died many years ago, and very small vegetative bamboo
sprouts had been part of the heavily grazed pasture, so that the local people did not
remember bamboo in these locations (Mathenge, personal communication, 1993).
Since no old culms were found, they must have been decomposed a long time ago, as
dead culms normally stay for many years especially in the drier areas. Therefore, the
time gap between two subsequent bamboo cycles appears to be very long in this area,
i.e., regeneration seems to be very slow.
• The region is fairly dry, i.e., Sinarundinaria alpina is at the limit of its distribution
• The Sirimon area in the Northwest has a very high population of herbivores,

In Ethiopia, no distinct Sambucus-stage could be observed. Apparently,

Sambucus africana does not occur naturally in Ethiopia (Friis, personal communi-
cation); however, after dieback of the flowered bamboo, other light demanding
species occur, until the pioneer stage takes over again.
During the presented study, on Mount Kenya and the neighboring Aberdare
Range, bamboo stands were mainly found on the wetter western, southern, and
southeastern slopes and were absent in the Northern areas. From this distribution, it
was concluded that Sinarundinaria alpina requires precipitation of about 800–
1000 mm/year although arid periods are tolerated. In clear correspondence to the
observations of Wimbush (1945), new culms were mainly produced after prolonged
rainy areas. During the research period of 1992–1996, the main culm production
occurred in large areas in May/June 1995, after 9 months of unusually heavy rainfall.

Fig. 7 Sinarundinaria alpina (Poaceae), mixed bamboo – young regrowing bamboo stands,
Mt. Kenya National Park, Naro Moru route. (Photo: R. W. Bussmann)

Naro Moru route. (Photo:
R. W. Bussmann)
This, and the fact, that the newly produced culms were mostly much larger and taller
than the old ones clearly depicts the dependency of bamboo growth on rainfall
duration and intensity.
Mt. Nyiro, Harenna Escarpment (Bale Mts., Ethiopia)) could be observed. In clear
climatic conditions. Therefore, it has to be concluded, that in contrast to many Asian
Although covering wide areas (Harenna, Bussmann 1997), Illubabor (Friis, personal
communication), these forests were not recognized as specific forest types or not

Naro Moru route. (Photo:
R. W. Bussmann)
route. (Photo: R. W. Bussmann)
Fig. 11 Sambucus africana (Caprifoliaceae) – Sinarundinaria alpina (Poaceae), stage after bam-
boo dye-back, with young resprouting bamboo in the background, Mt. Kenya National Park, Naro
Moru route. (Photo: R. W. Bussmann)
mentioned at all by earlier authors. The large bamboo populations of South and
Central Kenya (Mt. Kenya, Aberdares, Mau, about 1000 km South of Harenna) are
linked to Harenna only by a small population occurring on top of Mt. Nyiro at the
southern end of Lake Turkana, which is also at least 500 km away from the Bale
mountains. As mentioned above, viable seeds of Sinarundinaria alpina have never
been found, and the species is known to regenerate only vegetatively, thus a
propagation by animals seems very unlikely. Therefore the large gaps between the
main population and even these outposts pose some interesting questions regarding
the ecology and former distribution of Sinarundinaria, as influenced by the
palaeoclimate of the region. This requires further fieldwork.
mentioned by different authors (Agnew 1985; Schmitt and Beck 1992) are regarded
as clearly differentiated subassociations of the respective higher taxonomic units.
Local Food Uses
The fruits are edible (Gachati 1989).

Fig. 12 Sambucus africana

(Caprifoliaceae) –
Sinarundinaria alpina
(Poaceae), stage after bamboo
dye-back, with young
resprouting bamboo in the
background, Mt. Kenya
Bussmann)
References
Agnew ADQ. Cyclic sequences in plant communities of the Aberdare mountains, Kenya. J East Afr
Nat Hist Soc Natl Mus. 1985;75:1–12.
management of a tropical mountain forest ecosystem [dissertation]. Bayreuth: Universität
Bayreuth; 1994.
Bussmann RW. Notes on the vegetation of Menagesha forest (Shoa Province, Ethiopia). In: BITÖK,
Afrika, vol. 64. Bayreuth: Bayreuther Forum Ökologie; 1999a. p. 105–18.
In: BITÖK, editor. Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika, vol. 64. Bayreuth: Bayreuther Forum Ökologie; 1999b. p. 29–44.
Ambasht NK, editors. Modern trends in applied terrestrial ecology. Amsterdam: Kluwer; 2002a.
p. 195–223.
2006;11(1):41–66.
Phytocoenologia. 1995;25(5):467–560.
Bussmann RW, Beck E. Vegetation units of Mount Kenya Forest Reserve – comment. In: BITÖK,
Afrika, vol. 64. Bayreuth: Bayreuther Forum Ökologie; 1999. p. 17–28.
BITÖK, editor. Vegetationsökologische, ethnobotanische und -faunistische Beiträge über
Äquatorial-Afrika, vol. 64. Bayreuth: Bayreuther Forum Ökologie; 1999. p. 1–16.
Flora of Tropical East Africa. Rotterdam: Balkema; 1970.
Dale IR, Greenway PJ. Kenya trees and shrubs. Nairobi: Buchanan; 1961.
Hedberg O. Vegetation belts of the East African mountains. Sven Bot Tidskr. 1951;45:140–99.
Mensch und Großwild. Jahrb. Verein z. Schutze der Bergwelt. 1992;57:207–26.
Speck H. Soils of the Mount Kenya Area, their formation, ecological, and agricultural significance.
In: Winiger M, editor. Mt. Kenya area. Contributions to ecology and socio-economy. African
studies series A1. Universität Bern; 1986. pp. 47–72.
Wimbush SH. The African alpine bamboo. Empire For J. 1945;24:33–9.
Schefflera volkensii (Harms) Harms
ARALIACEAE

Grace N. Njoroge
Synonyms
Schefflera volkensii (Harms) Harms: Astropanax volkensii (Harms) Lowry,

G.M. Plunkett, Gostel & Frodin.
Local Names
Schefflera volkensii: Kikuyu: Muthai; Kamba: Ndambili; Marakwet: Tingwa,

Tinwot; Maa: Ottini; Nandi: Ewait; Sebei: Kwelet, Kwalet; Nandi: Jinet; Kipsigis:
Dinet, Tinet (Beentje 1994, Gachati 1989, Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Schefflera volkensii

(Araliaceae), Mt. Kenya
National Park forest (Photo
R.W. Bussmann)
Botany and Ecology
Schefflera volkensii (Harms) Harms: Scandent shrub or tall (sometimes spreading

and much branched) tree up to 24–30 m tall, or an epiphyte upon other trees.
Flowers sessile, up to 12–20 together. Fruits more or less urceolate, up to 5.5 mm
long by 4(–5) mm diameter, more or less 5-ribbed, minutely puberulous or gla-
brous, stylopodium up to 7 mm long, styles 5, free and spreading for up to 0.5 mm
terminally. Stipules sheath-like, up to about 1–2(–l.4) cm long. Inflorescence
(Fig. 5/8) a more or less extended or compressed raceme of bracteate racemes of
small pedunculate more or less globular capitula up to 7 mm diameter when flowers
in bud, primary branches up to 23 cm long by 4.0(–5.5) mm diameter, generally
sparsely lenticellate, secondary branches (peduncles of capitula) (5–)10(–17) mm
long, sometimes lenticellate, borne in the axils of ovate to oblate bracts up to 4 mm
long. Petiole up to 13(–17) cm long by 3.2(–3.8) mm diameter, sometimes some-
what expanded at base, glabrous or sparsely hairy at tip, leaflets 4–6(–7), generally
coriaceous, narrowly obovate or elliptic, occasionally oblanceolate or obovate,
rarely narrowly or broadly elliptic, up to 15 cm long by 7 cm wide, acute to rounded
or rarely slightly retuse at apex, with a generally acute, sometimes acute to rounded
or broadly to narrowly cuneate base, with entire to very slightly repand, sometimes
Schefflera volkensii (Harms) Harms 945

(Araliaceae), Mt. Kenya
National Park forest (Photo
R.W. Bussmann)

(Araliaceae), flowers,
Mt. Kenya National Park
forest (Photo R.W. Bussmann)
very slightly crisped margins, glabrous, petiolules up to 2.1(–2.8) cm long by

1.9 mm diameter (Tennant 1968). Commonly found in Afromontane forests
(Bussmann 2002a, b, 2006a) (Figs. 1, 2, and 3).
Schefflera volkensii: The gum can be used for colds, cough, lung problems, and
bronchitis, often mixed with honey (Kokwaro 2009). Used to treat dermatitis (Luizza
et al. 2013; Yineger et al. 2008). In Madagascar, Schefflera longipedicellata is used
to treat epilepsy, colds, gonorrhea, and serves as anthelminthic and to expel the
placenta. Also used to treat plague (Razafindraibe et al. 2013). Schefflera bojeri is
sold in markets to treat ulcers, hepatitis, back pain, and neurasthenia
(Randriamiharisoa et al. 2015). In Pakistan, Schefflera arboricola is used for
wounds, cuts, indigestion, abdominal pain, and to improve the blood circulation
Schefflera volkensii: In rare cases used as firewood and eaten by colobus monkeys
(Bussmann 2006b; Bussmann et al. 2011). Schefflera abyssinica serves also for
firewood and as honey plant (Bussmann et al. 2011).
References
2006a;11(1):41–66.
Res Appl. 2013;11:315–39.
2015;11:60. https://doi.org/10.1186/s13002-015-0046.
women from Agnalazaha littoral forest southeastern Madagascar. J Ethnobiol Ethnomed.
2013;9:73.
Schefflera volkensii (Harms) Harms 947
Tennant JR. Flora of tropical East Africa: Araliaceae. London: Crown Agents for Overseas
10.1186/s13002-019-0285-4.
2008;26:132–53.
Schkuhria pinnata (Lam.) Kuntze ex Thell.
ASTERACEAE

Grace N. Njoroge
Synonyms
Schkuhria pinnata (Lam.) Kuntze ex Thell.: Amblyopappus mendocinus Phil.;

Chlamysperma polygama Triana; Hopkirkia anthemoides DC.; Mieria virgata La
Llave; Pectis pinnata Lam.; Rothia pinnata (Lam.) Kuntze; Rothia pinnata var.
pallida Kuntze; Rothia pinnata var. purpurascens Kuntze; Schkuhria abrotanoides
Roth; Schkuhria abrotanoides var. pomasquiensis Hieron.; Schkuhria advena Thell.;
Schkuhria anthemoides (DC.) J.M. Coult.; Schkuhria anthemoides fo. flava (Rydb.)
Heiser; Schkuhria anthemoides var. guatemalensis (Rydb.) Heiser; Schkuhria
anthemoides var. wisliseni (A. Gray) Heiser; Schkuhria anthemoides var. wrightii
(A. Grey) Heiser; Schkuhria bonariensis Hook. & Arn.; Schkuhria coquimbana
Phil.; Schkuhria guatemalensis (Rydb.) Standl. & Steyerm.; Schkuhria hopkirkiana
A. Gray; Schkuhria isopappa Benth.; Schkuhria octoaristata DC.; Schkuhria
pinnata fo. pringlei (S. Watson) Heiser; Schkuhria pinnata var. abrotanoides
(Roth) Cabrera; Schkuhria pinnata var. guatemalensis (Rydb.) McVaugh; Schkuhria
pinnata var. octoaristata (DC.) Cabreta; Schkuhria pinnata var. virgata (La Llave)
R. W. Bussmann (*)
G. N. Njoroge

Heiser; Schkuhria pinnata var. wislizeni (A. Gray) B.L. Turner; Schkuhria pringlei
S. Watson; Schkuhria virgata (La Llave) DC.; Schkuhria wislizeni A. Gray;
Schkuhria wislizeni fo. flava (Rydb.) S.F. Blake; Schkuhria wislizeni var. frustrata
S.F. Blake; Schkuhria wislizeni var. wrightii (A. Gray) S.F. Blake; Schkuhria
wrightii A. Gray; Tetracarpum anthemoideum (DC.) Rydb.; Tetracarpum flavum
Rydb.; Tetracarpum guatemalense Rydb.; Tetracarpum pringlei (S. Watson) Rydb.;
Tetracarpum virgatum (La Llave) Rydb.; Tetracarpum wislizeni (A. Gray) Rydb.;
Tetracarpum wrightii (A. Gray) Rydb.
Local Names
Schkuhria pinnata: Luo: Maua-Matindo, Onyalo-Biro (Kokwaro 2009).
Botany and Ecology
Schkuhria pinnata (Lam.) Kuntze ex Thell.: Annual herb 5–50 cm high; much
branched, the branches glandular and pilose. Leaves alternate, pinnatisect or the upper
simple and filiform, in outline 1–6 cm long, 0.1–2 cm wide, with 3–7 lobes, these
filiform or themselves divided, pilose or glabrous, glandular. Capitula turbinate or ovoid,
on stalks to 3.5 cm long; involucre 4.5–6 mm long; phyllaries 2–6 mm long with yellow
apex, glandular. Ray floret 1, sometimes absent, yellow, 1–2 mm long, glandular; disc
florets 4–8, 2 mm long with glandular lobes. Achenes black, 3–4.5 mm long, ribbed,
hairy; pappus scales 8, pale brown with purple marks or purplish, lanceolate, 1–2.5 mm
long, hairy (Beentje 2000, 2002). Common weed in the undergrowth of Afromontane
forests, introduced from South America (Bussmann 2002).
The whole plant is used for chest, liver, and stomach pains (Kokwaro 2009). Also
applied as antimalarial (Njoroge et al. 2004, 2006a, b; Njoroge & Bussmann 2006a, b).
In contrast to Africa, the plant is of great medicinal importance in South America,
where it is used for blood cleansing, liver problems, blood purification, gallbladder,
bad breath, diabetes, menstrual delay, allergies, menstruation, blood irrigation, and
inflammation of the urinary tract (Bussmann & Sharon 2006).
References
Schkuhria pinnata (Lam.) Kuntze ex Thell. 951
larial ethnophytotherapeutic remedies among the kikuyus Central Kenya. J Ethnobiol
Scutia myrtina (Burm. f.) Kurz
RHAMNACEAE

Grace N. Njoroge
Synonyms
Scutia myrtina (Burm. f.) Kurz: Adolia alba Lam.; Adolia capensis Kuntze; Adolia
myrtina (Burm.f.) Kuntze; Adolia obcordata (Biv. ex Tul.) Kuntze; Adolia rubra
Lam.; Blepetalon aculeatum Raf.; Catha zeylanica G.Don; Ceanothus capensis DC.;
Ceanothus circumcissus (L. f.) Gaertn.; Rhamnus capensis Thunb.; Rhamnus
circumcissa L. f.; Rhamnus myrtina Burm. f.; Scutia capensis fo. obcordata (Boivin
ex Tul.) Radkl.; Scutia circumcissa (L. f.) W. Theob.; Scutia commersonii Brongn.;
Scutia eberhardtii Tardieu; Scutia hutchinsonii Suess.; Scutia indica Brongn.; Scutia
lucida G. Don; Scutia myrtina var. emarginata Bhandari & Bhansali; Scutia
natalensis Hochst.; Scutia obcordata Boivin ex Tul.; Ziziphus capensis Thunb.
ex Poir.
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Scutia myrtina: Kikuyu: Mulangari, Murangari, Mutanda Mbogo; Kamba:

Kitumbii; Boni: Tsina; Chagga: Mrega; Giriama: Munyahi; Kipsigis: Simbejwet;
Luo: Migodha, Osiri; Maa: Ol-Sanan-gurudei; Samburu: Sanunguri; Sebei:
Letwa; Tugen: Sumbioyawa (Beentje 1994, Gachati 1989, Kokwaro 2009).
Botany and Ecology
Scutia myrtina (Burm. f.) Kurz: Shrubs or rarely small trees 2–5(–10) m tall,
usually scandent. Bole to 30 cm thick; older trunks with thick dark corky longitu-
dinally cracked bark; younger herbage and young inflorescences puberulent; branch-
lets angular, usually emerging at right-angles, numerous. Thorns prickle-like,
recurved axillary, solitary (but often 2 per node), 2–10(–15) mm long. Leaf blade,
ovate to obovate, 2–6 cm long, 15–40 mm wide, at base rounded to cuneate, at apex
(rarely slightly acuminate to) rounded (sometimes apiculate) or refuse or emarginate,
always mucronulate, entire or in the distal two-thirds with a few (up to 9 on each
side) indistinct crenulations or appressed teeth, on each side of midrib with 5–8
obscure secondary nerves; petioles 3–10 mm long. Stipules 2–3 mm long, quickly
deciduous. Cymes often fascicle-like, 2–20-flowered (never more than 1 flower per
cyme maturing fruit); peduncles 2–7 mm long; pedicels 1–2(–3) mm long in flower,
2–3 mm in fruit. Sepals 1.2–2 mm long. Petals 0.7–1 mm long. Drupe 7–9 mm long
and thick, pallid when very young, ripening through red to purplish black; endocarps
6 mm long, readily separating into 2(–3) free indehiscent stones. In a wide variety
of situations from forest margins to bushland, thicket and wooded grassland; from
sea level to 2700 m (Johnston 1972). The species is characteristic component of drier
Afromontane forests (Bussmann 2002a, b) (Figs. 1, 2, and 3).
Scutia myrtina: A root decoction is used as anthelminthic and for gonorrhea. A leaf
paste is used for ringworm and wounds (Kokwaro 2009). Used also as antimalarial
(Njoroge and Bussmann 2006).
Scutia myrtina (Burm. f.) Kurz 955
Fig. 1 Scutia myrtina

(Rhamnaceae), Mt. Kenya
Kenya. (Photo
R.W. Bussmann)

Kenya. (Photo
R.W. Bussmann)

Kenya. (Photo
R.W. Bussmann)
Local Food Uses
Scutia myrtina: The fruit is edible (Beentje 1994; Bussmann 2006). The leaves are
boiled for soup (Bussmann 2006).
Scutia myrtina: Used for firewood. The leaves are browsed by livestock (Bussmann
2006).
References
Scutia myrtina (Burm. f.) Kurz 957
Johnston MC. Flora of tropical East Africa: Rhamnaceae. London: Crown Agents for Overseas
Governments & Administrations; 1972. 40 pp. ISBN 0-85592-019-X.
Ethnomed. 2006;2:8.
Solanecio mannii (Hook. f.) C. Jeffrey
ASTERACEAE

Grace N. Njoroge
Synonyms
Solanecio mannii (Hook. f.) C. Jeffrey: Crassocephalum mannii (Hook. f.) Milne-
Redh.; Crassocephalum multicorymbosum (Klatt) S. Moore; Senecio acervatus
S. Moore; Senecio bogoroensis De Willd.; Senecio congolensis De Willd.; Senecio
mannii Hook. f.; Senecio mannii var. kikuyuensis Chiov.; Senecio morrumbalensis
De Willd.; Senecio multicorymbosus Klatt
Local Names
Solanecio mannii: Kikuyu: Muthakwa-Wathi, Musariki, Mwathathi, Muthariti;

Kamba: Mooa, Ilangala; Pare: Mdaa; Shambaa: Gulanguso; Kipsigis: Chepkurbet;
Luo: Maroo, Marowo; Maa: Ol-musakawa-entim, Ol-airamirami; Marakwet:
Tergekwa, Yerekegwa; Meru: Mutumboro; Samburu: Lugugutt; Taita:
Mukorombosha; Tiriki: Kaibaraki (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Solanecio mannii (Hook. f.) C. Jeffrey: Erect softly woody shrub or small tree 1–8
(–10) m high, with foul smell, stems rather thick and succulent, green, with prom-
inent persistent leaf-scars, branching in a dichotomous manner (like frangipani) to
form a rounded crown, trunk up to 15 cm in diameter and with thin, smooth grayish
or greenish bark. Achenes 2–3.5 mm long, ribbed, shortly hairy in the grooves,
pappus 3.5–8.5 mm long. Ray florets absent, disc florets yellow, less often orange or
white, corolla 4.5–8.5 mm long, tube expanded above the middle, glabrous or
shortly hairy in upper part, lobes 0.7–2 mm long. Capitula discoid, very numerous
in large thyrses of about congested sub-umbelliform corymbs, stalks of individual
capitula glabrous or shortly pubescent, scented, involucre cylindrical, 4–7.5 mm
long, 1.5–2 mm in diameter, bracts of calyculus 3–4, lanceolate or oblong-
lanceolate, pubescent at least on the margins, 1–3 mm long, phyllaries 5–8, green,
sometimes with purple tips, 3.5–7 mm long, glabrous or thinly pubescent. Leaves
crowded towards the apices of the branches, slightly succulent, obovate, oblanceo-
late, or elliptic, 10–55 cm long, 2–16 cm wide, attenuate into an exauriculate
petioloid base, margins shallowly to prominently sinuate-serrate or sinuate-laciniate,
apex acute, acuminate-apiculate, glabrous or pubescent to thinly floccose especially
on main veins above, paler and glabrous to floccose or tomentose beneath, some-
times the lower midrib purplish. (Beentje 2000, 2002). Common at the edge of semi-
deciduous Afromontane forests (Bussmann 2002).
Solanecio mannii: The roots are used as anthelminthic and purgative, for diarrhea
and indigestion. Pounded leaves are applied to snakebites (Kokwaro 2009). Used to
treat anaplasmosis, theileriosis, and upper respiratory tract infections (Njoroge and
Bussmann 2006). The leaves of Solanecio angulatus are used to treat hepatitis
Solanecio mannii: Often used in hedges as ornamental (Beentje 1994). A root

decoction is used for the treatment cancer, pneumonia, epilepsy, cough, and typhoid
(Jeruto et al. 2008). The leaves of Solanecio angulatus are boiled in Ethiopia, and
the steam in haled to expel spirits (Bussmann et al. 2011). Also, 5–7 days after giving
birth the new mother’s body is washed with the plant to protect from evil. Inhaled
vapors during delivery induce sweating which helps rid the body of evil spirits and
keeps the mother warm (Luizza et al. 2013). Also burnt in social settings
Solanecio mannii (Hook. f.) C. Jeffrey 961
References
Res Appl. 2013;11:315–39.
Solanum aculeastrum Dunal
Solanum anguivi Lam.
Solanum incanum L.
Solanum nigrum L.
SOLANACEAE

Grace N. Njoroge
Synonyms
Solanum anguivi Lam.: Solanum indicum L.; Solanum indicum var. lividum (Link)
Bitter; Solanum indicum var. maroanum Bitter; Solanum lividum Link; Solanum
richardii Dunal; Solanum sodomeum L.
Solanum incanum L.: Solanum bojeri Dunal; Solanum sanctum L.
Solanum nigrum L.: Solanum humile Lam.; Solanum nigrum var. atriplicifolium
G. Mey.
Local Names
Solanum aculeastrum: Kikuyu/Embu: Gitura, Mutura; Kipsigis: Siganet; Maa:

Osigawai; Marakwet: Sikawa (Beentje 1994; Gachati 1989; Kokwaro 2009).
Solanum anguivi: Kikuyu: Mutongo, Mukenye; Marakwet: Jibokemnerkeny
(Beentje 1994, Gachati 1989, Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Solanum incanum: Kikuyu: Mutongu; Teso: Etulerut; Maa: Endulelei;

Irawq: Hangali; Luhya: Indalandalwa, Maduranzura; Marakwet: Labotwa;
Luo: Machoge, Ochok; Taita: Mrongo; Digo: Mtunguza Koma; Kamba:
Mukondu, Mutungu; Meru: Mutongo; Hehe: Ndula; Swahili: Mtunguja,
Mtunguja-mwitu, Mnyanya-mwitu, Mnyanya-porini, Mtua, Mtula (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: Thorn apple, Gray bitter apple, Bitter
tomato, Sodom’s apple
Solanum nigrum: Kikuyu: Managu; Giriama: Munavu; Kamba: Ndulu; Acholi:
Ocuga; Luo: Osuga (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Solanum aculeastrum Dunal: Erect shrub to small tree, 1–6 m, prickly. Young stems
terete, densely stellate-pubescent and prickly, with porrect to multangulate, sessile or
variously stalked trichomes, the stalks to 2 mm long, the rays 8–12, 0.1–0.5 mm long,
the midpoints shorter than the rays or up to 1 mm long, the prickles 8–17 mm long,
3–10 mm wide at base, curved, occasionally straight, strongly flattened, prominent,
orange to orange-brown, glabrous to sparsely stellate-pubescent in the lower half,
spaced 5–20 mm apart; bark of older stems stellate-pubescent to glabrescent, gray to
red-brown or brown. Sympodial units plurifoliate. Leaves lobed, the blades 8–15 cm
long, 6–12 cm wide, 1.5–2 times longer than wide, ovate, sometimes elliptic,
chartaceous, drying strongly discolorous; adaxial surface red-green to yellow-green,
glabrescent; abaxial surface white-gray, densely stellate-pubescent, with porrect and
multangulate, sessile or stalked trichomes, the stalks to 0.1 mm long, the rays 8–12,
0.15–0.8 mm long, the midpoints same length as the rays or up to 1 mm long, with 0–2
prickles on both surfaces; the primary veins 3–6 pairs, the tertiary venation visible on
both surfaces; base cordate to truncate, sometimes auriculate; margins lobed, the lobes
2–3 on each side, 1.5–4.5 cm long, extending 1/4–3/4 of the distance to the midvein,
broad-deltate, often with secondary lobing, apically obtuse, sometimes rounded; apex
acute; petiole 1.2–3 cm long, ca. 1/6 of the leaf blade length, densely stellate-
pubescent, with 0–2 prickles. Inflorescences apparently terminal or lateral, 3–6 cm
long, unbranched or rarely branched, with 4–12 flowers, 1(–5) flowers open at any one
time; axes densely stellate-pubescent, unarmed; peduncle 0–3.5 cm long; pedicels
1–3.5 cm long, erect, articulated at the base, densely stellate-pubescent, with 0–2
prickles on long-styled flowers, unarmed on short-styled flowers; pedicel scars spaced
1–4 mm apart. Flowers 5-merous, heterostylous and the plants andromonoecious, with
the lowermost 1(–2) flowers long-styled and hermaphrodite, the distal flowers short-
styled and staminate. Calyx 3–15 mm long, with 2–10 prickles in long-styled flowers,
unarmed in short-styled flowers, the lobes 2–7 mm long, deltate, apically acute to
apiculate, densely stellate-pubescent. Corolla 1.5–3.2 cm in diameter, white, some-
times mauve, stellate, lobed for 1/2–4/5 of the way to the base, the lobes 6–15 mm
long, 2–5 mm wide, broad-deltate, spreading, densely stellate-pubescent abaxially, the
trichomes porrect, sessile, the rays 8–12, 0.1–0.3 mm long, the midpoints shorter than
the rays. Stamens equal, with the filament tube 2–3 mm long and the free portion of the
Solanum aculeastrum Dunal . . . 965
filaments 0.5–1.5 mm long; anthers 4.5–6.5 mm long, connivent, tapering, poricidal at

the tips. Ovary densely stellate-pubescent in the upper 1/5; style 0.7–1 cm long in
long-styled flowers, slender to stout, straight to recurved, sparsely stellate-pubescent
near the base. Fruit a spherical to ellipsoid berry, often apiculate or acute at the distal
end, retaining the same shape throughout development or widening with age, 1(–3) per
infructescence, 3–5 cm long, 2–4.5 cm wide, the pericarp smooth to warty, evenly
green when young, green-yellow to yellow at maturity; fruiting pedicels 2.5–4.5 cm
long, 1.8–3.5 mm in diameter at base, woody, erect to pendulous, with 0–5 prickles;
fruiting calyx lobes not elongating, covering ca. 1/6 of the mature fruit or less,
moderately reflexed, with 0(–10) prickles. Seeds ca. 50–100 per berry, 4–5 2.7–
4 0.3–0.8 mm, flattened-reniform, almost round, often somewhat irregular in
outline, yellowish to pale orange-brown, the surface smooth or with raised outlines
of cells or small pits (Edmonds 2012). In disturbed, open parts of Afromontane forests
(Bussmann 2002) (Fig. 1).
Solanum anguivi Lam.: Shrub, 0.7–2 m, erect, armed or unarmed; young stems
stellate-pubescent; trichomes porrect, orange-translucent, sessile or shortly stalked,
stalks to 0.1 mm, rays 6–8, 0.1–0.25 mm, midpoints same length as rays or to
2 mm; prickles up to 5(–10) 1–4 mm at base, straight to curved, rounded to flattened.
Leaf blades drying discolorous, green-brown, ovate to elliptic, (5–)11–25 5–17 cm,
1.5–2.5 times longer than wide, base cuneate to rounded, usually unequal or oblique,
margin usually lobed, sometimes subentire, the lobes up to 5 on each side, broadly
Fig. 1 Solanum aculeastrum

(Solanaceae), Mt. Kenya
R.W. Bussmann)
deltate, to 2(–3) cm long and rounded to acute, extending up to 1/3(–1/2) of the

distance to the midvein, apex acute(acuminate); sparsely to densely stellate-pubescent
on both sides with trichomes on abaxial surface porrect, sessile or shortly stalked,
stalks to 0.1 mm, rays 6–8, 0.1–0.3 mm, midpoints same length as rays or up to 1.5 mm
long, adaxially with reduced rays and elongated midpoints; primary veins 4–7 pairs;
petiole 1–5 cm, 1/4–1/6 of the leaf length. Inflorescences 2.5–6 cm long, not branched
or branched once, with 5–22 flowers, peduncle 2–20 mm long; rachis 0.6–5 cm long;
peduncle and rachis unarmed; pedicels 0.5–1.2 cm long in flower, in fruit 0.8–1.3 cm
long, usually unarmed. Flowers perfect, 5-merous. Calyx 3–5 mm long, the lobes
deltate to somewhat ovate, 1–2 mm long, acute, unarmed or with 1–4 prickles. Corolla
white to mauve, 0.8–1.5 cm in diameter, lobed for 2/3 of its length, the lobes deltate,
3–6 2–3 mm. Stamens equal; anthers 3.5–5 mm. Ovary with a few stellate trichomes
towards the apex; style 6–8 mm long. Berries 6–22 per infructescence, evenly green
when young, red at maturity, spherical, 6–9 mm in diameter; fruiting calyx unarmed.
Seeds 1.8–2.5 1.8–2.3 mm. Forest and forest edges, wooded grassland, grassland,
bushland, disturbed areas, old cultivation, and roadsides; (40–)1000–2200(–3100)
m. Widespread in Madagascar, Senegal, Gambia, Guinea Bissau, Sierra Leone, Libe-
ria, Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Equatorial Guinea, Central African
Republic, Congo-Kinshasa, Rwanda, Burundi, South Sudan, Ethiopia, Angola, Zam-
bia, Malawi, Mozambique, Zimbabwe, and South Africa (Edmonds 2012). In dis-
turbed, open parts of Afromontane forests (Bussmann 2002).
Solanum incanum L.: Erect or spreading shrub up to 3 m tall, occasionally a small
tree; stems and leaves with stellate hairs and pale yellow to brown prickles, up to
1 cm long. Leaves alternate, simple; stipules absent; petiole 0.5–8.5 cm long; blade
almost round to lanceolate, 1–30 cm 1–17 cm, base rounded, truncate or cordate,
often unequal, apex acute or obtuse, margin entire to pinnately lobed, densely hairy.
Inflorescence a 2–15(–26) flowered cyme, inserted above the leaf axil. Flowers
bisexual or functionally male, nodding or pendent, regular, (4–)5–7(–9)-merous;
pedicel 0.5–4 cm long; calyx campanulate, lobes up to 1.5 cm long, enlarging and
splitting in fruit; corolla campanulate to rotate, 1–4.5 cm in diameter, with ovate or
broadly triangular lobes, blue, pink, purple or violet, rarely white; stamens inserted
near the base of the corolla tube and alternating with corolla lobes, filaments short,
anthers slender; ovary superior, 2(–4)-celled, style up to 15 mm long, densely hairy.
Fruit a globose or depressed globose, occasionally ovoid-ellipsoid berry
2.5–3.5 cm 2–3 cm, yellow, orange or brown when ripe, many-seeded. Seeds
lentil-shaped to almost kidney-shaped, up to 3.5 mm 3 mm, pale yellow to brown
(Edmonds 2012). In disturbed, open parts of Afromontane forests (Bussmann 2002)
(Figs. 2 and 3).
Solanum nigrum L.: Erect or widely spreading annual to short-lived perennial,
sometimes forming dense clumps up to 2 m across and 1.5 m high, (5)25–70(150)
cm tall, sometimes flushed with purple, pubescent to villous with simple, curved,
usually appressed, eglandular or glandular hairs, glabrescent. Branches angular to
narrowly winged, the edges smooth or with widely spaced short teeth or prickle-like
hairs. Leaves solitary or sometimes paired; petiole 0.3–6.5 cm long, narrowly
winged distally; lamina 2–14 0.7–7.3 cm, elliptic to ovate-lanceolate or ovate,
Fig. 2 Solanum incanum

R.W. Bussmann)
Fig. 3 Solanum incanum

R.W. Bussmann)
base subtruncate or rounded to cuneate, oblique, decurrent into the petiole, apex
acute or obtuse, acuminate, entire to sinuate-dentate, both surfaces pubescent,
with more numerous hairs along the veins and towards the margins, glabrescent.
Cymes lateral, unbranched, lax and often extended, (3)5–10-flowered; peduncle
0.3–2 cm long, slender, erect or ascending, in fruit occasionally elongated a little,
ascending to deflexed; pedicels 3–8.5 mm long, slender, decurved to ascending, in
fruit elongated to 12 mm, deflexed. Calyx 1.2–2.5 mm long, campanulate, in fruit
Fig. 4 Solanum nigrum

(Solanaceae). Fruits.
elongated to 3.5 mm; lobes 0.3–1.2 0.3–1.2 mm, obovate, semi-elliptic or ovate
to triangular, rounded to acute, sparsely pubescent with more numerous hairs
towards the apex outside, in fruit enlarged to 3 2 mm, appressed or scarcely
reflexed. Corolla white, cream or yellowish, with or without a conspicuous yellow-
green eye, rarely flushed with purple or pale blue, (4)5–7(9) mm across, rotate
or stelliform; lobes 1.5–4 0.8–2.3 mm, oblong or ovate to lanceolate, rarely
linear, acute, puberulous outside. Stamen filaments 0.3–1.5 mm long; anthers
1.5–2.5(2.8) mm long, oblong. Ovary 1 mm in diameter, globose to ellipsoid,
glabrous; style 2–4.5 mm long, straight or sigmoidal, the stigma level with the tip of
the anthers or protruding by up to 2 mm. Fruits often dull, black or purplish-black to
yellow-green, opaque, 6–10 mm in diameter, usually broadly ovoid, smooth, usually
without sclerotic granules, glabrous, edible when mature. Seeds with light bone-
color and sometimes flushed with purple, 1.7–2.4 mm long, obovate in outline,
minutely pitted all over. In disturbed, open parts of Afromontane forests (Bussmann
2002) (Figs. 4, 5, 6, and 7).
Solanum aculeastrum: Root decoction to treat gonorrhea (Beentje 1994). Roots

also used to treat bronchitis (Kokwaro 2009). For back pain, diarrhea, tonsillitis,
toothache, and wounds (Njoroge et al. 2004).
Solanum anguivi: The fruits are chewed for chest pains and cough. Roots chewed
for toothache (Kokwaro 2009). Used to treat tinea versicolor, earache, eczema, fever,
hemorrhoids, stomach problems, tuberculosis (Yineger et al. 2008), and for lymph-
adenitis (Giday et al. 2009).
Solanum incanum: Widely used as analgesic. Throughout tropical Africa used for
sore throat, angina, stomachache, colic, headache, painful menstruation, liver pain
and pain caused by onchocerciasis, pleurisy, pneumonia, and rheumatism The leaf,
root, and fruit decoctions are gargled or drunk, roots are chewed and sap swallowed,

(Solanaceae). Flowers.
leaf paste, root infusions, and pounded fruits are applied externally or rubbed into
scarifications, leaf sap is used for washing painful areas, and ash of burnt plants is
mixed with fat and applied externally. For relief of toothache, a root infusion is used
as mouth wash, fruit or root is rubbed on the gums or smoke of burning seeds is
inhaled. Hiccups are suppressed by licking a mixture of the ash of burned leaves and
salt. Another widespread use is in the treatment of venereal diseases. The root
powder is mixed with food or rubbed into scarifications, root infusions or decoctions
are drunk, roasted pulverized roots are taken in water, leaf decoctions and fruit sap
are drunk, and fruit sap is applied externally. Different plant parts are also widely
used in the treatment of skin problems, including skin infections, whitlow, ring-
worm, burns, sores, rashes, wounds, warts, carbuncles, ulcers, and benign tumors. In
Senegal, a maceration of the leaves is used as an eye bath to cure ophthalmia; in
Malawi, fruit sap is rubbed into scarifications around the eye to treat conjunctivitis.
In Senegal, pounded seeds are mixed with pulped fruits to massage aching ears. In
Uganda, Tanzania, and South Africa, extracts of leaves or flowers are used as ear
drops to cure inflammations. In Senegal, Kenya, Uganda, and Zimbabwe, different
plant parts are used to treat snakebites: a decoction of the roots is drunk, roots are
chewed, and sap is swallowed, and young chewed leaves or pulped fresh roots are
applied to the bite wound. In Kenya, the fruit pulp is applied to bleeding wounds,
warts, and for toothache (Beentje 1994). The root decoction is also used for
abdominal pain, dyspepsia, fever, stomachache, indigestion, and toothache. Young

(Solanaceae). Bakuriani,
Georgia. (Photo:
R. Bussmann)
Fig. 7 Solanum
sp. (Solanaceae), Mt. Kenya
R.W. Bussmann)
leaves are chewed and applied to snakebites, and leaf juice is applied to earache.
The fruit juice is applied to fresh wounds, for skin problems, as well as ringworm,
and taken for syphilis. The fruits are used as emetic but are toxic and need to be used
with care. Formerly also used to treat polio (Kokwaro 2009). The juice is applied to
wounds and cults (Bussmann et al. 2006). Used for fever, throat infections, and
wounds (Bussmann 2006). Used also to treat cough (Bekalo et al. 2009; Wondimu
et al. 2007). For chest pain, diarrhea, amoebic dysentery with blood, stomachache
(Nankaya et al. 2019, 2020; Njoroge et al. 2004; Teklehaymanot 2009;
Teklehaymanot and Giday 2010), and external wounds (Nankaya et al. 2019,
2020; Teklehaymanot 2009), also for heartburn, body pain, colds and cough, dys-
pepsia, ringworm (Lulekal et al. 2008; Nankaya et al. 2019, 2020), and especially
throat infections (Muthee et al. 2011). Leaves are chewed for digestion (Luizza et al.
2013). Sometimes used also to remedy snakebites and chest pain (Giday et al. 2003).
Solanum nigrum: The stems and roots are chewed to relieve toothache and inflam-
mations of the gums (Batsatsashvili et al. 2017; Bussmann et al. 2014, 2016a, b, c,
2017a, b; 2018; Bussmann 2017). In Pakistan, the species is used to treat leprosy
(Ur-Rahman et al. 2018), as well as breast cancer, diarrhea, febricity, ulcers, chicken
pox, hyperglycemia, piles, cardiac pain, sore eyes, cuts and wounds (Umair et al. 2019),
gastralgia, and infertility (Sher et al. 2016). In the Himalayas, Solanum nigrum is
employed for diarrhea, vomiting, asthma, fever, bronchitis, cuts and wounds, and
involuntary urinary discharge (Raj et al. 2018), while the species serves for typhoid
treatment in Kenya (Njoroge et al. 2004). In traditional Indian medicine, the species
serves as antidote for opium, ingestion, boils, cough, diarrhea, dysentery, ear complaints,
eye complaints, fever, goiter, heart ailment, jaundice, liver complaints, inflammation
(scrotum, kidney, and bladder), sores, nostril complaints, piles, rheumatism, skin dis-
eases, sprain, stomachache, swelling, throat trouble, ulcers in the mouth and urine
complaints (Verma et al. 2007), and the fruits are used for heart disease (Kunwar et al.
2012). Used for fever, throat infections, and wounds (Bussmann 2006). Used also to
treat typhoid (Njoroge et al. 2004).
Solanum marginatum has medicinal uses in Ethiopia, especially for swellings
(Balemie and Kebebew 2006; Mekonnen et al. 2015; Mohagheghzadeh and Faridi
2006; Teklehaymanot 2009; Teklehaymanot and Giday 2007, 2010). Solanum
aculeastrum, Solanum bendnerianum, Solanum renschii, Solanum
schumannianum, and Solanum terminale are used for fever, throat infections,
and wounds (Bussmann 2006). Solanum dasyphyllum is used to remedy cough
and trypanosomiasis (Bekalo et al. 2009), and for nosebleed (Giday et al. 2010).
Solanum adoense serves for nosebleeds, stomach problems, and scabies (Yineger
et al. 2008). Solanum hastifolium serves for diarrhea, amoebic dysentery with
blood, and stomachache (Teklehaymanot and Giday 2010). Solanum mauritianum
is applied to skin rashes (Njoroge et al. 2004). Solanum campylacanthum serves for
scabies and eczema (Gedif and Hahn 2003).
Local Food Uses
Solanum aculeastrum: Fruits are edible but bitter (Beentje 1994).

Solanum incanum: Selection for larger, less toxic fruits and leaves has taken place
over time whereas the wild types produce small, bitter, and toxic fruits; cultivated
types are available, especially in West Africa, of which the fruits and leaves can be
safely eaten in soup and as a vegetable. Leaves are added to soup to improve the
flavor. The large variation in toxicity makes it dangerous to transfer specific uses
from one region to another. The fruit juice can be used to clot milk (Borana) (Beentje
1994).
Solanum nigrum: Mature fruits with sweetish taste are used in food raw (limited)
(Bussmann et al. 2011) and to make jam. Young leaves are used like spinach
(Balemie et al. 2006; Batsatsashvili et al. 2017; Bussmann et al. 2014, 2016a, b, c,
2017a, b, 2018).
Solanum aculeastrum: Used for hedges (Beentje 1994) and as fishing poison
(Neuwinger 2004).
Solanum incanum: In Niger, Sudan, Rwanda, and Namibia, the fruits are used as an
ingredient of arrow poison and in Mozambique as fish poison. In Ethiopia, fruit sap
is mixed with butter and applied to cattle to control ticks. The fruit juice is squeezed
into sheep’s nostrils to treat cough, and into the eyes for conjunctivitis, as well as to
wounds (Kokwaro 2009). Used to treat nasal bots in livestock and is used as
acaricide (Wanzala 2017). Makes for very bad firewood (Tian 2017). Treats blackleg
in livestock (Yineger et al. 2007) and is used as fishing poison (Neuwinger 2004). In
livestock employed to treat hearty water (Kioko et al. 2015). Leaves are applied to
baking pans to avoid sticking of the dough (Luizza et al. 2013).
Solanum nigrum: A dye solution is prepared from flowers, leaves, and fruits to
obtain yellow and orange colors and used for dyeing wool yarn as well as products
made of wool (Batsatsashvili et al. 2017; Bussmann et al. 2014, 2016a, b, c, 2017a,
b, 2018).
Solanum marginatum is used for dermatophytes (Yineger et al. 2008).
References
Batsatsashvili K, Mehdiyeva N, Kikvidze Z, Khutsishvili M, Maisaia I, Sikharulidze S, Tchelidze D,
Alizade V, Paniagua Zambrana NY, Bussmann RW. Solanum nigrum L. In: Bussmann RW, editor.
Ethnobotany of the caucasus. Cham: Springer International Publishing; 2017.
doi.org/10.1186/1746-4269-5-26.
Bussmann RW. Ethnobotany of the Samburu of Mt. Nyiru, South Turkana. J Ethnobiol Ethnomed.
2006;2:35.

Shanshiashvili T, Chelidze D, Batsatsashvili K, Bakanidze N. Wine, Beer, Snuff, Medicine and
2014;12:237–313.
Samtskhe-Javakheti, Tusheti, Svaneti, and Racha-Lechkhumi, republic of Georgia
(Sakartvelo), Caucasus. J Ehnobiol Ethnomed. 2016a;12:43. https://doi.org/10.1186/
s13002-016-0110-2.
(Republic of Georgia), Caucasus. Med Aromat Plants. 2016b;5:5.
Khutsishvili M, Batsatsashvili K, Hart RE. Medicinal and food plants of Tusheti, Khevsureti and
Pshavi, Sakartvelo (Republic of Georgia), Caucasus. Acta Soc Bot Pol. 2016c;
(Republic of Georgia), Caucasus. Indian J Tradit Knowl. 2017a;16(1):25–34.
Edmonds, J.M.. Flora of tropical East Africa: solanaceae. Ed. Beentje, H. Royal Botanic Gardens,
Kew. 2012. ISBN 978-1-84246-395-6. Reference page. [Fl. Trop. E. Africa, Solan.]
doi.org/10.1186/1746-4269-5-34.
borne diseases among Maasai people in northern Tanzania. Veterinary World. 2015;812
Res Appl. 2013;11:315–39.
4269-4-10.
Mekonnen, T., Giday, M., Kelbessa, K. 2015. Ethnobotanical study of homegarden plants in
Sebeta-Awas District of the Oromia Region of Ethiopia to assess use, species diversity and
management practices J Ethnobiol Ethnomed 1164.
1746-4269-3-12.
10.1186/s13002-016-0131-x.
livestock industry with focus on Acari Ticks. Evid Based Complement Alternat Med. Article
ID 8647919, 33 pages. 2017; https://doi.org/10.1155/2017/8647919.
2008;26:132–53.
Sonchus asper (L.) Hill
Sonchus oleraceus L.
ASTERACEAE

Grace N. Njoroge
Synonyms
Sonchus asper (L.) Hill: Picrosia australis Decne; Picrosia longifolia D. Don.;
Sonchus asper (L.) Vill.; Sonchus carolinianus Walter; Sonchus gigas subsp. medius
Boulos; Sonchus oleraceus var. asper L.; Sonchus spinosus Lam.; Tragopogon
fritillarioides Less.
Sonchus oleraceus L.: Sonchus ciliatus Lam.; Sonchus gracilis Phil.; Sonchus
mairei H. Lév.; Sonchus maritimus Sessé & Moc.; Sonchus plumieri Sessé &
Mac.; Sonchus rivularis Phil.
Local Names
Sonchus asper/Sonchus oleraceus: Kikuyu: Mahiu; Luo: Achak; Shambaa:

Kwake; Hehe: Sungasunga (Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Sonchus asper L.: Annual. Root slender, narrowly conical or fusiform, usually
branched. Stem (10)30–75(100) cm high, finely sulcate, fistular, simple or branched,
glabrous or covered with dark, stalked, glandular hairs above. Leaves somewhat stiff
or almost coriaceous, less often thin and soft, dark green or sometimes light bluish,
oblong or obovate, 6–15 cm long and 1.5–8 cm wide, undivided, sinuate-pinnatifid,
or, less often, pinnatisect, with unevenly spinescent teeth; lower leaves on winged
petiole, middle and upper leaves sessile, with roundish auricles appressed to stem.
Capitula less numerous (usually 5–10), in corymbose or umbellate-corymbose
inflorescence; peduncles glabrous or glandular hairy, often much longer than usually
undeveloped capitula; fully developed capitula with 100–150 florets. Involucre
campanulate or bowl-shaped, 10–12(15) mm long, (6)10–15(25) mm wide, bracts
dark green, narrowly ovate, lanceolate or oblong, usually three-rowed. Corolla
yellow. Achenes oblong-obovoid, flat, light- or dark-brown, 2.5–3(3.5) mm long
and 0.75–1 mm wide, not rugose transversely (or weakly rugose), usually winged,
weakly toothed (under high magnification!) along margin, mostly with three prom-
inent longitudinal ribs on each side; pappus of numerous thin and soft, white, almost
smooth, weakly toothed (under high magnification), fragile hairs, two times or
slightly more as long as achenes. Flowering June–September. Ural, Caucasus,
Altai, Middle Asia, as weed in gardens, vineyards, along roads, in thickets of
steppes, river valleys, up to the middle mountain belt (Boborov and Tzvelev 1964)
(Figs. 1, 2, and 3).
Sonchus oleraceus L.: Annual (sometimes biennial). Root narrowly conical or
fusiform. Stem 30–100 cm high, finely sulcate, fistular, simple or branched, glabrous
or glandular-hairy above. Leaves not stiff, dull, 10–18(22) cm long and 5–7(12) cm
wide, pinnatipartite or lyrate with large terminal, triangular lobe, finely sinuate-
toothed, with somewhat spinescent teeth; lower leaves on winged petiole with
wide amplexicaul base; upper leaves sessile, with sagittate auricles at base. Capitula
mostly few (usually 5–15), in corymbose or umbellate-corymbose inflorescence;
peduncles and bases of capitula arachnoid- or flocculose-lanate when young, some-
times glandular hairy; some capitula often undeveloped, normally with 100–125
(150) florets. Involucre campanulate or bowl-shaped, 10–12 mm long and (6)10–15
(25) mm wide; bracts dark green, usually three-rowed, sometimes dorsally with few
glandular hairs. Corolla yellow, less often whitish. Achenes oblong-obovoid, some-
what narrowed toward base, more or less strongly compressed, light brown, brown,
or dark brown, 2.5–3, mm long, 1 mm wide, weakly toothed along margin (under
high magnification!), with three or less often more prominent longitudinal ribs on
each side, finely rugose transversely; pappus of numerous thin, soft, white, almost
smooth bristles (slightly barbed under high magnification), two times as long as
achene. Flowering June–October. Ural, Caucasus, Altai, Middle Asia, as weed in
field, kitchen gardens, irrigated areas, along roads, along river banks, to the middle
mountain belt (Boborov and Tzvelev 1964) (Figs. 4, 5, and 6).
Sonchus asper (L.) Hill. . . 977
Fig. 1 Sonchus asper

(Asteraceae), garden, Chicani,
Bolivia. (Photo
R.W. Bussmann and

Bolivia. (Photo
R.W. Bussmann and

Bolivia. (Photo
R.W. Bussmann and
Fig. 4 Sonchus oleraceus

Bolivia. (Photo
R.W. Bussmann and
Phytochemistry
Latex, aliphatic alcohols (taraxasterol, lupeol, n-nonocyanate, n-rentriacontane,

n-tri-triacontane), triterpenoids (taraksasterol), steroids (sitosterol, stigmasterol,
campesterol), sequiterpenoids (uzanIn C, macrocliniside A, crepidiazide A,

Bolivia. (Photo
R.W. Bussmann and
picriside B, picriside C), flavonoids (apigenine, luteoline, quercetine, quercimeritrin,

cinaroside, isocinaroside luteoline), vitamins (C), coumarins (scopoletine,
cichorianine), fatty acids (Sokolov 1993).
Sonchus asper: Used for wound healing (Muhammad et al. 2019) and diabetes
(Ullah et al. 2019).
Sonchus oleraceus: The roots are used as purgative. The crushed leaves soaked in
water are used for earache (Kokwaro 2009). Used medicinally in Ethiopia (Bekalo
et al. 2009). Very frequently used medicinally in the Americas, e.g., for cholera and
high blood pressure (Bussmann and Sharon 2006). Used to treat chest pain and
cough (Njoroge and Bussmann 2006b). Also to remedy gonorrhea and other sexu-
ally transmitted disease (Njoroge and Bussmann 2009), and sometimes for typhoid
(Njoroge et al. 2004).
Used ceremonially to remedy embarrassment, shame, hangover, negative energy,
anger, and to calm strong character (Bussmann and Sharon 2006; Paniagua-
Zambrana et al. 2020).
Sonchus arvensis is used in India for cholera, cuts, wounds, dysentery, and jaundice
(Verma et al. 2007).

Bolivia. (Photo
R.W. Bussmann and
Sonchus asper: Used in fodder as digestive (Ali et al. 2019).

Sonchus oleraceus: The leaf extract is used for snakebites in livestock and for
nervous system problems (Kokwaro 2009). Used as fodder additive to serve as
galactagogue (Ali et al. 2019). Used ceremonially to remedy embarrassment,
shame, hangover, negative energy, anger, and to calm strong character (Bussmann
and Sharon 2006; Paniagua-Zambrana et al. 2020). Used against anaplasmosis in
cattle (Njoroge and Bussmann 2006a).
Sonchus afromontanus and Sonchus bipontini are sometimes browsed by livestock
(Bussmann 2006; Bussmann et al. 2011). Sonchus bipontini is also used for scabies
and blackleg (Yineger et al. 2007).
References
Sher H, Bussmann RW, Shao J-W. Traditional Uses of plants by Indigenous Communities for
Veterinary Practices at Kurram District; 2019.

doi.org/10.1186/1746-4269-5-26.
Boborov EG, Tzvelev NN. (English 2001). Flora of the USSR, Volume 29: Compositae; Akademia
Nauk, Leningrad; 1964. 832 pages, 34 plates
Sikharulidze S, Tchelidze D. Sonchus asper (L.) hill; Sonchus oleraceus L. In: Batsatsashvili K,
Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions far Eastern Europe.
Cham: Springer International Publishing; 2020. https://doi.org/10.1007/978-3-319-77088-
8_129-2.
of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006a;108:332–9.
2009;8(2):262–9.
Paniagua-Zambrana NY, Bussmann RW, Romero C. Sonchus asper (L.) hill; Sonchus oleraceus
L. In: Paniagua-Zambrana NY, Bussmann RW, editors. Ethnobotany of mountain regions –
Ethnobotany of the Andes. Cham: Springer International Publishing; 2020. https://doi.org/
10.1007/978-3-319-77093-2_272-1.
Sokolov PD, editor. Plant resources of the USSR: flowering plants, their chemical composition, use;
Volume 7. Family Asteraceae (Compositae). Leningrad: Akademia Nauk; 1993, 352p. (in
Russian).
the southern and tribal regions of Khyber Pakhtunkhwa province, Pakistan. Ethnobot Res Appl.
2019;188 https://doi.org/10.32859/era.18.8.1-20.
of Banares Hindu university, Varanasi, Uttar Pradesh. J Ethnobiol Ethnomed. 2007;3:35.
Spilanthes mauritiana (Rich. ex Pers.) DC.
ASTERACEAE

Grace N. Njoroge
Synonyms
Spilanthes mauritiana (Rich. ex Pers.) DC.: Acmella caulirhiza Delille; Spilanthes

acmella (L.) L.; Spilanthes africana DC.
Local Names
Spilanthes mauritiana: Kikuyu: Gathara-Ita (Gachati 1989).
Botany and Ecology
Spilanthes mauritiana (Rich. ex Pers.) DC.: Annual creeping or trailing herb,

upper parts sometimes erect, up to 30 cm. Leaves opposite, ovate, up to 5 3 cm,
3-veined from the base, sparsely hairy on both surfaces particularly on the veins
R. W. Bussmann (*)
G. N. Njoroge

when young; margin serrate-crenate to almost entire. Capitula solitary, axillary and
terminal on long peduncles, deep yellow. Ray florets usually present, 3–4 mm long,
3-toothed at the apex, yellow or rarely white. Weed in lawns and gardens in high
rainfall areas; elsewhere in wet habitats such as by streams and rivers or along
roadsides. Widespread in Tropical Africa, South Africa, Swaziland, and Mauritius.
Spilanthes mauritiana: Used to remedy toothache and stomach problems (Njoroge

et al. 2004).
In Peru, the species is also used for toothache, anesthetic for pain, and for skin
rashes (Bussmann and Sharon 2006). Crushed leaf paste is applied to injuries
(Teklehaymanot et al. 2007). Spilanthes paniculata is used for toothache and
boils (Debbarma et al. 2017).
References
source of traditional medicines in Central Kenya: Optimizing resource efficiency R.U.E. in
agro-ecosystems. Lyonia. 2004;72:71–87.
Stephania abyssinica (Quart.-Dill. & A. Rich.)
Walp.
MENISPERMACEAE

Grace N. Njoroge
Local Names
Stephania abyssinica: Kikuyu: Muriira, Muguu; Shambaa: Futeute; Luhya:

Ibokho; Taita: Lufiafia; Kipsigis: Tabarariet, Labororiet; Maa: Oloiyobo-Igeek,
Ndabararia; Samburu: Lokitimu; Tiriki: Lihoru; Chagga: Mdekafuko (Beentje
Botany and Ecology
Stephania abyssinica (Quart.-Dill. & A. Rich.) Walp.: Twining liana, woody at the
base; stem covered with a thin bark; branchlets glabrous or more or less densely
pubescent to tomentose when young. Leaf-lamina 5–20 4–13 cm, ovate to broadly
ovate, rarely suborbicular, rounded at the base, obtuse or subacute at the apex, mem-
branous or papery, slightly discolorous, glabrous or tomentellous, basal nerves 8–10,
palmate, petiole 4–12 cm long. Male inflorescences of false compound umbels, axillary,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Stephania abyssinica

R.W. Bussmann)
Fig. 2 Stephania abyssinica (Menispermaceae), Mt. Kenya National Park, Kenya. (Photo
R.W. Bussmann)
solitary or clustered 2–4 together; axes glabrous or tomentellous; peduncle 4–10 cm

long with 3–6 rays ending in umbel-like cymes; involucre of 3–5 caducous bracts. Male
flowers with 6–8 obovate or subobovate sepals 1.2–2.5 0.6–1.2 mm, purplish, their
bases often violet; petals 3–4, 0.8–1.2 mm long, broadly ovate or suborbicular; syn-
andrium 6–8-locular. Female inflorescences similar to the male. Female flowers with
3–4 sepals; carpel glabrous. Drupe subspherical-flattened, 0.5–0.8 cm in diameter,
glabrous; endocarp with small prickles or thick tubercles arranged in three lines; condyle
not perforated. Seeds up to 0.8 cm long (Troupin 1956). Very common especially in
drier Afromontane forests (Bekalo et al. 2009; Bussmann 2002, 2006a; Bytebier and
Bussmann 2000) (Figs. 1, 2, 3, and 4).
Stephania abyssinica (Quart.-ill. & A. Rich.) Walp. 987

R.W. Bussmann)

(Menispermaceae), flowers,
Kenya. (Photo
R.W. Bussmann)
Leaves are crushed and applied to wounds, especially tortoise bites. The roots serve
as aphrodisiac. The plant powder is rubbed into small cuts on aching body parts
(Kokwaro 2009). Used to treat headaches and fever (Singh et al. 2019). Root
decoction is used to remedy gonorrhea (Yineger et al. 2008). The leaf decoction is
used to treat rabies (Giday et al. 2010), to treat stomachache, and expel a retained
placenta after birth (Giday et al. 2009). Root powder is applied to wounds, and the
root tonic is used against impotence (Giday et al. 2007).
The extract of the whole plant is used for mastitis in cattle (Kokwaro 2009). The
stem fibers are used for baskets (Beentje 1994). Eaten by livestock (Bussmann
2006b). The stems are used to make ropes to tie house posts, tools, and to make
milk containers (Bussmann et al. 2011). Various uses in ethnoveterinary medicine
(Teklehaymanot et al. 2007). The leaf decoction is used to treat rabies (Giday et al.
2010). Stephania dinklagei is used as fish poison (Neuwinger 2004).
References
doi.org/10.1186/1746-4269-5-26.
Bussmann RW. Vegetation zonation and nomenclature of African Mountains – An overview.
Lyonia. 2006a;111:41–66.
syntaxonomical survey. J East Afr Nat Hist. 2000;891-2:45–71.
Kirtinagar Block of Tehri Garhwal in Western Himalaya. Ethnobot Res Appl. 2019;1814 https://
doi.org/10.32859/era.18.14.1-11.
Troupin G. Menispermaceae. In: Turrill WB, Milne-Redhead E, editors. Flora of Tropical East
Africa. 32 pp. London: Crown Agents for Oversea Governments and Administrations; 1956.
ISBN 978-1-84246-286-7 Online Reference page.
2008;26:132–53.
Stoebe kilimandscharica O. Hoffm
ASTERACEAE

Grace N. Njoroge
Local Names
Stoebe kilimadscharica: Kikuyu: Muhato, Muhatu, Muthithinda; Marakwet:

Chepsikara; Maa: Ol-kibejus; Sebei: Sesindot, Sabuldet (Beentje 1994; Gachati
1989; Kokwaro 2009).
Botany and Ecology
Stoebe kilimandscharica O. Hoffm.: Shrub, 1.2–6 m high, much branched; bark gray,
longitudinally fissured; branches tomentose when young. Capitula 3–5 mm long,
sessile, solitary or in few-headed fascicles and axillary in leaf-axils in the distal part of
the branches, appearing as large compound leafy spikes; phyllaries gray-green, the inner
yellowish, oblong, 1–4 mm long, obtuse, tomentellous. Florets golden brown or yellow,
one per capitulum, tube cylindrical, 2.3–2.7 mm long, lobes about 0.2 mm long, anthers
1.7–1.8 mm long, style 2.7–3 mm long. Achenes cylindric, 0.6–1.2 mm long, glandular-
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Stoebe
kilimandscharica
(Asteraceae), Mt. Kenya
National Park, Sirimon
moorlands, Kenya. (Photo
R.W. Bussmann)
puberulous; pappus 1.8–2.7 mm long. Leaves gray-green, fascicled and crowded on

abbreviated shoots, recurved, linear, 1–4.5 mm long, 0.5 mm wide (to 8 by 1 mm on
seedlings), margins involute, apex mucronate, tomentellous (Beentje 2000, 2002;
Bussmann 2006) (Fig. 1).
Stoebe kilimandscharica is used by a variety of local names, indicating the
exactitude of traditional nomenclature. However, no uses whatsoever have been
reported. This makes the species an excellent example for the fact that many species
of African mountain regions simply grow in habitats that traditionally were rarely
accessed, and thus no uses were discovered for such species.
References
Beentje H, editor. Flora of Tropical East Africa: Compositae (Part 1). Rotterdam: Balkema; 2000.
Beentje H, editor. Flora of Tropical East Africa: Compositae (Part 2). Rotterdam: Balkema; 2002.
Bussmann RW. Vegetation zonation and nomenclature of African Mountains - An overview.
Lyonia. 2006;111:41–66.
Strychnos henningsii Gilg.
Strychnos spinosa Lam.
Strychnos usambarensis Gilg.
LOGANIACEAE

Grace N. Njoroge
Synonyms
Strychnos henningsii Gilg.: Strychnos albersii Gilg & Busse; Strychnos barbata
Chiov.; Strychnos elliottii Gilg & Busse; Strychnos holstii fo. condensata Duvign.;
Strychnos holstii fo. laxiuscula Duvign; Strychnos holstii Gilg; Strychnos holstii var.
reticulata (Burtt Davy & Honoré) J. Duvign.; Strychnos innocua subsp. dysophylla
(Benth.) I. Verd.; Strychnos ligustroides Gossw. & Mendoça; Strychnos myricoides
S. Moore; Strychnos pauciflora Gilg; Strychnos procera Gilg & Busse; Strychnos
reticulata Burtt Davy & Honoré; Strychnos sennensis Baker.
Strychnos spinosa Lam.: Brehmia spinosa (Lam.) Harv. ex A. DC.; Strychnos
buettneri Gilg; Strychnos cardiophylla Gilg & Busse; Strychnos carvalhoi Gilg;
Strychnos cuneifolia Gilg & Busse; Strychnos djalonis A. Chev.; Strychnos
emarginata Baker; Strychnos euryphylla Gilg & Busse; Strychnos flacurtii Desv. &
Thouars; Strychnos gillettii De Wild.; Strychnos gracillima Gilg.; Strychnos
gracillima var. paucispinosa De Wild.; Strychnos harmsii Gilg & Busse; Strychnos
laxa Soer.; Strychnos leiosepala Gilg & Busse; Strychnos lokua A. Rich.; Strychnos
R. W. Bussmann (*)
G. N. Njoroge

madagascariensis Poir.; Strychnos megalocarpa Gilg & Busse; Strychnos mini-

ungansamba Gilg; Strychnos muehge Chiov.; Strychnos omphalocarpa Gilg &
Busse; Strychnos radiosperma Gilg & Busse; Strychnos rhombifolia Gilg & Busse;
Strychnos sansibarensis Gilg; Strychnos schweinurthii Gilg; Strychnos spinosa subsp.
lokua Bruce; Strychnos spinosa subsp. volkensii (Gilg.) E.A. Bruce; Strychnos sinosa
var. pubescens Baker; Strychnos tonga Gilg; Strychnos unguacha var. retusa Chiov.;
Strychnos volkensii Gilg; Strychnos vuntac Bjoer.
Strychnos usambarensis Gilg.: Strychnos cerasifera Gilg.
Local Names
Strychnos henningsii: Kikuyu/Kamba: Muteta; Pokot: Chapkamkam; Borana:

Kara, Karrah; Kipsigis: Mase; legutuet; Maa: Olduyesi; Meru: Muchambe;
Samburu: Chibulukwa; Somali: Herdesa; Tugen: Turubupwa, Turkukwa;
Turkana: Yopoliss (Beentje 1994; Gachati 1989; Kokwaro 2009); English: Red
bitterberry.
Strychnos usambaransis: Kikuyu: Mutikani; Kamba: Gitarongui (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: Blue bitterberry, Stipe-fruited strychnos,
Stipe-fruited monkey orange.
Botany and Ecology
Strychnos henningsii Gilg.: Shrub or small tree, 2–10(20) m high with spreading
rounded crown. Trunk 40–50 cm in diam. (more or less); bark pale grey or pale
brown, rough. Branches pale grey or pale brown, sometimes shallowly fissured, not
lenticellate; branchlets conspicuously sulcate when dry, glabrous. Leaves petiole
short, glabrous 1–3 mm long; lamina pale to dark green and shining above, less
shining and paler beneath, coriaceous, very variable in shape and size even in a
single branchlet, elliptic, oblong, narrowly elliptic, or ovate, (1.2)1·5–3(3·5) x as
long wide, (1.5)2–6(10) (0.6)1–3(6) cm, rounded to acuminate at the apex,
cuneate, rounded, or sometimes on main axis subcordate (and then comparatively
wider); glabrous on both sides; one pair of secondary veins from above the base
curved along the margin and one, rarely two, faint submarginal pairs; tertiary
venation reticulate, prominent on both sides, especially in thick leaves. Inflorescence
axillary and sometimes also terminal, much shorter than the leaves, 1 1.2 2 cm,
congested, few- or many-flowered. Peduncle often very short, sparsely pubescent to
glabrous like the branches and pedicels. Bracts small, upper sepal-like, lower larger,
sparsely pubescent to glabrous beneath, often with colleters in the axils. Flowers
fragrant, 5-merous, sessile or subsessile. Sepals pale green, connate at the base,
broadly orbicular or nearly so, 1–1.4 1–1.4 mm, rounded or obtuse at the apex,
minutely ciliate, glabrous on both sides, without colleters. Corolla in the mature bud
2.3–3 x as long as the calyx, 2.8–4 mm long, greenish-yellow, creamy, or white,
subrotate and 4–5 mm in diam. When open, glabrous outside, inside pilose or villose
Strychnos henningsii Gilg. . . . 993
at the base of the lobes or sometimes entirely glabrous; tube short, 0.7–1.2 x as long
as the calyx, 0.8–1.5 mm wide; lobes thick, triangular to ovate, 1.7–2.5 x as long as
the tube, 1.7–2 x as long as wide, 2–2.5 1–1.6 mm, acute, spreading. Stamens just
exserted; inserted at the mouth of the corolla tube; anthers elliptic, 0.8–
1 0.5–0.8 mm, glabrous. Pistil glabrous, 1.6–2.2(3) mm long; ovary globose,
depressed-globose, or sometimes ovoid, 0.8–1.2(1·5) 0.9–1.4 mm, rounded or
sometimes acuminate at the apex, 2-celled; style 0.7–1 mm long; stigma capitate. In
each cell 8–12 ovules. Fruit yellow, orange, or red, ellipsoid, 1 0.8–2 1.5 cm,
one-seeded. Wall thin. Seed pale brown, ellipsoid, not flattened,
0.8 0.5 0.5–1.2 0.7 0.7 cm, glabrous, smooth, with a deep closed groove
at one side (like coffee-bean), very minutely fovelate. Occurs from DR Congo east to
Ethiopia and south to Angola and South Africa. It also occurs in Madagascar (Bruce
and Lewis 1960). A characteristic component of drier Afromontane forests, in
particular together with Coffea arabica and Croton sp. (Bussmann 2002).
Strychnos spinosa Lam.: Shrub or small tree with spreading habit, up to 6.5 m high;
bark sometimes reticulate and irregularly corky but never thick; young branchlets
varying from pale to dark mottled or concolorous, glabrous to shortly pubescent,
sometimes longitudinally ridged and often powdery corky; spines slender to stout,
straight or curved, sometimes absent. Leaves shortly petiolate; lamina membranous to
coriaceous, ovate, obovate or elliptic, 3–8 cm long, 1.3–7 cm wide, rounded, subacute or
more rarely acute or emarginate and often apiculate at the apex, cuneate or more rarely
rounded at the base, matt or shiny, 3–7-nerved at or just above the base, with hair-
pockets sometimes visible in the angles, indumentum of lamina varying on both surfaces
from pubescent to glabrous. Cymes terminal; peduncle and pedicels spreading-
pubescent. Calyx-lobes narrowly deltoid to linear, 3.5–5.5 mm long, shorter or longer
than the corolla, acuminate, usually glabrous except for the base, very rarely pubescent
up to the lobes. Corolla greenish-white; lobes deltoid, subacute, about half the length of
the corolla-tube, with dense hairs forming a ring at the base. Anthers densely bearded at
the base. Ovary broadly ovoid, unilocular; ovules numerous. Fruit globose, 5–12 cm in
diameter; rind woody, yellow; seeds numerous, flat and disk-like, 1.5–2 cm long,
embedded in fleshy yellow edible pulp. (Bruce and Lewis 1960).
Strychnos usambarensis Gilg.: Large climber up to 70 m long, climbing with solitary
tendrils, or shrub to small tree up to 10( 15) m tall; stem up to 25 cm in diameter; bark
pale or dark grey or grey-brown with darker patches, smooth, inner bark orange;
branches with lenticels, usually very dark brown, often covered with a pale skin
which splits and peels off, branchlets pale brown, glabrous or short-hairy. Leaves
opposite, simple and entire; stipules absent; petiole 2–6 mm long, glabrous; blade
ovate to elliptical, 3–8( 16) cm 1–3.5( 7) cm, base cuneate to rounded, apex
acuminate, 3–5-veined from the base. Inflorescence an axillary lax or congested thyrse,
solitary or several together, 1–2.5 cm long, few-flowered. Flowers bisexual, regular,
4–5-merous; sepals fused at base, ovate to triangular, up to 1 mm long; corolla tube up to
1.5 mm long, lobes oblong to ovate or triangular, c. 1 mm long, acute, recurved from
somewhat below the middle, glabrous or minutely hairy outside, inside with a ring of
hairs in the throat, white, yellow, or sometimes orange; stamens inserted at the mouth of
the corolla tube, exserted; ovary superior, ovoid, 0.5–1 mm long, glabrous, 2-celled,
style up to 1.5 mm long, stigma small, head-shaped or sometimes obscurely 2-lobed.

Fruit a globose berry 1–2 cm in diameter, often laterally compressed, soft, orange or
orange-yellow when ripe, often glaucous, 1-seeded. Seed depressed-globose or ellip-
soid, 9–12 mm 7–11 mm 5–8 mm, short and densely hairy, smooth. Occurs from
Guinea east to Nigeria, and from Congo east to Kenya and south to South Africa (Bruce
and Lewis 1960).
Strychnos henningsii: The Maasai and Batemi peoples in Tanzania add the plant to
milk and meat-based soups as a general tonic. In Kenya, a decoction of the branches
in soup is taken to treat rheumatism and gynecological complaints; a root decoction
is taken to treat chest pain and internal injuries. A stem decoction is taken to treat
malaria. The root decoction is used to treat chest pain and internal injuries. Fresh
roots are chewed for snake bites. Root and leaf decoctions serve for malaria and
rheumatism, normally given with honey. The bark serves for back ache (Kokwaro
2009). The bark decoction also treats rheumatism and arthritis (Beentje 1994) In
Kenya and Tanzania, fresh roots are chewed to treat snakebites. In Tanzania pounded
roots are taken in food to treat hookworm infections. In South Africa, a bark extract
is taken in small doses to treat colic, stomach-ache, dizziness, and as a purgative and
anthelminthic. The powdered bark in water is taken to treat nausea. A bark decoction
is given to children to treat internal worms and it is given to cattle to treat diarrhea
and heartwater caused by the rickettsial parasite Cowdria ruminantium. In Mada-
gascar the stem bark and roots are used to poison rats and mice, and sometimes also
for criminal purposes. Used as antimalarial (Njoroge and Bussmann 2006). Used for
back pain in women and postpartum pain (Njoroge and Bussmann 2009). Also for
diarrhea, general postpartum care, and in pregnancy (Randrianarivony et al. 2016,
2017). In Kenya, used for arthritis and stomachache (Muthee et al. 2011). Com-
monly traded in local markets (Delbanco et al. 2017).
Strychnos henningsii: In Kenya, the wood is used to make arrow shafts and poles
for building huts and cattle enclosures (Beentje 1994). Used as fish poison
(Neuwinger 2004).
Strychnos usambarensis: The Banyambo of Rwanda and Tanzania use the root bark
and leaves to produce arrow poison, sometimes in combination with other plants. In
DR Congo, the Nduye people mix the powdered root with water and apply this to the
nostrils of hunting dogs to improve their scent. In DR Congo and western Kenya, the
wood is used for house construction (Beentje 1994). Used as fish poison (Neuwinger
2004).
Strychnos henningsii Gilg. . . . 995
References
Bruce EA, Lewis J. Flora of Tropical East Africa: Loganiaceae. London: Crown Agents for
Overseas Governments & Administrations; 1960. 47 pp. ISBN 978-1-84246-282-9
Delbanco AS, Burgess ND, Cuni-Sanchez A. Medicinal plant trade in Northern Kenya: Economic
importance, uses and origin. Econ Bot. 2017;71(1):12–31.
anthelmintic and other medicinal plants traditionally used in Loitoktok district of Kenya. J
Ethnopharmacol. 2011;135:15–21.
Njoroge GN, Bussmann RW. Phytotherapeutic management of Diversity and utilization of antima-
Ethnomed. 2006;2:8.
2009;82:262–9.
Madagascar. Indian J Tradit Knowl. 2016;15(1):68–78.
2017. https://doi.org/10.1186/s13002-017-0147-x.
Syzygium cordatum Hochst.
Syzygium guineense (Willd.) DC.
MYRTACEAE

Grace N. Njoroge
Synonyms
Syzygium guineense (Willd.) DC.: Calyptranthes guineense Willd.; Eugenia

guineensis (Willd.) Baill. ex Laness. Syzygium fleuryi A. Chev.
Local Names
Syzygium cordatum: Kikuyu: Muriru, Mukui; Digo: Muziahi; Kipsigis: Sumoiyot,

Lemuguyet; Lumbwa: Lemejwet; Luo: Mukutan-Achak; Maa: Ol-Olobioroni;
Swahili: Mzambarau-ziwa, Myamayu, mlati; Nandi: Lemaiwet; Samburu:
Lairakai, Ngilenyai; Marakwet: Reberwa; Sebei: Lemeyuet (Beentje 1994; Gachati
1989; Kokwaro 2009); English: Water berry, Water wood, Water tree
Syzygium guineense: Kikuyu: Mukoe, Mukui; Swahili: Mshiwi, Mzuari,
Mzambarau, Mzambarau mwitu, Lubale, Mkongoro, Mlungiro; Digo: Muzaihi,
Mkulu; Kipsigis: Lemeiywet, Lemechwet; Maa: Ol-Ertagai; Marakwet/Tugen/
R. W. Bussmann (*)
G. N. Njoroge

Sebei: Lemaiyua; Meru: Muiru; Nandi: Lamayiowet; Samburu: Lamulii;

Ndorobo: Laikarai, Leperoi (Beentje 1994; Gachati 1989; Kokwaro 2009); English:
Water berry, Water pear, Snake bean tree.
Botany and Ecology
Syzygium cordatum Hochst.: Small or medium-sized evergreen tree up to 20 m tall;

foliage very dense, slightly glaucous. Bark dark brown, rough, flaking. Young stems
quadrangular and slightly winged. Leaf-lamina 4–8 2.2–13.5 7 cm, lanceolate-
elliptic to oblong-elliptic or suborbicular, apex broadly rounded to subacute or, very
rarely very shortly subacuminate, or emarginate, base deeply cordate and
amplexicaul; petiole up to 0.2 cm long. Receptacle (including pseudopedicel) and
calyx 0.6–0.9 cm long; calyx and upper receptacle 0.35–0.5 cm long; filaments (0.8)
1.1–1.5 cm long. Fruit c. 1.8 0.9 cm, purple-black, urceolate, persistent calyx +
upper receptacle 0.3–0.4 cm long 0.5 cm wide. Syzygium cordatum is distributed
from DR Congo eastward to Kenya and southward to South Africa (Verdcourt 2001).
A characteristic species in lower Afromontane forests dominated by Ocotea
usambarensis and Pouteria adolfi-friederici (Balemie and Kebebew 2006;
Bussmann 2002a, b, 2006).
Syzygium guineense (Willd.) DC.: Trees, shrubs, or pyrophytic subshrubs 0.2–30 m
tall with smooth or rough bark; sometimes buttressed; young branches terete or
somewhat 4-angled, mostly only strongly 4-angled in hybrids with Syzygium
cordatum. Leaves elliptic, oblong-elliptic, or elliptic-lanceolate to obovate-elliptic,
(3–)4–16 cm long, 1–7.7 cm wide, acute to acuminate to broadly rounded, sub-
truncate or even emarginate at the apex, cuneate at the base; petiole 0.6–2.5(–4.5) cm
long. Inflorescence short to extensive, 5–19 cm long. Calyx-tube with pseudo-
pedicels usually distinct. Filaments 3.5–9 mm long. Fruits variously red or purple,
subglobose or ellipsoid, 0.8–3.5 cm long, 0.6–2.5 cm wide. Very widespread
throughout tropical Africa from Senegal to Somalia and Arabia and south to S &
SW Africa (Verdcourt 2001). A characteristic species in lower Afromontane forests
dominated by Ocotea usambarensis and Pouteria adolfi-friederici (Balemie and
Kebebew 2006; Bussmann 2002a, b, 2006) (Figs. 1, 2, 3, 4, 5, and 6).
Syzygium cordatum: In traditional medicine, a root decoction is drunk against

amenorrhea. Ash of the burnt wood is rubbed on the forehead against headache.
Decoctions of the root bark and stem bark are taken for treatment of malaria. Root
and bark infusions are taken to treat cough; root and bark decoctions to treat
indigestion, abdominal pain, stomachache, and venereal diseases (Kokwaro 2009).
The bark is used as an emetic and to treat diarrhea, stomach problems, headache,
amenorrhea, wounds, and respiratory problems. A leaf extract is drunk against
cough, and an infusion of the leaves against diarrhea and stomach complaints, and
Syzygium cordatum Hochst. . . . 999
Fig. 1 Syzygium guineense

(Myrtaceae), flowering
specimen, Bale Mountains
Bussmann)

(Myrtaceae), bark cut, Bale
as a purgative. Ground leaves, bark, and roots steeped in water are applied as a
poultice as a galactagogue.
Syzygium guineense: With ample application in African traditional medicine, but
can be dangerous, as the bark may be poisonous, and death after its use has been
recorded. The root is soaked in water for drinking and bathing to treat epilepsy. A
root infusion is drunk for treatment of stomach-ache. Root extracts are taken as a
purgative, anthelmintic, and taeniacide, also for stomach-ache. Bark decoctions are
used against stomach-ache, diarrhea, and malaria; they are considered mildly laxa-
tive, and are applied in draught or in baths as a tonic. An infusion is taken against
coughs, asthma, throat problems, and intercostal pain. The powdered bark is used as
an antispasmodic, purgative, and anthelmintic, and used for treatment of diarrhea,
stomach-ache, broken bones, and wounds. The fruits are eaten for diarrhea

(Myrtaceae), flowering
branch, Bale Mountains
Bussmann)

(Myrtaceae), flowers, Bale
(Kokwaro 2009). In Cameroon, the bark is used for the treatment of snakebites. Twig
bark preparations are applied against paralysis. A decoction of twigs and leaves is
drunk or used as an enema for its purgative properties and against colic, diarrhea, and
abdominal pain. It is also used as drink or bath against insanity, amenorrhea, and


cerebral malaria. The crushed leaf is applied on wounds and boils. Leaf decoctions
are taken against intestinal parasites and stomach-ache, used as an enema against
diarrhea, and used as an embrocation to bathe and then massage into areas of sprain.
Leaf decoctions or pulverized leaves are given as tonic to pregnant women. The leaf
is chewed against stomach-ache. A liquid of chewed leaves mixed with water is used
as eye drops to treat ophthalmia. The fruit is used for treating dysentery. The bark
decoction is used for diarrhea (Bekalo et al. 2009). A leaf paste is applied to
swellings, and the powder ingested for diarrhea (Flatie et al. 2009).
Syzygium aromaticum is part of a mixture used in Indian medicine for tuberculosis,
smallpox, measles, skin diseases, rheumatism, cardiac problems, and as antifungal
Local Food Uses
Syzygium cordatum: The fruit is edible, but it has a rather bland taste. It is popular
with children. The fruit is also made into jellies and alcoholic drinks (Beentje 1994).
Sometimes an extract is used as fish poison (Neuwinger 2004).
Syzygium guineense: The fruit is edible. It has been described as sweet and juicy,
but also as having a rather bland taste and not being appreciated. It is sought after
especially by children, and in Ethiopia and Kenya, the fruit is sold on markets
(Bussmann et al. 2011). The fruit is made into a beverage, vinegar, and added to
spirits for flavoring. In Sudan, a meal is made from roasted and ground fruits.
Sometimes an extract is used as fish poison (Neuwinger 2004).
Syzygium cordatum: The wood is used for mortars, utensils, construction, beams,
rafters, poles, furniture, window frames, and beehives (Beentje 1994). Its durability
in water makes it especially suitable for boat building, and in South Africa, the logs
are traditionally used to make the jetties and slipways around the swamps in the Kosi
Bay area. The wood is also suitable for flooring, interior trim, joinery, toys, novelties,
turnery, railway sleepers, mine props, veneer, and plywood. It is popular as fuelwood
and used for charcoal making. Smoke from burning wood is used to season milk
containers. The flowers provide nectar for honeybees. The pulverized bark is
sprinkled on water as fish poison.
Syzygium guineense: The wood is used for construction, flooring, panelling, furni-
ture, utensils, tool handles, plates, stools, carvings, and poles (Beentje 1994). Its
flexibility makes it suitable for bows and ribs of canoes. The bole is made into
dugout canoes. In East Africa, the wood has been used for railway sleepers. It is also
suitable for vehicle bodies, interior trim, joinery, toys, novelties, boxes, crates, mine
props, veneer, plywood, hardboard, and particle board. It is good fuelwood and used
for charcoal making. The bark is used for tanning and dyeing. Bark extracts are
sometimes used to harden lateritic floors or to glaze pottery. The leaves and fruits are
used as fodder for livestock, and the flowers are a source of nectar for honeybees.
The tree is used as a shade tree in coffee cultivation in Ethiopia. The crushed leaf is
taken to treat insanity and possession. Used as firewood, for construction, and is an
excellent honey resource (Bussmann et al. 2011). The root is tied around the neck
against evil spirits (Giday et al. 2007).
References

doi.org/10.1186/1746-4269-5-26.
Bussmann RW. Vegetation zonation and nomenclature of African Mountains an overview. Lyonia.
2006;11(1):41–66.
zone, Benishangul-Gumuz regional state, mid-West Ethiopia. J Ethnobiol Ethnomed.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Verdcourt B. Flora of tropical East Africa: Myrtaceae. Rotterdam: Balkema; 2001.
Tabernaemontana stapfiana Brizten
APOCYNACEAE

Grace N. Njoroge
Local Names
Tabernaemontana stapfiana: Kikuyu: Mwelele, Mwerere; Kipsigis: Erendet,

Terendet; Kisii: Ombondo; Luhya: Kunandere; Meru: Muerere; Nandi: Mabondet;
Ndorobo: Derendet; Tugen: Cheboition.
Botany and Ecology
Tabernaemontana stapfiana Brizten: Tree 5–25(–35) m high; trunk 25–90 cm in

diameter; bark pale to dark grey brown, rough, thick, corky; wood yellow or light
brown. Fruits dark green, densely speckled yellow or white, subglobose, obliquely
ovoid and angular when young, 10–20 cm long, 8–20 cm in diameter, rounded,
dehiscent, several-many-seeded; pericarp 2.5–6 cm thick; aril orange; seeds dark
brown, slightly coffee-bean-like, obliquely ellipsoid, 15–21 mm long. Inflorescence
lax, 10–28 cm long, few- to many-flowered, glabrous in all parts; peduncle 3–15 cm
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Tabernaemontana
stapfiana (Apocynaceae),
Chogoria, Kenya (Photo
R.W. Bussmann)
long; pedicels 5–30 mm long. Flowers with sepals suborbicular to oblong, 5–7 mm
long, ciliate or not, with colleters within; corolla white, thick, fleshy; mature bud
with a comparatively large ovoid head conspicuously wider than the tube with a
subacute apex; tube pale green, almost cylindrical, throat pale yellow, 21–42 mm
long, not angular, twisted over the entire length of the tube; lobes spreading, later
recurved, obliquely elliptic, sometimes falcate, 17–60 mm long, 8–35 mm wide,
rounded, obscurely auriculate at the left side of the base; stamens included for
2–15 mm, inserted 11–15 mm from the base; pistil glabrous, 13–17 mm long;
style 5–10 mm long; ovary almost cylindrical; ovules 100–200 in each carpel.
Leaves petiolate; blade narrowly elliptic to slightly obovate, 12–42 cm long,
3–17 cm wide, acuminate, apiculate or rounded at the apex, cuneate or decurrent
at the base, with scattered dots beneath; petiole 30 mm long (Omino 2002; Goyder
et al. 2012). Characteristic species in Ocotea usambarensis and Pouteria adolfi-
friederici dominated wet Afromontane forests (Bussmann 2002a, b, 2006) (Fig. 1).
Tabernaemontana stapfiana: The root and bark decoction is used to treat pneumo-
nia, chest pains, and speeds delivery (Jeruto et al. 2008). The smoke of
Tabernaemontana elegans is used to treat headaches (Mohagheghzadeh and Faridi
2006). In Madagascar, Tabernaemontana ciliata is used got stomach pain, malaria,
abdominal pain, and intestinal parasites (Rabearivony et al. 2015).
Tabernaemontana coffeoides serves for similar purposes (Rakotoarivelo et al.
2015). The flower sap of Tabernaemontana divaricata serves to treat conjunctivitis
(Raj et al. 2018).
Tabernaemontana stapfiana Brizten 1007
Tabernaemontana pachysiphon is used as fish and arrow poison (Neuwinger 2004).
References
2006;11(1):41–66.
Goyder d, Harris T, Masinde S, Meve U, Venter J. Flora of tropical East Africa: Apocynaceae (Part
E. Africa, Apocyn. (Part 2)]
Omino EA. Flora of tropical East Africa: apocynaceae (Part 1). Rotterdam: Balkema; 2002. 114 pp.
ISBN 90-5809-409-X
018-0208-9.
Tagetes minuta L.
ASTERACEAE

Grace N. Njoroge
Synonyms
Tagetes minuta L.: Tagetes bonariensis Pers.; Tagetes glandulifera Schrank;

Tagetes glandulosa Link; Tagetes porophyllum Vell.; Tagetes riojana M. Ferraro
Local Names
Tagetes minuta: Kikuyu: Mubangi; Luhya: Inasbutsaka (Beentje 1994; Gachati

1989; Kokwaro 2009).
Botany and Ecology
Tagetes minuta L.: Annual herbs, 10–250 cm, plant aromatic. Stems much branched
in larger plants and almost woody, ribbed, glabrous, glandular. Leaves mostly
opposite, often alternate in upper part, dark green, pinnatisect, elliptic in outline,
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Tagetes minuta

(Asteraceae), Chicani,
Bolivia. (Photo
R.W. Bussmann and
3–30 0.7–8 cm, rachis narrowly winged, lobes up to 17, linear-oblong, to

11 1 cm, with orange glands. Synflorescence a dense terminal corymb. Capitula
narrowly cylindric; involucre 8–12 mm; phyllaries 3 or 4, yellow-green, fused,
glabrous, with brown or orange linear glands. Ray florets 2 or 3, pale yellow to
cream, lamina 2–3.5 mm. Disk florets 4–7, yellow to dark yellow, 4–5 mm. Achenes
black, narrowly ellipsoid, 6–7 mm, pilose; pappus of 1 or 2 setae to 3 mm and 3 or
4 scales to 1 mm, apices ciliate (Wu et al. 1994–2013) (Figs. 1 and 2).
Tagetes minuta: Used in Kenya to treat asthma (Njoroge et al. 2004; Njoroge and
Bussmann 2006b), wounds (Njoroge and Bussmann 2006c), to increase fertility
(Njoroge and Bussmann 2009). Also employed for wound healing and ulcers (Jeruto
et al. 2008), and other medicinal purposes (Yineger et al. 2008). Sometimes burnt
and the smoke inhaled to treat snakebites and mood disorders (Mohagheghzadeh and
Faridi 2006).
Taretes erecta: used in Peru used for respiratory problems, bronchitis, colds, and
congestion (Bussmann and Sharon 2006). Tagetes filifolia: used in Peru for severe
colic, stomach, stomach pain, diarrhea (Bussmann and Sharon 2006). Similar uses
are reported from India (Raj et al. 2018).
Tagetes minuta L. 1011
Fig. 2 Tagetes minuta

(Asteraceae), Chicani,
Bolivia. (Photo
R.W. Bussmann and
Leaf decoction used as insecticide (Jeruto et al. 2008; Kokwaro 2009; Wanzala
2017), and to treat cattle diseases (Njoroge and Bussmann 2006a). Tagetes patula is
plated as ornamental in Ethiopia (Mekonnen et al. 2015).
References
of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006a;108:332–9.
Central Kenya. J Ethnopharmacol. 2006c;111:303–7.
and reproductive health conditions in Central Province, Kenya. Indian J Tradit Knowl. 2009;8
(2):262–9.
018-0208-9.
Wanzala W. Potential of traditional knowledge of plants in the Management of Arthropods in
livestock industry with focus on Acari ticks. Evid Based Complement Alternat Med. 2017:
Article ID 8647919, 33 pages. https://doi.org/10.1155/2017/8647919.
2008;26:132–53.
Tamarindus indica L.
FABACEAE

Grace N. Njoroge
Synonyms
Tamarindus indica L.: Cavaraea elegans Speg., Tamarindus erythraeus Mattei,

Tamarindus occidentalis Gaertn., Tamarindus officinalis Hook., Tamarindus
somaliensis Mattei, Tamarindus umbrosa Salisb.
Local Names
Tamarindus indica: Swahili/Digo/Taita: Mkwaju; Bajun: Ukwayu; Borana:

Groha; Boni: Mukai; Ilwana: Morhoqa; Kamba: Kithumula; Luo: Chwaa,
Ochwa; Maa: Oloisiojoi; Marakwet: Aron; Meru: Muthithi; Oromo: Rhoka;
Pokot: Aron, Oron; Samburu: Rogei; Sanya: Roka; Somali: Hamar, Rakhai;
Taveta: Muzumura, Musumera; Tugen: Arwe; English: Tamarind (Beentje 1994;
Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Tamarindus indica L.: Tree 3–24 m. high, bark rough, grey or grey-black, crown
rounded, young branchlets pubescent or puberulous pairs, narrowly oblong, (0.8–)
1.2–3.2 cm long, 0.3–1.1 cm wide, rounded to rounded-subtruncate rarely slightly
emarginate at apex, rounded and asymmetric at base, glabrous except for a tuft of
yellowish hairs at base, sometimes pubescent up midrib and margins, rarely all
over both surfaces, venation reticulate-raised on both surfaces. Racemes 1–15(–
22) cm long, axis subglabrous to densely pubescent, pedicels 3–14 mm long,
glabrous to pubescent. Flower-buds red. Hypanthium 3–5 mm long. Sepals 8–
12 mm long, pale yellow inside, reddish outside. Large petals 10–13 mm long,
elliptic or obovate-elliptic, gold with red veins. Pods curved or sometimes straight,
sausage-like, (3–)6.5–14 cm long, 2–3 cm in diameter, usually obtuse at base and
apex, sometimes irregularly constricted, closely covered outside with brown scurf,
1–10-seeded. Seeds chestnut-brown, rhombic to trapeziform, 11–17 mm long,
10–12 mm wide. Woodland, wooded grassland, deciduous bushland, near sea-level
to 1520 m.
A decoction of leaves and twigs is used to treat diarrhea. The root decoction serves to
remedy fevers and cough. The bark decoction can be used as gargle for sore throat.
The leaf decoction is used for baths and internally in cases of measles and
chickenpox, also for stomach-ache. A decoction of roots and barks serves for
gonorrhoea, asthma, malaria, leprosy, liver problems, rheumatism, hookworm,
ulcers, and as laxative (Kokwaro 2009). The fruits are used to treat malaria and
diarrhea (Flatie et al. 2009). Very widely used all over Africa form abdominal pain,
diarrhea, as laxative, for wound healing, as anthelminthic, for malaria and fever
(Havinga et al. 2010). Dried fruits are ingested for eye infections (Muthu et al. 2006).
Sometimes used to treat cancer and diabetes, as well as as panacea (Delbanco et al.
2017).
In India used for anasarca, anthelmintic, boils, cough, digestion, eye com-
plaints, fever, headache, muscular pain, scorpion stings, stomach-ache, sun
stroke, swelling, syphilis, tooth ache, and urine complaints (Raj et al. 2018;
Verma et al. 2007). Also used for the treatment of hypertension (Malik et al.
2015). In Madagascar used to treat headache, flu, colds, fever, cough, measles,
dizziness during pregnancy and children’s epilepsy, back pain and fatigue
(Rabearivony et al. 2015; Randrianarivony et al. 2016a), and in pregnancy
(Randrianarivony et al. 2016b and for malaria (Randrianarivony et al. 2017). In
Peru used as laxative, for blood circulation, epilepsy, and heart disease
(Bussmann and Sharon 2006).
Tamarindus indica L. 1015
Local Food Uses
From the sausage-shaped fruits of the tamarind tree comes the sticky acidic pulp that
has been used as a food ingredient for thousands of years. The pulp was traded
widely in ancient times leading to the extensive use and cultivation. The edible fruits,
and especially the pulp, can be eaten raw or used as an ingredient in curries, pickles,
confectionery, and in fermented drinks. The seeds can be eaten raw or cooked
(Beentje 1994; Kokwaro 2009). The fruits are used as appetizer (Flatie et al.
2009). Fruits widely eaten in Madagascar (Randrianarivony et al. 2016a).
Tamarind wood is used as timber, firewood, and for charcoal. Used as dye. The
leaves are fed to livestock (Borana) (Beentje 1994; Kokwaro 2009). Extracts are
used as insect repellent (Wanzala 2017). Used for shade and ritual purposes (Balemie
and Kebebew 2006). Used to make beds, mortars and pestles, and for cultural
purposes in Madagascar (Randrianarivony et al. 2016a).
References
Delbanco AS, Burgess ND, Cuni-Sanchez A. Medicinal plant trade in northern Kenya: economic
zone, Benishangul-Gumuz regional state, mid-West Ethiopia. J Ethnobiol Ethnomed.
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Havinga RM, Hartl A, Putscher J, Prehsler S, Buchmann C, Vogl CR. Tamarindus indica
L. Fabaceae: patterns of use in traditional African medicine. J Ethnopharmacol.
2010;127:573–88.
018-0208-9.

vol. 1: William L. Brown Center, MBG, St. Louis; 2016a. ISBN 978-0-9960231-4-6.
Mahaboboka, Amboronabo, Mikoboka, southwestern Madagascar. J Ethnobiol Ethnomed.
2017; https://doi.org/10.1186/s13002-017-0147-x.
Wanzala W. Potential of traditional knowledge of plants in the Management of Arthropods in
livestock industry with focus on Acari ticks. Evid Based Complement Alternat Med. 2017:
Article ID 8647919, 33 pages. https://doi.org/10.1155/2017/8647919.
Tarchonanthus camphoratus L.
ASTERACEAE

Grace N. Njoroge
Synonyms
Tarchonanthus camphoratus L.: Tarchonanthus abyssinica Sch. Bip.,

Tarchonanthus camphoratus var. litakunensis Harv., Tarchonanthus minor Less.
Local Names
Tarchonanthus camphoratus: Kikuyu: Mururicua; Maa: Esentyio, Kileleshwa,

Ol-Leleshwa’ Kipsigis/Tugen: Lelechuet, Elewa Swahili: Mkalambati’ (Beentje
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Tarchonanthus camphoratus L.: Dense shrubs or small trees up to 8 m tall, with a

strong camphor odor. Trunk up to c. 40 cm in diameter, bark greyish, fissured, twigs
leafy, closely greyish- or pale brown-felted. Leaves subsessile or with a petiole up
to c. 6 mm. long and tomentellous, lamina very variable in size about 2 0.5 to 8.5
(12) 3.5(5) cm, narrowly oblong-elliptic, elliptic or oblanceolate to narrowly
obovate, apex subacute obtuse or rounded, base cuneate, margins entire, upper
surface glabrescent, finely tessellate-reticulate with numerous golden glandular-
globules along deeply depressed veins, under surface white-felted, the tomentum
about obscuring minute golden glands, midrib and nerves prominent beneath. Heads
numerous in large leafy terminal panicles. Female capitula, involucres
3–3.5 6 mm, broadly turbinate to subglobose, phyllaries 2–3-seriate, imbricate,
rotund to narrowly ovate, shortly connate at base or free, tomentose-araneose outside
and often long ciliate about the upper margins, florets 4–5, corollas 1 mm. long,
shortly funnel-shaped, lanate outside, stigma shortly bifid exserted, achene 2.5–3
(4) 1.5 mm, somewhat compressed ellipsoid, densely covered in a mass of long
sericeous-woolly hairs. Male capitula, involucres 2.5–3.5 3.5–5 mm, broadly
turbinate to shallowly cupuliform, phyllaries 5–6, ovate, connate below, sometimes
also with several free shorter narrowly oblong-ovate phyllaries, tomentose-araneose
outside, florets (3)10–25, corollas 2.5 mm long broadly funnel-shaped, densely
woolly outside, anther thecae 1.5–2 mm long, exserted, style linear exserted by
1.5 mm above the anther tube, ovary rudimentary. Male capitula, involucres
2.5–3.5 3.5–5 mm, broadly turbinate to shallowly cupuliform, phyllaries 5–6,
ovate, connate below, sometimes also with several free shorter narrowly oblong-
ovate phyllaries, tomentose-araneose outside, florets (3)10–25, corollas 2.5 mm long
broadly funnel-shaped, densely woolly outside, anther thecae 1.5–2 mm long,
exserted, style linear exserted by 1.5 mm above the anther tube, ovary rudimentary.
Female capitula, involucres 3–3.5 6 mm, broadly turbinate to subglobose, phyl-
laries 2–3-seriate, imbricate, rotund to narrowly ovate, shortly connate at base or
free, tomentose-araneose outside and often long ciliate about the upper margins,
florets 4–5, corollas 1 mm long, shortly funnel-shaped, lanate outside, stigma shortly
bifid exserted, achene 2.5–3(4) 1.5 mm, somewhat compressed ellipsoid, densely
covered in a mass of long sericeous-woolly hairs (Beentje 2000, 2002) (Figs. 1 and 2).
Inhalation of smoke for headache, fever, rheumatism. Asthma. A decoction of leaves

and twigs serves to remedy asthma, gastrointestinal problems, bronchitis, and is used
as mouthwash. Hot leaf poultices are used for asthma, bronchitis, and general
inflammation (Kokwaro 2009). The dry leaves in water are used against tapeworm
(Maundu et al. 2001). Used for abdominal disorders, diarrhea, as anthelminthic, for
asthma and bronchitis, as well as headache (Nankaya et al. 2019; Njoroge and
Bussmann 2006).
Tarchonanthus camphoratus L. 1019
Fig. 1 Tarchonanthus
camphoratus (Asteraceae).
Mt. Longonot caldera rim,
Kenya. (Photo R.W:
Bussmann)
Fig. 2 Tarchonanthus
camphoratus (Asteraceae).
Kenya. (Photo R.W:
Bussmann)
Local Food Uses
The leaves are used in soups for strengthening (Bussmann et al. 2006, 2018).
Used for bedding because it has a good smell and protects from bed bugs, also used
as perfume (Bussmann et al. 2006, 2018; Maundu et al. 2001), as fire-starter
(Bussmann et al. 2006), and firewood, as well as construction (Beentje 1994;
Maundu et al. 2001).
References
Beentje H. Flora of tropical East Africa: Compositae (Part 1). Rotterdam: Balkema; 2000.
Maundu P, Berger D, Ole Saitabau C, Nasieku J, Kipelian M, Mathenge S, Morimoto Y, Höft
R. Ethnobotany of the Loita Maasai people and plants working paper 8. Paris: Unesco; 2001.
Njoroge GN, Bussmann RW. Phytotherapeutic management of Diversity and utilization of antima-
larial ethnophytotherapeutic remedies among the; 2006
Teclea simplicifolia Engl.
Vepris dainelii (Pic. Serm.) Mziray
Vepris nobilis Delile.
RUTACEAE

Grace N. Njoroge
Synonyms
Teclea simplicifolia Engl.: Teclea salicifolia Engl.; Teclea simplicifolia (Engl.)

Engl.; Teclea viridis I. Verd.; Toddalia eugeniifolia Engl.
Vepris dainelii (Pic. Serm.) Mziray: Diphasia dainellii Pic.Serm.
Vepris nobilis Delile.: Aspidostigma acuminatum Hochst.; Cranzia nobilis (Delile)
Kuntze; Teclea diversifolia Lanza; Teclea nobilis Delile; Toddalia nobilis (Delile)
Hook. f. ex Oliv.
Local Names
Teclea simplicifolia: Kikuyu: Munderendu; Borana: Mike; Digo: Muchimi wa

tsakani; Ndorobo: Ikirai; Kipsigis / Ndorobo / Marakwert: Kuriot; Meru: Muretu;
Tugen: Kurionde; Turkana: Edapalakuyen; Maa: Oleglai; Samburu: Lgelai,
Ngolei orok, Nkilaiorok (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Vepris dainelii: Kikuyu: Munderendu (Beentje 1994; Gachati 1989; Kokwaro 2009).
Vepris nobilis: Kikuyu: Munderendu; Teso: Ekudep; Kipsigis / Ndorobo: Kuriot,
Kurisyot; Luhya: Mumutare, Mutaro, Muzo; Luganda: Nzo; Luo: Midat, Odar;
Maa: Ol-Gelai; Marakwet: Koorea, Lugumwa; Pokot: Kurion; Turkana: Ekodek
Botany and Ecology
Vepris dainellii: Endemic to southern Ethiopia, a small understorey tree up to 15 m tall

in moist montane forest. Differs from Vepris nobilis by its opposite leaves and 2-lobed
fruit. The tough wood is used for furniture and agricultural implements, and as
firewood. The fruit is edible (Kokwaro 1982). An important component of semi-
deciduous Afromontane forests in southern Ethiopia (Bussmann 1997, 2002a, b)
(Figs. 1, 2, 3, and 4).
Vepris nobilis: Evergreen shrub or tree (3–)5–12 m high or much taller in rain-forest;
bark smooth and grey; branchlets glabrous. Leaves trifoliolate, occasionally 2- or
1-foliolate; petiole 1.5–6( 8) cm long, terete or sometimes slightly grooved at the
apex, usually glabrous; leaflets subsessile or with a petiolule up to 10 mm long,
elliptic to oblong-elliptic, 5–15( 18) cm long, 1.5–4( 5.5) cm broad, acute to
acuminate at the apex, narrowly cuneate at the base, entire, glabrous except some-
times puberulous on the midrib; lateral nerves numerous. Inflorescence of terminal
and axillary panicles 4–15( 21) cm long, glabrous. Flowers polygamous. Bisexual
flowers with 4( 5) stamens. Female flower with 4 or 5 staminodes 0.5–1.2 mm long
(variable in length on the same flower); ovary subglobose, 1–1.4 mm in diameter,
glabrous, unilocular, 2-ovulate; style up to 0.5 mm long; stigma disk-shaped and
peltate, 1 mm. in diameter. Male flowers with 4( 5) stamens, 3–5.5 mm long
(variable in length on the same flower); anthers basifixed; rudimentary ovary slender
Fig. 1 Vepris dainellii

(Rutaceae), Bale Mountains
National Park, Demaro Forest,
Ethiopia. (Photo R:W.,
Bussmann)
Teclea simplicifolia Engl. . . . 1023

Bussmann)
Fig. 3 Vepris dainellii (Rutaceae), Bale Mountains National Park, Demaro Forest, Ethiopia. (Photo
R:W., Bussmann)
and glabrous. Sepals 4, united into a cupuliform calyx 0.6–0.8 mm long; lobes small,
ovate, ciliate. Petals 4( 5), narrowly elliptic, 3.5–4 mm long, 1.5–1.7 mm broad.
Male flowers with 4( 5) stamens, 3–5.5 mm long (variable in length on the same
flower); anthers basifixed; rudimentary ovary slender and glabrous. Female flower with
4 or 5 staminodes 0.5–1.2 mm long (variable in length on the same flower); ovary
subglobose, 1–1.4 mm in diameter, glabrous, unilocular, 2-ovulate; style up to 0.5 mm
long; stigma disk-shaped and peltate, 1 mm in diameter. Bisexual flowers with 4( 5)
stamens. Fruit obovoid, 6–8 mm long, 5–6 mm in diameter, red, glabrous, barely
foveolate, wrinkled when dry, 1-seeded. Seed ovoid, 5.5–6 mm long. Evergreen forest,

Bussmann)
Fig. 5 Vepris nobilis

Bussmann)

Bussmann)

Bussmann)
riverine forest and woodland; 900–2600 m. (Kokwaro 1982). An important component

of semideciduous Afromontane forests (Bussmann 2002a, b) (Figs. 5, 6, 7, and 8).
Vepris simplicifolia: A small 1-foliolate tree up to 10( 20) m tall, occurring in
Ethiopia, Kenya and Tanzania (Beentje 1994). An important component of semi-
deciduous Afromontane forests (Bussmann 2002a, b) (Figs. 9 and 10).
Vepris dainellii: The roots are crushed and applied topically to boils (Giday et al.
2010).

Bussmann)
Fig. 9 Vepris sp. (Rutaceae), Bale Mountains National Park, Demaro Forest, Ethiopia. (Photo R:
W., Bussmann)
Vepris nobilis: The leaves are used in a vapor bath to treat fever, and leaf and root
decoctions are drunk to treat pneumonia, rheumatism, and itching. The roots serve as
an anthelmintic and for treatment of pneumonia. Pounded root bark is applied to
syphilitic ulcers, whereas the stem bark is taken as an expectorant. Bark and leaves
are used as an analgesic. Roots and twigs are used as toothbrushes (Beentje 1994;
Gachati 1989; Kokwaro 2009). A decoction of roots or leaves with honey is used for
pneumonia, rheumatism, arthritis, measles, and externally for scabies (Kokwaro
2009). Used as toothbrush (Bussmann et al. 2006). Used to treat malaria and to
make strengthening soups (Bussmann 2006). Used also as febrifuge
(Mohagheghzadeh and Faridi 2006). Also used for colds (Gafna et al. 2017).
Fig. 10 Vepris
sp. (Rutaceae), Bale
Demaro Forest, Ethiopia.
(Photo R:W., Bussmann)
Vepris simplicifolia: In traditional medicine, a bark decoction is drunk to treat chest

complaints, malaria, and hepatitis, and a root decoction to treat stomach-ache,
backache, leprosy, gonorrhea, and brucellosis. Leaves and twigs are used for treating
pleurisy. A leaf decoction is taken against pneumonia, leaf ash is applied externally
against leprosy, and fruits are chewed to relieve toothache. Twigs are used as
toothbrushes (Beentje 1994; Gachati 1989; Kokwaro 2009). Used to treat malaria
and to make strengthening soups, as well as toothbrush (Bussmann 2006). To treat
pneumonia, allergies and gonorrhea (Muthee et al. 2011), for stomachache, diarrhea
and hepatitis, back ache, joint pains, and general health (Nankaya et al. 2019, 2020).
Vepris ampody is applied to wounds and used for abdominal pain (Rabearivony et al.
2015). Vepris boviniana is used for epilepsy (Randrianarivony et al. 2016a, b, c),
Vepris unifoliolata is used for postpartum hemorrhages and colic (Randrianarivony
et al. 2016c).
Local Food Uses
Vepris dainellii: The fruit is edible (Beentje 1994; Gachati 1989; Kokwaro 2009).
Vepris nobilis: The fruits are edible (Beentje 1994; Gachati 1989; Kokwaro 2009).
Vepris dainellii: The tough wood is used for furniture and agricultural implements,
and as firewood (Beentje 1994; Bekalo et al. 2009; Gachati 1989; Kokwaro 2009;
Bussmann et al. 2011).
Vepris nobilis: The wood is used for poles and posts in house building, fences, tool
handles, and utensils, e.g., bows, walking sticks, clubs, spear shafts, bows, and

(Rutaceae), Maasai war club
(rungu), Sekenani camp,
R:W., Bussmann)
spoons. The wood is excellent for turning and inlay work, and is also suitable for
heavy construction, flooring, joinery, shipbuilding, vehicle bodies, furniture, cabinet
work, mine props, sporting goods, agricultural implements, toys, novelties, and vats.
It is used as firewood and for charcoal production. The flowers are a source of nectar
for honeybees. The tree is occasionally planted for improving the soil by its leaf
litter, and as a shade and amenity tree (Beentje 1994; Gachati 1989; Kokwaro 2009).
Used to make tools and ploughs (Bussmann et al. 2011). Excellent for making
rungus and sticks (Bussmann et al. 2006). Eaten by livestock, for the construction
of fences, as firewood, and excellent to make rungus (Bussmann 2006) (Fig. 11).
Vepris simplicifolia: The wood is very similar to that of Vepris nobilis and used for
similar purposes (Beentje 1994; Gachati 1989; Kokwaro 2009). Used as firewood
(Bussmann et al. 2011; Tian 2017). Eaten by livestock, for the construction of
fences, as firewood, and excellent to make rungus (Bussmann 2006).
Vepris verdooriana is used as fishing poison (Neuwinger 2004). Vepris macrophylla
is used for construction (Rakotoarivelo et al. 2013). Vepris boviniana and Vepris
unifoliolata are used for protection spells, to destroy spells that prevent pregnancy,
and to improve agricultural production (Randrianarivony et al. 2016a, b, c).
References
doi.org/10.1186/1746-4269-5-26.
Bussmann RW. The forest vegetation of the Harenna escarpment (Bale Province, Ethiopia) sand
phytogeographical affinities. Phytocoenologia. 1997;27(1):1–23.
Plants (Basel). 2020;9:44. https://doi.org/10.3390/plants9010044.
gascar Conserv Dev. 2016b;11(2):44–51.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016c. ISBN 978-0-9960231-4-6.
pastoral Maasai girls in southern Kenya. J Ethnobiol Ethnomed. 2017;13:2. https://doi.org/
10.1186/s13002-016-0131-x.
Themeda triandra Forssk.
POACEAE

Grace N. Njoroge
Synonyms
Themeda triandra Forssk.: Andropogon tenuipedicellatus Steud., Anthistiria

argentea Nees, Anthistiria australis R. Br., Anthistiria brachyantha Boiss.,
Anthistiria ciliata Cav. ex Spreng., Anthistiria ciliata Nees, Anthistiria glauca
Desf., Anthistiria imberbis Retz., Anthistiria japonica Willd., Anthistiria paleacea
(Vahl.) Ball, Anthistiria punctata Hochst. ex A. Rich., Calamina imberbis (Retz.)
Roem. & Schult., Stipa arguens Thunb., Themeda barbinodis B.S. Sun & S. Wang,
Themeda forskahlii (Kunth) Hack., Themeda imberbis (Retz.) Cooke, Themeda
japonica (Willd.) Tanaka, Themeda triandra var. burchellii (Hack.) Stapf., Themeda
triandra var. japonica (Willd.) Makino, Themeda triandra var. punctata (Hochst. ex
A. Rich.) Stapf, Themeda triandra var. sublaevigata Chiov., Themeda triandra var.
syriaca (Boiss.) Hack., Themeda triandra var. trachyspathea Gooss., Themeda
triandra var. vulgaris auctt., non Hackel
R. W. Bussmann (*)
G. N. Njoroge

Local Names
Themeda triandra: Kikuyu: Nyaragita (Gachathi 1989).
Botany and Ecology
Themeda triandra Forssk: Perennial grass rising from a knotty rootstock. Culms
tussocky, yellowish, usually farinose near nodes, 0.5–1.5 m tall. Leaf sheaths usually
hispid with tubercle-based hairs; leaf blades 10–50 0.4–0.8 cm, glabrous or pilose,
finely acuminate; ligule 1–2 mm, truncate, ciliate. Compound panicle lax, open, with
spaced, often nodding spathate fascicles; spathes and spatheoles narrowly lanceo-
late, glabrous, or thinly to densely tuberculate-hispid, innermost 2–3.5 cm. Raceme
composed of a triad of 1 sessile and 2 pedicelled spikelets above the involucre of
2 homogamous pairs. Homogamous spikelets all sessile, arising at same level,
staminate, both glumes present, 7–14 mm, oblong-lanceolate with lateral scarious
wings, glabrous or hispid with tubercle-based hairs. Sessile spikelet 5–7 mm; callus
1.5–3 mm, pungent, brown bearded; lower glume dorsally rounded, dark brown,
glossy, hispidulous in upper 1/3, smooth below; awn 3.7–7 cm. Pedicelled spikelet
7–12 mm, male or barren. Flowering and fruiting June to December (Wu et al. 1994–
2013). An important component of African mountain grasslands (Bussmann 2006).
Themeda triandra: Used as fodder (Gachathi 1989).
References
2006;111:41–66.
Thunbergia alata Sims
ACANTHACEAE

Grace N. Njoroge
Local Names
Thunbergia alata: Kikuyu: Kanyanja; Luhya: Indeleresia, Kimira, Tsindelesia;

Luo: Nuawend-Agwata (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Thunbergia alata Sims: Vines, herbaceous. Stems more or less 4-angled to flat-
tened, bisulcate, pubescent. Petiole 1.5–3 cm, winged, sparsely pubescent, leaf blade
sagittate to deltoid ovate, 2–7.5 2–6 cm, abaxially hirsute, adaxially sparsely
strigose, palmately 5-veined, base hastate to cordate, margin entire or undulate, apex
acute. Flowers axillary, solitary, peduncle 2.5–3 cm, sparsely strigose, bracteoles
ovate, 1.5–1.8 1–1.4 cm, abaxially hirsute, 5–7-veined, apex acute, acuminate, or
obtuse. Calyx annular, unequally 10–13-lobed. Corolla orange with dark purple
R. W. Bussmann (*)
G. N. Njoroge

glandular “eye” in throat, 2.5–4.5 cm, tube basally cylindric for 2–4 mm, throat
1–1.5 cm, lobes obovate, apically truncate. Staminal filaments ca. 4 mm, glabrous,
anther thecae 3.5–4 mm, unequal, pubescent at margin and base. Ovary glabrous,
style ca. 8 mm, glabrous, stigma funnel-shaped, unequally 2-lobed, lower lobe
spreading, upper lobe erect. Capsule pubescent, basal part ca. 7 10 mm,
2-seeded, beak ca. 1.4 cm, ca. 3 mm wide at base. Seeds reticulate on dorsal surface.
Flowering October–March, fruiting February–May. (Wu et al. 1994–2013). Com-
mon in disturbed parts of Afromontane forests (Bussmann 2002; Bytebier and
Bussmann 2000) (Figs. 1 and 2).
Fig. 1 Thunbergia alata

(Acanthaceae), Bale
Fig. 2 Thunbergia alata

(Acanthaceae), Bale
Thunbergia alata Sims 1035
Thunbergia alata: Leaves crushed and extracted in water. This given to children for
mouth and tongue pain. Leaves and buds pounded in fat applied for backache and
joint pains. A decoction of the whole plant drunk by expectant mothers to strengthen
the placenta (Bekalo et al. 2009; Kokwaro 2009). Leaves boiled to treat cough,
backache, and problematic fetus placement in the womb (Jeruto et al. 2008).
Thunbergia alata: Sometimes browsed by cattle and wildlife, but generally useless
(Bussmann 2006; Bussmann et al. 2006, 2011). The leaves are boiled to treat
anaplasmosis and theileriosis in cattle (Njoroge and Bussmann 2006).
References
doi.org/10.1186/1746-4269-5-26.
Bytebier B, Bussmann RW. Vegetation of mount Nyiru Samburu District, Kenya: a checklist and
ment of cattle diseases among the kikuyus Central Kenya. J Ethnopharmacol. 2006;108:332–9.
Toddalia asiatica (L.) Lam.
RUTACEAE

Grace N. Njoroge
Synonyms
Toddalia asiatica (L.) Lam.: Aralia labordei H. Lév., Paullinia asiatica L.,
Toddalia aculeata Pers., Toddalia angustifolia Lam., Toddalia asiatica var. flori-
bunda (Wall.) Kurz, Toddalia asiatica var. gracilis Gamble, Toddalia asiatica var.
obtusifolia Gamble, Toddalia floribunda Wall., Toddalia rubricaulis Roem. &
Schult., Toddalia tonkinensis Gauillaumin, Zanthoxylum asiaticum (L.) Applehans,
Groppo & J. Wen.
Local Names
Toddalia asiatica: Kikuyu: Mururue; Digo: Kikombe-Cha-Chui; Kamba: Maluia;

Kipsigit: Chepindorwet; Luhya: Luabare; Luo: Ajua, Nayaluet-Kwach; Maa:
Ole-Pormunyo, Ole-Barmonyo; Marakwet: Kipkeres; Kipkutai; Meru:
R. W. Bussmann (*)
G. N. Njoroge

Mukongura; Nandi: Usuet; Ndorobo: Womboriot; Sanya: Kikucha; Samburu:

Llaramunyo; Tugen: Katemwe; Turkana: Etokebengu; Swahili: Mdakakomba
Botany and Ecology
Toddalia asiatica (L.) Lam.: A scrambling or climbing, retrorsely aculeolate shrub;

young branches minutely rusty pubescent or glabrescent. Branches with small
recurved prickles. Leaves 3-foliolate, petiole grooved above, leaflet Inflorescence
ferruginous-pubescent; bracts linear, 0.8–1.3 mm. long, caducous. Leaflets sessile,
elliptic or obovate or oblanceolate, 3–8 cm. long, 1.3–3 cm. wide, acuminate or
obtuse at the apex, entire or slightly crenulate; midrib prominent and occasionally
aculeolate on the lower surface; lateral nerves numerous but not raised; conspicu-
ously glandular on both surfaces; petiole 1–4 cm. long, grooved above and persis-
tently pubescent in the groove, sometimes aculeolate. Calyx small, with 5 acute
lobes. Flowers usually in cymose clusters on the panicle branches, unisexual by
abortion; pedicels 1–3 mm. long. Seeds 3–4 mm. long, dark brown, smooth. Male
flowers with ovate anthers 1 mm. long; rudimentary ovary glabrous, with a long
5-ribbed style. Petals triangular-lanceolate, about 2.5 mm. long, yellow. Fruit 7–
10 mm. across, orange when ripe. Female flowers with a very short style and 3–5-
lobed stigma; 5 staminodes present (Kokwaro 1982). Frequently collected especially
in Northern Kenya (Delbanco et al. 2017). Characteristic for disturbed Afromontane
forests (Bussmann 2002) (Figs. 1 and 2).
Fig. 1 Toddalia asiatica

National Park, Odo-Bulu
foreat, Ethiopia. (Photo
R.W. Bussmann)
Toddalia asiatica (L.) Lam. 1039
Fig. 2 Toddalia asiatica

National Park, Odo-Bulu
foreat, Ethiopia. (Photo
R.W. Bussmann)
Toddalia asiatica: The fruit is chewed for cough, the root decoction is used as emetic
and purgative (Maasda) (Beentje 1994; Gachati 1989; Kokwaro 2009). Leaves
boiled, and steam inhaled to treat nasal and bronchial problems. Fruit decoction
for cough and colds. The roots are chewed for cough and stomachache. Root juice
rubbed into incisions in ankles and wrists for snakebite treatment, and sometimes
used orally as emetic for the same purpose. Root decoction with milk to treat malaria.
The bark decoction is used for stomach problems, and externally to treat infected
wounds (Kokwaro 2009). The whole plant serves to treat malaria, fever, and colds in
Kenya (Bussmann et al. 2006; Njoroge and Bussmann 2006a, b), as well as backpain
in women (Njoroge et al. 2004; Njoroge and Bussmann 2009). The bark decoction
serves as emetic for digestion, respiratory problems, cold, and stomachache
(Nankaya et al. 2019, 2020). In Madagascar, Toddalia is also used to treat back
pain as well as stomach pain (Rabearivony et al. 2015; Rakotoarivelo et al. 2015),
epilepsy in infants, pulmonary infections, and externally as cataplasm
(Randrianarivony et al. 2016a, 2017), and as antimalarial (Rakotoarivelo et al.
2015). It is sold in markets as remedy for malaria, digestive complaints, fever,
cholera, diarrhea, rheumatism, syphilis (Randriamiharisoa et al. 2015), as well as
during pregnancy to avoid complications (Randrianarivony et al. 2016b, c). The
species is regarded useless in Southern Ethiopia (Bussmann et al. 2011).
Toddalia asiatica: In Madagascar, the species has cultural uses for disenchantment,
to protect oneself from evil spells, and to succeed in agricultural activities
(Randrianarivony et al. 2016a).
References
Delbanco AS, Burgess ND, Cuni-Sanchez A. Medicinal plant trade in northern Kenya: economic
2009;82:262–9.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016a. ISBN 978-0-9960231-4-6
gascar Conservation Dev. 2016b;11(2):44–51.
Madagascar. Indian J Tradit Knowl. 2016c;151:68–78.
2017; https://doi.org/10.1186/s13002-017-0147-x.
Trichilia emetica Vahl
MELIACEAE

Grace N. Njoroge
Synonyms
Trichilia emetica Vahl: Trichilia roka (Forrsk.) Chiov.
Local Names
Trichilia emetica: Kikuyu: Mururi; Chagga: Mchengo; Digo: Munwa, Madzi;

Luhya: Musinzi; Meru: Mutugati; Swahili: Muwamaji (Beentje 1994; Gachati
1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Fig. 1 Trichilia cf. emetica

(Meliaceae), flowering
specimen, Bale Mountains
National Park, Odio-Bulu
R.W. Bussmann)

(Meliaceae), flowers, Bale
Odio-Bulu forest, Ethiopia.
Botany and Ecology
Trichilia emetica Vahl: Evergreen or semi-evergreen tree, usually 8–20( 25) m tall;
crown very dense, wide-spreading in open; foliage very dark green; bark dark grey
or dark brown, rough or smooth. Staminal tube usually 0.8–1.1 cm long. Petals
(0.7–)1–1.6 cm long. Capsule (unopened but mature) 1.8–2.5 cm in diameter,
usually crowded at the ends of the branchlets. Leaves up to 28 cm long; leaflets
usually 9–11, elliptic or oblong-elliptic, up to 15 5 cm, nearly always broadest
near the middle and with the apex of lateral leaflets rounded, emarginate or broadly
acute without a hollow curve; lateral nerves usually in 11–18 closely set pairs; lower
surface sparsely to densely puberulous with short, weak curly or flexuous hairs,
Trichilia emetica Vahl 1043

(Meliaceae), young fruits,
Park, Odio-Bulu forest,
Ethiopia. (Photo
R.W. Bussmann)

(Meliaceae), fruit, Bale
Odio-Bulu forest, Ethiopia.
usually drying olive-green or pale yellow-brown. Calyx 0.35–0.5 cm long,

tomentellous, lobed almost to the base; lobes suborbicular, imbricate. Inflorescence
usually condensed and many-flowered (Styles and White 1991). In the undergrowth
of Afromontane forests (Bussmann 2002a, b). Generally only germinating under
light conditions (Bussmann and Lange 2000) (Figs. 1, 2, 3, and 4).
Trichilia emetica: The root infusion is used as emetic but is very toxic (Beentje
1994). Pounded bark infusion used for pneumonia as emetic and to cause sweating.
The root decoction also taken for colds, lumbago, as diuretic, and to induce labor.
Seeds are fried, and the seed oil applied to wounds from which jiggers were
extracted. The oil is also applied to the skin to prevent jigger infections, rheumatism,
and leprosy (Kokwaro 2009). The leaves of Trichilia dregeana are applied to skin
problems (Giday et al. 2009). The crushed roots of Trichilia prieuriana serve to treat
diarrhea (Yineger et al. 2007).
Trichilia emetica: The seed oil is used for soap, the timber for furniture (Beentje
1994). All species used as firewood (Bussmann et al. 2011). Used as fish poison,
together with Trichilia dregeana (Neuwinger 2004).
References
Nat Hist. 2000;891–2:101–11.
doi.org/10.1186/1746-4269-5-34.
Styles BT, White F. Flora of tropical East Africa: Meliaceae. Rotterdam: Balkema; 1991.
Typha domingensis Pers.
Typha latifolia L.
TYPHACEAE

Grace N. Njoroge
Synonyms
Typha domingensis Pers.: Typha angustata Bory. & Chaub., Typha angustifolia L.,
Typha angustifolia subsp. angustata (Boty. & Chaub.) Briq., Typha angustifolia var.
angustata (Bory & Chaub.) Jordanov, Typha angustifolia var. domingensis (Pers.)
Hemsl., Typha angustifolia var. virginica Tidestr., Typha australis Schumach., Typha
bracteata Greene, Typha domingensisis var. eudomingensis Geze, Typha tenuifolia
Kunth, Typha truxillensis Kunth
Typha latifolia L.: Typha crassa Raf., Typha elatior Raf., Typha latifolia fo. remota
Skvortsov, Typha latifolia subsp. eulatifolia Graebn., Typha latifolia var. typica
Rothm.
Local Names
Typha domingensis: Kikuyu: Ndothua (Gachati 1989; Kokwaro 2009)
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Typha domingensis Pers.: Erect shoots 1.5–4 m, not glaucous, flowering shoots 1–
2 cm thick in middle, stems 3–4 mm thick near spike. Leaves: sheath sides mem-
branous, margin broadly clear, summit tapered to blade or with persistent, membra-
nous auricles, mucilage glands at sheath-blade transition orange-brown, numerous
on entire sheath and proximal 1–10 cm of blade, widest blades on shoot 6–18 mm
wide when fresh, 5–15 mm when dry, distal blade about equaling inflorescence.
Inflorescences: staminate spike separated from pistillate by (0–)1–8 cm of naked
axis, about 1.4 X longer than pistillate, 1 cm thick at anthesis, staminate scales straw-
colored to mostly bright orange-brown, variable in same spike, linear to cuneate,
often laciniate distally, to 3–4 0.3 mm, pistillate spikes in flower when fresh bright
cinnamon-brown with whitish stigmas (drying brownish), later orange- (to medium)
brown, in fruit generally paler as stigmas and often bracteole blades wear off, about
6–35 cm 5–6 mm in flower, 15–25 mm in fruit, compound pedicels in fruit
peg-like, about 0.6–0.9 mm, pistillate bracteole blades forming spike surface before
flowering, later slightly exceeded by stigmas and slightly exceeding pistil hairs,
straw-colored to bright orange-brown, much paler than to nearly same color as
stigmas, irregularly narrowly to broadly spatulate or lanceolate, 0.8 0.1–
Fig. 1 Typha cf. domingensis

(Typhaceae), Naro Moru,
Kenya. (Photo
R.W. Bussmann)
Typha domingensis Pers. . . . 1047
0.3 mm, mostly wider than stigmas, apex variable in same inflorescence or different
plants, acute or acuminate. Staminate flowers 5 mm, anthers 2–2.5 mm, thecae
yellow, apex bright orange-brown, pollen in single grains. Pistillate flowers 2 mm
in flower, 8–9 mm in fruit, pistil-hair tips straw-colored to orange-brown in mass,
Fig. 2 Typha angustifolia

(Typhaceae), Kvemo Svaneti,
Georgia. (Photo
R.W. Bussmann)

Georgia. (Photo
R.W. Bussmann)

Georgia. (Photo
R.W. Bussmann)
usually with 1 subapical bright orange-brown, generally enlarged cell, stigmas often
deciduous in fruit, in flower erect, elongating, bending to form surface mat, white in
flower when fresh, later bright orange-brown, narrowly linear-lanceolate, about
1 0.1 mm, carpodia slightly exceeded by pistil hairs, usually evident at fruiting
spike surface, straw-colored, orange-spotted, apex broadly rounded (Napper 1971).
Rare in ponds at the edges of Afromontane forests (Bussmann 2002) (Figs. 1, 2, 3, 4,
5, and 6).
Typha latifolia L.: Erect shoots 150–300 cm, flowering shoots 1–2 cm thick in
middle, stems 3–7 mm thick near inflorescence. Leaves: usually glaucous when
fresh, sheath sides papery or membranous, margins narrowly clear, summit tapered
into blade to distinctly shouldered, or rarely with firm, papery auricles, mucilage
glands at sheath-blade transition usually colorless, obscure, absent from sheath
center and blade, widest blades on shoot 10–23( 29) mm wide when fresh,
5–20 mm when dry, distal blades about equaling inflorescence. Inflorescences:
staminate spikes contiguous with pistillate or in some clones separated by to 4
( 8) cm of naked axis, about as long as pistillate, about 1–2 cm thick at anthesis,
staminate scales colorless to straw-colored, filiform, simple, about 4 0.05 mm,
pistillate spikes in flower pale green drying brownish, later blackish brown or

Georgia. (Photo
R.W. Bussmann
reddish brown, in fruit often mottled with whitish patches of pistil-hair tips,
5–25 cm 5–8 mm in flower, 24–36 mm thick in fruit, compound pedicels in
fruit bristle-like, variable in same spike, 1.5–3.5 mm, pistillate bracteoles absent.
Staminate flowers 5–12 mm, anthers 1–3 mm, thecae yellow, apex dark brown,
pollen in tetrads. Pistillate flowers 2–3 mm in flower, 10–15 mm in fruit, pistil-hair
tips colorless, whitish in mass, not enlarged, stigmas persistent, forming solid layer
on spike surface, pale green in flower, drying brownish, then reddish brown or
usually distally blackish, spatulate, ovate to ovate-lanceolate, 0.6–1 0.2–0.25 mm,
carpodia exceeded by and hidden among pistil hairs, straw-colored, apex rounded.
Seeds numerous (Napper 1971).
Typha sp.: The pollen is used to purify breast milk, for urinary problems, and to stop
internal bleeding (Muhammad et al. 2019; Raj et al. 2018) and diarrhea (Umair et al.
2019). Also used to as a remedy for mumps, measles, gonorrhea,

Georgia. (Photo
R.W. Bussmann)
Local Food Uses
Typha sp.: Pollen sometimes used as food.
Typha sp.: Dried leaves are sometimes used to weave floor mats.
References
Kokwaro JO. Medicinal plants of East Afriabout. Nairobi: University of Nairobi Press; 2009.
Napper DM. Flora of tropical East Africa: Typhaceae. London: Crown Agents for Overseas
Governments & Administrations; 1971. 5 pp. ISBN 0-85592-009-2.
018-0208-9.
10.1186/s13002-019-0285-4.
Urtica maasaica Mildbr.
URTICACEAE

Grace N. Njoroge
Local Names
Urtica maasaica: Kikuyu: Hatha, Thabai; Maa: Endamejoi; Shambaa: Tufia (Gachati
1989; Kokwaro 2009); English: Maasai stinging nettle, forest nettle; Swahili: Mpupu.
Botany and Ecology
Urtica maasaica Mildbr.: Erect perennial dioecious herb up to 2 m tall, with little-
branched about quadrangular stems, forming loose clumps from a creeping rhizome;
all parts of plant with fiercely stinging hairs 1.5–2 mm long, raised on protuberances
about 1 mm high, otherwise glabrous to puberulous. Leaves: stipules fused,
interpetiolar, (0.8–)1–2 cm long, 0.4–1 cm wide, with rounded apex and broadly
cuneate to cordate base, chartaceous, brown; petioles 1.5–4.5 cm long; lamina ovate,
7–13 cm long, 6–10.5 cm wide, base cordate, margin serrate or usually double
serrate, with numerous teeth on each side, apex acute to shortly acuminate; lateral
R. W. Bussmann (*)
G. N. Njoroge

nerves 6–9 pairs, lower 2–3 pairs running into subcordate leaf-base, all pairs
extending to leaf-margin and linked by a fine net of anastomosing nerves; upper
surface with stinging hairs and sometimes also a fine pubescence, cystoliths very
numerous, punctiform, lower surface with stinging hairs along the nerves. Inflores-
cences paired in the axils of the upper leaves, spike-like, 2.5–4.5 cm long, with the
flowers in small, densely cymose, bracted glomerules, sometimes the inflorescence
with a few side-branches near the base. Male flowers on pedicels 0.8–1.5 mm long;
perianth about 1 mm in diameter, puberulous to pubescent, occasionally with a few
stinging hairs, depressed globose in bud. Female flowers sessile; perianth about
1 mm long, pubescent, with numerous stinging hairs on the large pair of tepals. Male
flowers on pedicels 0.8–1.5 mm long; perianth about 1 mm in diameter, puberulous
to pubescent, occasionally with a few stinging hairs, depressed globose in bud.
Female flowers sessile; perianth about 1 mm long, pubescent, with numerous
stinging hairs on the large pair of tepals. Achene flattened, ovoid, about 1 mm
long, 0.8 mm wide, white, long enclosed in the persistent perianth but dispersed
without it. Clearings and natural open glades in upland rain-forest and moist
evergreen bushland, often near human dwellings, especially where cattle are kept;
(400–)1500–3250 m, especially in semideciduous, broad-leaved Afromontane for-
ests (Beetje 1994; Bussmann 2002) (Figs. 1 and 2).
Urtica maasaica: The Maasai use the leaves to cure stomach-ache (Maundu et al.
2001). In the Kisii area of Kenya, the leaves are used to treat malaria. In Tanzania,
the macerated roots and leaves are used for the treatment of hepatic diseases. In
Rwanda and Burundi, Urtica massaica is used alone and in mixtures with other plant
species to treat numerous ailments: bruises, injuries, fractures, venereal diseases,
rheumatism, and urethral leak. Macerated roots and leaves to treat liver problems, the
boiled root infusion for stomachache (Kokwaro 2009). Serves as antimalarial and to
Fig. 1 Urtica maasaica

(Urticaceae), Bale Mountains
forest. (Photo
R.W. Bussmann)
Urtica maasaica Mildbr. 1055
Fig. 2 Urtica maasaica

(Urticaceae), Bale Mountains
forest. (Photo
R.W. Bussmann)
treat ear infections (Njoroge and Bussmann 2006a, b). Sometimes as aphrodisiac, for
back pain in women, and menstrual regulation (Njoroge and Bussmann 2009).
Urtica dioica is also used for sciatica, as rheumatism, boils, and skin ailments, as
diuretic, for jaundice, as astringent and anthelminthic, and to treat kidney bleeding
(Bhat et al. 2013; Joshi et al. 2010; Malik et al. 2015; Njoroge et al. 2004).
Sometimes the plant is used to treat fractures and blood pressure instabilities
(Kunwar and Bussmann 2009, 2010, 2013, 2019, 2020). In Pakistan the plant is
used to treat headache, fever, and myalgia (Sher et al. 2016) and as emollient
(Ur-Rahman et al. 2018) for arthritis and nausea (Wali et al. 2019). Used against
muscle pain, for blood purification, fever, rheumatism, arthritis, blood circulation,
hemorrhages, hair loss, asthma, hemorrhoids, inflammation (general) (Bussmann
and Sharon 2015; Paniagua Zambrana et al. 2020; Li et al. 2020), for stomach
disorders (Ali et al. 2019), and for anemia and headache (Singh et al. 2017, 2019). In
Pakistan, it is used as astringent and anthelminthic (Muhammad et al. 2019).
Urtica ardens is also used for sciatica, as rheumatism, boils, and skin ailments (Bhat
et al. 2013).
Local Food Uses
Urtica maasaica: The leaves of Urtica massaica are wilted, boiled and eaten, or
used young and uncooked, as is the case with several other Urtica species through-
out the world. In Tanzania, this vegetable is considered a famine food, but in Uganda
it is locally more popular and frequently eaten.
Urtica dioica leaves are often eaten as salad or boiled for herb pies, especially in
spring, and often mixed with a large number of other species (Abbas et al. 2019;
Batsatsashvili et al. 2017; Bussmann 2017; Bussmann et al. 2014, 2016a, b, c,
2017a, b, 2018; Kunwar et al. 2009).
Urtica maasaica: In Uganda the leaves are used as a repellent against rats and for
protection of crops from grazing cattle. Also used to treat dietary deficiencies in
cattle (Njoroge and Bussmann 2006c).
Leaves of Urtica dioica are used as fodder (Bussmann et al. 2014, 2016a, b, c,
2017a, b). Leaves are used in traditional carpet weaving as source of green, and roots
as yellow dye.
References
Abbas Z, Alam J, Muhammad S, Bussmann RW, Mulk Khan S, Hussain M. Phyto-cultural diversity
of the Shigar valley Central Karakorum Baltistan, Northern Pakistan. Ethnobot Res Appl.
2019;18:32. https://doi.org/10.32859/era.18.31.1-18.
org/10.32859/era.18.24.1-19.
Tchelidze D, Aleksanyan A, Alizade V, Paniagua Zambrana NY, Bussmann RW. Urtica dioica
L. In: Bussmann RW, editor. Ethnobotany of the Caucasus. Cham: Springer International
Publishing; 2017. https://doi.org/10.1007/978-3-319-49412-8_140.
in Kedarnath Wildlife Sanctuary of Garhwal Himalaya, India. J Ethnobiol Ethnomed 2013;9:1
http://www.ethnobiomed.com/content/9/1/1
2017; XXVII, 746p. ISBN 978-3-319-49411-1.
978-0-9960231-2-2.
loss of diversity – Ethnobotanical travels in the Georgian Caucasus. Ethnobot Res Appl.
2014;12:237–313.
2167-0412.1000266.
Batsatsashvili K, Hart RE. Ethnobotany of Samtskhe-Javakheti, Sakartvelo Republic of Geor-
gia, Caucasus. Indian J Tradit Knowl. 2017a;161:7–24.
Urtica maasaica Mildbr. 1057

Sakartvelo Republic of Georgia, Caucasus. Indian J Tradit Knowl. 2018;171:7–33.
Biodiversität, Naturausstattung im Himalaya, vol III. Universität Erfurt; 2009. p. 475–89
macology of medicinal plants in Far-west Nepal. Ethnobotany Res Appl 2009;7:5–28.
Kunwar RM, Fadiman M, Thapa S, Acharya RP, Cameron M, Bussmann RW. Plant use values and
phytosociological indicators: implications for conservation in the Kailash sacred landscape,
Nepal. Ecol Indic. 2020;108:105679. https://doi.org/10.1016/j.ecolind.2019.105679.
Li W, Liu B, Bussmann RW, Batsatsashvili K, Kikvidze Z. Urtica cannabina L.; Urtica dioica
L. In: Batsatsashvili K, Kikvidze Z, Bussmann RW, editors. Ethnobotany of mountain regions
Central Asia and Altai. Cham: Springer International Publishing; 2020. https://doi.org/10.1007/
978-3-319-77087-1_146-1.
R. Ethnobotany of the Loita Maasai people and plants working paper 8. Paris: UNESCO; 2001.
Njoroge GN, Bussamnn RW. Ethnotherapeutic management of sexually transmitted diseases STDs
and reproductive health conditions in Central Province, Kenya. Indian J Tradit Knowl
2009;82:262–269.
larial ethnophytotherapeutic remedies among the Kikuyus (Central Kenya). J Ethnobiol
ment of cattle diseases among the Kikuyus Central Kenya. J Ethnopharmacol. 2006c;108:332–9.
Njoroge GN, Gemill, B, Bussmann RW, Newton LE, Ngumi VW. Utilization of weed species as
Paniagua Zambrana NY, Bussmann RW, Echeverría J, Romero C. Urtica dioica L.; Urtica echinata
Benth.; Urtica magellanica Juss. ex Poir.; Urtica urens L. In: Paniagua Zambrana NY,
Bussmann RW, editors. Ethnobotany of mountain regions – ethnobotany of the Andes. Cham:
Springer International Publishing; 2020. https://doi.org/10.1007/978-3-319-77093-2_293.
https://doi.org/10.1007/978-3-319-77093-2_47-1-1.
2017;13:49. https://doi.org/10.1186/s13002-017-0178-3.
https://doi.org/10.32859/era.18.14.1-11.
Ethnobot Res Appl. 2019;18:35. https://doi.org/10.32859/era.18.35.1-30.
Uvaria acuminata Oliv.
Uvaria scheffleri L.L. Zhou, Y.C.F. Su & R.M.K
Saunders
ANNONACEAE

Grace N. Njoroge
Synonyms
Uvaria acuminata Oliv.: Uva acuminata Kuntze; Uvaria holstii Engl.
Local Names
Uvaria acuminata: Swahili: Mganda-simba; Boni: Tomorr; Digo: Mudzala,

Mumbweni; Giriama: Mrori; Sanya: Shinyole; Sukuma: Ihulungula; Ngoni:
Mhuani; Shambaa: Mleko (Beentje 1994; Gachati 1989; Kokwaro 2009).
Uvaria scheffleri: Kamba: Muguguma; Marakwet: Murguiyo; Pokot:
Tamingetwo (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Uvaria acuminata Oliv.: Much-branched aromatic shrub, small tree or liane 1.8–9 m
long or tall; branchlets with rather coarse granular stellate and often simple hairs when
young, later glabrescent or glabrous, blackish, wrinkled and lenticellate. Leaf-blades
oblanceolate, obovate or oblong, (0.7–)3–8( 12) cm long, (0.5–)1.3–3.9( 4.9) cm
wide, mostly distinctly acuminate at the apex or less often obtuse, rounded or more
usually cordate at the base, somewhat coriaceous, above mostly deep green and
sparsely pubescent with simple or stellate hairs or glabrous save on midrib, sparsely
to densely covered with coarse openly spaced ferruginous stellate hairs beneath and
with simple hairs on the nerves, rarely glabrous beneath; lateral nerves slightly
impressed above; venation reticulate, usually prominent on both surfaces; petiole
1–2.5( 4) cm long, stellate-pubescent. Flowers terminal or extra-axillary, solitary or
in 2-flowered cymes, hanging; peduncle more or less 2–3 mm long; pedicels 0.5–2 cm
long, stellate-tomentose; bracts oblong, 2–3 mm long, 1 mm wide; bracteoles minute,
1 mm long, soon deciduous. Sepals ovate or ovate-triangular, 4–6 mm long and wide,
obtuse to acute, shortly united at the base, often remaining as a loose persistent collar,
completely covering the petals in bud, pubescent with coarse ferruginous stellate hairs.
Petals white or yellow, oblong or oblong-lanceolate, subequal, 0.6–1.3 cm long,
3–6 mm wide, acute or obtuse, thin or slightly fleshy, puberulous on both surfaces
with stellate hairs. Stamens more or less 1 mm. long; connective-prolongation broad-
ened, truncate. Carpels mostly 20–25. Fruiting pedicels 1–2 cm long; monocarps
orange, 5–15, globose or ovoid, 0.8–1.6 cm long and wide, 1–4( 6)-seeded, not
constricted between the seeds, obtuse, densely tomentose with ferruginous or brown
hairs, rugose or verrucose; stipes 2–3 mm long. Seeds shining brown, irregularly
compressed-ovoid or shaped like the segment of an orange, 7–8 mm long, 4–5 mm
wide, 3–4 mm thick, usually with at least one sharp angle, with a paler irregularly
quadrate arillar collar. Thicket, bushland and dry rather scrubby forest, also in wetter
evergreen forest or woodland, predominantly coastal; 0–810 m.
Uvaria scheffleri L.L. Zhou, Y.C.F. Su & R.M.K Saunders: Scrambling shrub,
small tree or more usually a liane, 1.2–2.4(-several) m long or tall; young shoots
dark, flexuous, almost or quite glabrous, later grey-brown or purple-brown,
lenticellate. Leaf-blades oblong, oblong-elliptic or oblong-lanceolate, 1–14 cm
long, 0.7–5.8 cm wide, obtuse, acute, or somewhat acuminate at the apex, broadly
cuneate or rounded at the base, slightly coriaceous, deep shiny green, glabrous save
for a very few small hairs beneath when very young; venation closely reticulate,
slightly prominent on both surfaces; petiole 3 mm, long, channeled, with a few
ferruginous hairs; minute axillary buds are densely ferruginous pubescent. Flowers
more or less leaf-opposed, solitary; peduncle short; pedicels 0.8–1.8 cm long,
rugulose, glabrous or pubescent with scattered ferruginous stellate hairs or velvety;
bracteoles oblong, 2–3 mm long, 1 mm wide, very soon deciduous. Sepals almost
hemispherical or very broadly ovate, 2–2.5 mm long, 4–5 mm wide, obtuse, free, not
covering the petals in bud, sparsely to rather densely covered with stellate hairs.
Petals butter-yellow, outer more or less round, the inner narrower and more oblong,
clawed, 0.9–1.5 cm in diameter, very rounded at apex, not markedly fleshy, velvety
Uvaria acuminata Oliv. . . . 1061
ferruginous tomentose outside, sparsely so inside at apex. Stamens more or less

3 mm long; connective-prolongation thickened, truncate. Fruiting pedicels 1.2–2 cm
long; monocarps yellow to orange, 8–13( 16), subgloboseto cylindrical, (0.8–)1.8–
3.2 cm long, 6.5–11 mm wide, 1–8-seeded, mostly distinctly constricted between the
seeds or pairs of seeds, mostly apiculate, with scattered stellate hairs or more or less
glabrous, finely rugulose; stipes 0.4–1.3 cm long, often pubescent. Seeds brown,
somewhat shining, if 2 to a constriction then approximately semiglobular, with a
strongly convex surface and a more or less flat surface, but often quite ovoid, not
flattened, 8 mm long, 6–7 mm wide, 4–5.5 mm thick; hilum elliptic. (Verdcourt
1971). Evergreen thicket, scrub or forest; 900–1800 m. Often found in deciduous
Elaeodendron-Brachylaena forests and humid montane forests dominated by Lovoa
swynnertonii (Bussmann 2002).
Uvaria acuminata: Root decoction used for diarrhea (Digo) (Beentje 1994). Root
decoction to remedy painful menstruation, dysentery, snakebite, and painful breasts
(Kokwaro 2009). Smoke used to treat convulsions in children (Mohagheghzadeh
and Faridi 2006).
Uvaria scheffleri: Root bark decoction used for malaria (Digo) (Beentje 1994). Root
bark used for fever (Kamba), as well as for debility (Kokwaro 2009).
The roots of Uvaria leptocladon are used in Ethiopia for chest pain, tuberculosis,
cough, bloody diarrhea, malaria, boils, abscesses, and weight loss (Teklehaymanot
and Giday 2010).
Local Food Uses
Uvaria acuminata / Uvaria scheffleri: Fruit edible (Beentje 1994).
References
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi). J East Afr Nat Hist 2002;91 1/2:27–79,
Verdcourt B. Flora of tropical East Africa: Annonaceae. London: Crown Agents for Overseas
Vangueria apiculata K. Schum.
Vangueria infausta Burch.
Vangueria madagascariensis J.F. Gmel.
Vangueria volkensii K. Schum.
RUBIACEAE

Grace N. Njoroge
Synonyms
Vangueria apiculata K. Schum.: Vangueria longicalyx Robyns

Vangueria infausta Burch.: Canthium infaustum (Burch.) Baill.; Vangueria
tomentosa Hochst.
Vangueria madagascariensis J.F. Gmel.: Canthium edule (Vahl) Baill.; Canthium
maleolens Chiov.; Vangueria acutiloba Robyns; Vangueria commersonii Jacq.;
Vangueria cymosa C.F. Gaertn.; Vangueria edulis (Vahl) Vahl; Vangueria edulis
Lam.; Vangueria floribunds Robyns; Vangueria robynsii Tennant; Vangueria venosa
Robyns; Vavanga chinensis Rohr; Vavanga edulis Vahl
Local Names
Vangueria: Oromo: Sugurgura

Vangueria apiculata: Samburu: Lkoromosei; Kamba: Kikomoa, Mukomoa;
Kipsigis: Kimolwet; Luhya: Shikhomoli, Kumukomosi; Maa: Ol-gumi; Pokot:
R. W. Bussmann (*)
G. N. Njoroge

Tapirpirwa; Turkana: Emaler; Haa: Mugugunwa; Hehe: Msambalawe (Beentje

Vangueria infausta: Maa: Orgome,i Ol-gumi; Swahili / Digo: Mzango, Mviru,
Muiru; Kamba: Kikomoa, Mukomoa; Kipisigis: Kimolwet; Luo: Omuya; Nandi:
Kimoluet; Sanya: Bombo; Iraqw: Masalaramo; Hehe: Msaada; Luguru: Msada;
Bondei: Mvilu (Beentje 1994; Gachati 1989; Kokwaro 2009).
Vangueria madagascariensis: Hehe: Musambalawe; Kikuyu / Meru: Mubiru;
Borana: Bururi; Kamba: Kikomoa, Mukomoa; Kipsigis: Kimolwet; Luo: Anyuka,
Apindi; Maa: Ol-gumei; Pokot: Komolwo; Samburu: Rugumi, Ngoromusui;
Sebei: Kamolwet; Taveta: Mdaria; Tugut: Komolik (Beentje 1994; Gachati 1989;
Kokwaro 2009).
Vangueria volkensii: Samburu: Luguromosoi; Kamba: Kikomoa, Mukomoa;
Kikuyu: Mubiru, Mubiru-ngombe; Kipsigis: Kimolwet, Kimbuet; Luo: Anyuka,
Omugu; Ndorobo: Maldai (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Vangueria apiculata: Shrub, subscandent shrub or small spreading tree 1.8–12 m tall,
the stems sometimes several, virgate; branching often horizontal; bark grey, brown or
red-brown, smooth or finely ridged; young shoots mostly dark plum-colored and
lenticellate, quite glabrous. Leaf-blades elliptic, oblong, ovate or ovate-lanceolate to
lanceolate, 3–15( 17) cm long, 1.5–6( 8) cm wide, distinctly acuminate at the apex,
rounded, cuneate or occasionally subcordate at the base, often discolorous and venose
beneath when dry, quite glabrous or rarely slightly pubescent beneath; petiole 0.7–1
( 1.4) cm long; stipules with filiform part 3–9.5 mm long from a short broad base
1.5–2 mm long. Inflorescences typically short, 1–3.5 cm long including peduncle, fairly
lax to very condensed, the rhachis, etc., almost glabrous to usually pubescent or densely
shortly hairy; peduncle 0.5–1 cm. long, similarly hairy, with paired bracts 3 2 mm.
near apex; pedicels 2–4 mm long, pubescent. Calyx-tube subglobose, glabrescent to
densely shortly hairy, 1.5 mm in diameter; lobes oblong to linear, sometimes slightly
spathulate, (1.2–)3–7 mm long, glabrous or ciliolate. Buds distinctly apiculate at the
apex, the 5 appendages often quite long. Corolla glabrous, greenish white, or green to
yellow; tube 4–5 mm long; lobes narrowly to very narrowly triangular, 4–5 mm long.
Style 6–8 mm long; stigmatic club 1.2 mm long. Fruit green turning brown, subglobose
or sometimes irregularly ellipsoid where only one pyrene has developed, 1.7–2.2 cm
long, 1.4–2.2 cm wide, glabrous; pyrenes 0.9–1.7 cm long, 4–6 mm wide, 3–5
developing. Evergreen forest (including Juniperus and Podocarpus forest), riverine,
marsh-side and lake-side forest, bushland and thicket, grassland with scattered trees,
often on termite mounds or rocky outcrops; 900–2190 m. (Bridson and Verdcourt 1988,
1991; Verdcourt 1976; Bussmann 2002; Bytebier and Bussmann 2000).
Vangueria infausta: Shrub or small tree 1.5–8 m tall, very similar to the last species
save for the indumentum; trunk smooth, grey or eventually rough and ridged;
branches ridged, the young parts densely ferruginous, pubescent or velvety. Leaf-
blades elliptic, oblong-elliptic, ovate or sometimes almost round or lanceolate,
Vangueria apiculata K. Schum. . . . 1065
Fig. 1 Vangueria infausta

R:W. Bussmann)
4–30 cm long, 2.5–18 cm wide, rounded, subacute or acute to shortly acuminate at

the apex, rounded to cuneate or rarely subcordate at the base, often discolorous,
deeply pubescent to usually densely softly velvety on both surfaces, the hairs often
yellowish or ferruginous in dry state; petiole 0.3–1 cm long, similarly hairy; stipules
with base 2–4 mm long and apical part 0.3–1.2 cm long, hairy. Inflorescences
densely hairy, sometimes extensive and much branched but typically rather short,
branches 1.5–3.5 cm long, each 5–10-flowered; peduncles 6–8 mm long; pedicels
1–2.5 mm long, but those of the central flower of each dichasial element 3.5 mm
long. Calyx-tube 0.75–1.2 mm long; lobes obtusely triangular to narrowly oblong,
1–1.25( 2) mm long. Corolla green or yellow-green, typically densely spreading
hairy outside but glabrous in some variants; tube 3–4.5 mm long; lobes 3–4 mm long
with appendages up to 0.5 mm long or practically obsolete. Style 4.5–6 mm long;
stigmatic club yellow, 1 mm long. Fruit green, usually ripening to dull orange-brown
or purplish, depressed subglobose, 1.5–4.7 cm in diameter; pyrenes up to 1.3–2 cm
long, 0.6–1.3 cm wide, 5–8 mm thick (Bridson and Verdcourt 1988, 1991; Verdcourt
1976; Bussmann 2002). In humid Mountain Forests (Bussmann 2002; Bytebier and
Bussmann 2000) (Figs. 1 and 2).
Fig. 2 Vangueria infausta

R:W. Bussmann)
Vangueria madagascariensis: Shrub or small tree, 1.5–15 m tall, often multi-

stemmed and sometimes with a spreading crown; stems glabrous, longitudinally
ridged, with pale to dark bark, mostly smooth and unpeeling but peeling or powdery
in one variant. Leaf-blades narrowly to broadly elliptic or elliptic-lanceolate,
8–28 cm long, 3.2–15 cm wide, acute to shortly acuminate at the apex, cuneate to
rounded or less often subcordate at the base, entirely glabrous or sometimes very
young leaves pilose beneath and adult ones sparsely pubescent; petiole 0.8–1.8 cm
long; stipules with broad base 3–5 mm long, and narrow apex 0.4–1.8 cm long,
glabrous or pubescent. Inflorescence pubescent; branches 1–4.5 cm long, 7–10
flowered; main peduncles 1 cm long; pedicels 2 mm long, save those of central
flower of inflorescence which measure 4 mm. Calyx-tube 1.2–3 mm long; lobes
triangular-oblong to narrowly oblong, 0.5–1.5 mm long, pubescent. Corolla
greenish yellow, yellow or cream, glabrous or rarely with few hairs; tube
3–4.5 mm long; lobes 3.5–4.5 mm long, with appendages up to 0.5 mm long;
buds slightly to usually markedly acuminate or apiculate due to the corolla lobe-
appendages. Style 7–8 mm long; stigmatic club cylindrical, yellow, 1.2–1.5 mm
long. Fruits green to brownish, subglobose, 2.5–5 cm in diameter, with 4–5 pyrenes,
Vangueria apiculata K. Schum. . . . 1067
each 2 cm long, 1.2 cm wide, 8 mm thick, with thick woody walls 1–2 mm thick.
Seeds 1.6 cm long, 6 mm wide, 4.5 mm thick, narrowed at one end. Evergreen
forest, riverine forest and woodland, bushland, grassland with scattered trees, some-
times on rock outcrops and termite mounds; 0–2130 m (Bridson and Verdcourt 1988,
1991; Bussmann 2002). In humid Mountain Forests (Bussmann 2002; Bytebier and
Bussmann 2000).
Vangueria volkensii: Spreading shrub or small tree 2.4–9( 15) m tall, with dark
grey smooth or slightly fissured bark; shoots densely ferruginous velvety when
young, later dark and lenticellate. Leaf-blades ovate-oblong or elliptic to ovate,
3–17( 26) cm long, 1.5–10( 14) cm wide, acuminate at the apex, cuneate to
rounded or subcordate at the base, densely pubescent to velvety all over with
yellowish hairs; petiole 0.5–1.3 cm long; stipules with base 2 mm. long and filiform
part 0.5–1.2 cm long. Inflorescences short, 1–4 cm long, very densely shortly hairy;
peduncle 0.6–2 cm long, with bracts as in last species; pedicels 0.75–3 mm long,
densely shortly hairy. Calyx-tube subglobose, densely shortly hairy, 1–1.5 mm in
diameter; lobes linear or oblong, 1.5–6( 8) mm long, densely hairy. Buds distinctly
apiculate, often the 5 appendages very marked. Corolla bright green or yellow-green,
greenish cream inside, sparsely to densely hairy outside, sometimes only on the
lobes, the tube glabrous; tube 3.5–5.5 m. long; lobes narrowly triangular,
4–4.5 mm long. Fruit eventually turning brown, subglobose, 2–2.5 cm long or
sometimes asymmetric and ellipsoid if reduced to 1 pyrene by abortion; pyrenes
4–5 normally, up to 1.6 cm long, 7–8 mm wide, 6 mm thick. In humid Mountain
Forests (Bridson and Verdcourt 1988, 1991; Verdcourt 1976; Bussmann 2002;
Bytebier and Bussmann 2000).
Vangueria apiculata: Leaves chewed for stomachache. Boiled root decoction as

anthelminthic. The leaf infusion can be drunk for swellings and warts. Leaf ash for
tuberculosis (Kokwaro 2009; Muthee et al. 2011).
Vangueria infausta: Root decoction used to treat hookworm, also for cough,
gonorrhea, and infertility (Kokwaro 2009).
Vangueria madagascariensis: Root decoction as anthelminthic. The bark infusion is
used to treat malaria but shows toxicity (Kokwaro 2009; Muthee et al. 2011).
Local Food Uses
The fruits of Vangueria infausta are eaten, especially by children and baboons
(Bussmann et al. 2006, 2017). Fruits of Vangueria apiculata, V. madagascariensis,
and V. volkensii are being eaten in Northern Kenya (Bussmann 2006). The fruit of
V. madagascariensis is used to flavor beer (Beentje 1994).
Vangueria species are used for construction, to link roof beams and posts
(Bussmann et al. 2011). Vangueria apiculata, V. madagascariensis, and
V. volkensii are used for fence construction (Bussmann 2006).
References
1988.
1991.
Sekenani Valley, Maasai Mara, Kenya. Econ Botany. 2017;722:207–16.
Muthee J.K., Gakuya D.W, Mbaria J.M., Kareru P.G., Mulei C.M., Njonge F.K. 2011. Ethnobo-
tanical study of anthelmintic and other medicinal plants traditionally used in Loitoktok district
of Kenya. J Ethnopharmacol 135: 15–21.
Verdcourt B. Flora of Tropical East Africa: Rubiaceae (Part 1). London: Crown Agents for Overseas
Governments & Administrations; 1976. 414 pp. ISBN 0-85592-043-32 Invalid ISBN [Fl. Trop.
E. Africa, Rub. (Part 1)].
Vernonia amygdalina Deille
Vernonia auriculifera Hiern
Vernonia galamensis (Cass.) Less.
ASTERACEAE

Grace N. Njoroge
Synonyms
Vernonia amygdalina Deille: Gymanantheum amygdalinum (Delille) Walp.

Vernonia auriculifera Hiern: Vernonia laurentii De Wild., Vernonia uniflora
Hutch. & Dalziel, Gymnanthemum auriculiferum (Hiern) Isawumi
Vernonia galamensis (Cass.) Less.: Centraplus galamensis Cass., Centraplus
pauciflorus (Willd.) H. Rob., Vernonia pauciflora (Willd.) Less.
Local Names
Vernonia amygdalina: Acholi: Labwori; Nyankore: Omuburiri; Kikuyu:

Muthakawa; Kisii: Ormororia; Luhya: Omulusya, Musuritsa, Olumulusya,
Lisulushita; Luo: Olusia; Nandi: Cheburiander (Beentje 1994; Gachati 1989;
Kokwaro 2009); English: Bitterleaf, Common bitterleaf; French: Vernonie,
Vernonie commune, Ndole.
R. W. Bussmann (*)
G. N. Njoroge

Vernonia auriculifera: Luganda: Kikokooma; Kikuyu: Muthakawa, Muchatha,

Musakawa; Kisii: Musabakawa; Luo: Olusia; Maa: Ol-masakawa; Marakwet:
Turogogwa, Tbubagawa; Sebei: Tabenguet; Tugen: Tebinguet (Beentje 1994;
Gachati 1989; Kokwaro 2009); English: Bitterleaf, Common bitterleaf; French:
Vernonie, Vernonie commune, Ndole.
Vernonia galamensis: English: Ironweed
Botany and Ecology
Vernonia amygdalina Deille: Shrub or small tree up to 10 m tall, much branched,

trunk up to 40 cm in diameter, bark grey to brown, smooth, becoming fissured, young
branches densely pubescent. Leaves alternate, simple, stipules absent, petiole
0.2–4 cm long, blade ovate-elliptical to lanceolate, 4–15( 28) cm 1–4( 15) cm,
cuneate or rounded at base, shortly acuminate at apex, margin minutely toothed to
coarsely serrate, finely pubescent but often glabrescent, pinnately veined. Inflores-
cence a head, arranged in terminal, compound, umbel-like cymes, stalk of head up to
1 cm long, pubescent, involucre cylindrical to broadly ellipsoid, 3–5 mm long, bracts
3–7-seriate, 1–4.5 mm long, appressed. Flowers bisexual, regular, 5-merous, strongly
exserted from the involucre, pappus consisting of outer linear, caducous scales up to
1.5 mm long and of inner creamy or brownish bristles 4–7 mm long, corolla tubular,
5–8 mm long, whitish, glandular, with erect lobes, stamens with anthers united into a
tube, with appendages at apex, ovary inferior, 1-celled, pubescent and glandular, style
hairy, 2-branched. Fruit a 10-ribbed achene 1.5–3.5 mm long, pubescent and glandu-
lar, brown to black, crowned by the much longer pappus bristles. Seedling with epigeal
germination. Vernonia amygdalina occurs wild in most countries of tropical Africa,
from Guinea east to Somalia and south to north-eastern South Africa, and in Yemen
(Beentje 2000, 2002; Beentje et al. 2005). In the undergrowth of semidry Afromontane
forests (Bussmann 2002) (Figs. 1, 2, 3, 4, 5, and 6).
Vernonia auriculifera Hiern: Small tree, shrub, or woody herb up to 9 m tall, stems
branching from low down, at first brownish hairy, bark smooth, grey. Leaves
petiolate or especially the upper ones sessile, petiole up to 7.5 cm long, often
auriculate, blade elliptical or narrowly obovate to ovate, 10–45 cm 3–23 cm,
attenuate towards the rounded, truncate or cordate base, apex acute or shortly
acuminate, margin toothed, upper surface dull green, thinly fine-hairy mainly on
veins and glabrescent, lower surface whitish with often floccose tomentum. Inflo-
rescence composed of numerous heads in very large terminal compound thyrsoid
cymes, the whole inflorescence up to 150 cm in diameter, involucre narrowly
cylindrical, 4–6 mm long, phyllaries in 6–7 rows, appressed, green, especially the
inner rows often tinged purplish in the upper part, broadly ovate to ovate or elliptical,
apex obtuse to rounded, minutely apiculate, the inner rows elliptical, 4–6 mm long,
usually acute, the innermost eventually falling. Florets 1 per head, corolla 6.5–9 mm
long, scented, mauve, lilac or purplish, fading to white with age, lobes 2–3 mm long,
glabrous except the throat, anthers distally with linear or lanceolate appendage, style
branches long-tapering with stigmatic papillae above near the base. Fruit an achene
Vernonia amygdalina Deille. . . 1071
Fig. 1 Vernonia
cf. amygdalina (Asteraceae),
Park, Odo Bulu Forest,
Ethiopia. (Photo
R.W. Bussmann)
Fig. 2 Vernonia
cf. amygdalina (Asteraceae),
Ethiopia. (Photo
R.W. Bussmann)
3–4 mm long, 8–10-ribbed, shortly ascending-pubescent to almost glabrescent

except at the apex, outer pappus of narrow scales up to 1.5 mm long, inner pappus
white, 5–8 mm long (Beentje 2000, 2002; Beentje et al. 2005). In the undergrowth of
semidry Afromontane forests (Bussmann 2002).
Vernonia galamensis (Cass.) Less.: Usually annual herb up to 3( 5) m tall, but
mostly much smaller, stems ribbed, finely to coarsely hairy, sometimes branching
near the top. Leaves alternate, rather crowded, simple, sessile, blade elliptical to
linear, up to 25 cm 5 cm, base cuneate, apex acuminate, margins toothed, hairy on
both surfaces, but glabrescent. Inflorescence a head, solitary or few to many in a
terminal, lax to rather dense, leafy cyme, peduncle stout, pubescent, involucre ovoid
to nearly globose, 8–25 mm 1–15 mm, involucral bracts in 4–6 rows, pale green
often with darker tip, outer ones linear, short, middle ones often hardened at base,
Fig. 3 Vernonia
cf. galamensis (Asteraceae),
Ethiopia. (Photo
R.W. Bussmann)
tips usually leaf-like, inner ones oblong to narrow-lanceolate and acuminate, some-
what dry membranous. Flowers normally bisexual and fertile, long exserted, corolla
7.5–16 mm long, lower half tubular, gradually expanding above, bright blue to pale
mauve, pink, purple, violet, or almost white, sometimes flushed pale yellow or green,
lobes 5, linear, 2–7 mm long, glandular, stamens 5, slightly exserted, anthers united
into a tube, ovary inferior, style exserted, 2-branched. Fruit a narrowly obovoid
achene up to 8 mm long, with 10 equal, narrow ribs, dark brown to black, densely
appressed hairy, pappus in 2 whorls, outer pappus of up to 2 mm long barbed bristles,
inner pappus of up to 11 mm long barbed bristles. Occurs naturally from Cape Verde
and Senegal east to Eritrea and through East Africa south to Mozambique. The
greatest diversity is found in East Africa, in West Africa only a single variety occurs.
(Beentje 2000, 2002; Beentje et al. 2005). In the undergrowth of semidry
Afromontane forests (Bussmann 2002).
Vernonia amygdalina: Commonly used in traditional medicine. Leaf decoctions

are used to treat fever, malaria, diarrhea, dysentery, hepatitis, and cough, as a
laxative and as a fertility inducer. They are also used as a medicine for scabies,
Fig. 4 Vernonia
Ethiopia. (Photo
R.W. Bussmann)
headache, and stomach-ache. Root extracts are also used as treatment against
malaria and gastrointestinal disorders. In Nigeria, leaves are placed on a wound
as a substitute for iodine. One of the most common medicinal uses of Vernonia
amygdalina is as a treatment against intestinal worms including nematodes, espe-
cially Enterobius vermicularis (Kokwaro 2009). In Zimbabwe, a root infusion is
used to treat sexually transmitted diseases. Bark infusions are also taken to treat
fever and diarrhea, dried flowers against stomach disorders. Young twigs are used
as toothpicks or chewing sticks (Bekalo et al. 2009), and for skin lesions (Kumar
et al. 2011; Wondimu et al. 2007; Yineger et al. 2007), and as antimalarial (Flatie
et al. 2009; Tabuti 2008), and for diarrhea (Flatie et al. 2009). Also applied
externally to skin rashes and wounds (Gedif and Hahn 2003; Giday et al. 2003,
2009a, b), as anthelminthic (Giday et al. 2007), for intestinal disorders and skin
problems (Kumar et al. 2011).
Vernonia auriculifera: In Kenya used in traditional medicine in similar manners as
Vernonia amygdalina. The Meinit people in Ethiopia apply the root topically against
toothache. In Tanzania, the root soaked in water is a purgative for children. In DR
Congo, a drop of juice from the crushed stem bark is instilled in each nostril to treat
headache. Leaf preparations are used in various countries against dysentery and
stomachache. In Uganda, an infusion of the leaves is taken against worms and a leaf
decoction is drunk for the treatment of malaria. Leaf extracts are drunk as oxytocic
Fig. 5 Vernonia
Ethiopia. (Photo
R.W. Bussmann)
and abortifacient and against post-partum pains. Pulverized leaves are used against
impetigo, and in Congo, the dried and pounded leaves are applied on wounds. In
Cameroon, the leaf juice is used as eye drops for the treatment of cataract. The leaf
infusion is used to treat measles and for indigestion (Kokwaro 2009).
Vernonia galamensis: The leaf extract is drunk as a tea to treat chest pain. In Kenya,
the plant is used to treat stomach pain. Used for eye problems (Bussmann 2006).
Applied to external injuries and infected wounds (Teklehaymanot and Giday 2010),
for tonsillitis (Teklehaymanot 2009).
Vernonia anthelminthica serves for intestinal problems, fever, and skin ailments,
Vernonia cinerea is used for diarrhea, colds and cough (Bhat et al. 2013), also for
cholera, dysentery, constipation, fever, impotency, lactation, malaria, night blind-
ness, piles, skin diseases, threadworm, spleen complaints, and wounds (Verma et al.
2007). Vernonia poissonii sees medicinal use in Madagascar (Randrianarivony et al.
2017), where Vernonia appendiculata is sold in markets for the treatment for fever
and nervous system disorders (Randriamiharisoa et al. 2015), as well as dental
cavities (Rabearivony et al. 2015; Razafindraibe et al. 2013). Vernonia lasiopus is
used for postpartum pain, skin rashes, stomachache, back pain, diarrhea, and as
anthelminthic (Njoroge et al. 2004; Njoroge and Bussmann 2007) and for common
cold and otitis media (Njoroge and Bussmann 2006). Vernonia adoensis is used to
treat menstrual disorders (Teklehaymanot e complaints and wounds t al. 2007), as
Fig. 6 Vernonia
Ethiopia. (Photo
R.W. Bussmann)
well as gastrointestinal problems and poisoning (Giday et al. 2007). Veronica

exserta is used for parasites and chickenpox (Razafindraibe et al. 2013).
Local Food Uses
Vernonia amygdalina: A highly appreciated vegetable in West and Central Africa

and can be consumed in various dishes. In Nigeria, where the Yoruba name for this
crop is “ewuro” and the Igbo call it “onugbu,” leaves are boiled in soups. Leaves are
sometimes sold in the market after being shredded, parboiled, and made into fist-
sized balls. In Cameroon, the processed leaves are cooked with meat and/or prawns
mixed with ground peanuts to make a famous dish called “ndole.” Alternatively,
whole leaves are cooked together with cassava or yam tubers, whereas the leaves are
also dried and ground to powder for use in soups. In Cameroon, the leaves are
sometimes eaten unprocessed and raw mixed with palm oil and salt.
Vernonia galamensis: In Tanzania, the leaves are cooked in porridge.
Vernonia amygdalina: The leaves are browsed by goats. Dry stems and branches
provide fuel. The plant is sometimes grown as a hedge. The branches are used as stakes
to line fields. Vernonia amygdalina is also useful as a control agent against diseases in
plants. The ash from burnt branches is used to control seed-borne fungi (Curvularia,
Aspergillus, Fusarium and Penicillium spp.), thus ameliorating seed viability and
germination capacity. It has also been used for brewing beer as a substitute for hop
and as a well-known bee plant. Branches termite resistant (Beentje 1994; Gachati 1989;
Kokwaro 2009). The leaf juice is used as anthelminthic in livestock (Kokwaro 2009). It
is used for hut construction, fodder, firewood, and stomach problems in livestock
(Bussmann et al. 2011). Used to treat blackleg in livestock (Wondimu et al. 2007;
Yineger and Yewhalaw 2007), and for psychosomatic disoprders like “devil’s sickness”
and evil eye (Giday et al. 2010; Teklehaymanot and Giday 2007).
Vernonia auriculifera: In Kenya, Maasai use the stems and leaves in hut-construction
and Kipsigis use these materials to make platforms in hut-roofs for grain storage. In
Kenya, the leaves are also used for wrapping plant drugs to be roasted and used as a
poultice. In Uganda, the leaves are used as a substitute for toilet paper. The stems are
burnt as fuel. In Rwanda and DR Congo, the leaves and young twigs are eaten by
domestic animals. The stems, leaves, and flowers yield a dye: green colored without
mordant or with alum, and golden colored with chrome. Vernonia auriculifera is left to
grow or sometimes planted as a fallow plant to improve the soil or as a shade-providing
nurse tree. In Ethiopia, it is said that when the plant flowers, it is time to sow millet.
Leaves and stems used for hut construction (Beentje 1994; Gachati 1989; Kokwaro
2009). Used to make medicine containers (Kokwaro 2009).
Vernonia galamensis: The high oil content of the seed and the high content of vernolic
acid in the oil make it a potential oil crop. The oil, called “vernonia oil,” can be used in
the chemical (glue, paint and plastics), pharmaceutical, and agro-industrial industries.
In the paint industry, it is being tested as a component of low volatile-organic-solvent
paints. As a component of heat-baked films and coatings, vernonia oil provides
outstanding adhesion, flexibility, and chipping resistance, and good resistance to
alkaline, acid, and nonpolar solvents. In plastics, it can be used as a plasticizer of
PVC and as a structural component of polymers. The press-cake is suitable as animal
feed. The leaves have been smoked as a substitute for tobacco in Ethiopia.
A variety of Vernonia species are used as fishing poisons (Neuwinger 2004).
References
Beentje H, Jeffrey C, Hind DJN, editors. Flora of tropical East Africa: Compositae (Part 3). Kew:
Bekalo TH, Demissew Woodmata S, Asfaw WZ. An ethnobotanical study of medicinal plants used
by local people in the lowlands of Konta Special Woreda, southern nations, nationalities and
peoples regional state, Ethiopia. J Ethnobiol Ethnomed. 2009;5:26. https://doi.org/10.1186/

1746-4269-5-26.
http://www.ethnobiomed.com/content/9/1/1
appendices 1–7 @ http://www.naturekenya.org/JournalEANH.htm
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
group of Ethiopia: an ethnobotanical investigation. J Ethnobiol Ethnomed. 2009a;5:34. https://
doi.org/10.1186/1746-4269-5-34.
ethnobotanical study. J Ethnopharmacol. 2009b;124:513–21.
org/10.1186/s13002-015-0046-.
Mahaboboka, Amboronabo, Mikoboka, Southwestern Madagascar. J Ethnobiol Ethnomed
2017. https://doi.org/10.1186/s13002-017-0147-x.

2013;9:73.
1746-4269-3-12.
Vigna friesiorum Harms
Vigna membranacea A. Rich.
Vigna subterranea (L.) Verdc.
Vigna unguiculata (L.) Walp.
FABACEAE

Grace N. Njoroge
Synonyms
Vigna subterranea (L.) Verdc.: Arachidna trifolia Adans. ex A. Chev.; Arachis

africana Burm. f.; Cryptolobus africanus Spreng.; Cryptolobus subterraneus (L.)
Spreng.; Geolobus flavus Raf.; Glycine subterranea L.; Tetrodea subterranea (L.)
Raf.; Voandzeia subterranea (L.) Thouars
Vigna unguiculata (L.) Walp.: Dolichos biflorus L.; Dolichos capitatus Wall.;
Dolichos catjang Burm. f.; Dolichos catjang L.; Dolichos cylindricus Moench;
Dolichos echinatulus Blanco; Dolichos hastatus Lour.; Dolichos hastifolius
Schnizl.; Dolichos hastifolius Stokes; Dolichos leucomelas Kunze; Dolichos lubia
Forssk.; Dolichos melanophthalmus DC.; Dolichos monachalis Brot.; Dolichos
obliquifoliu Schnizl.; Dolichos sesquipedalis L.; Dolichos sinensis Forssk.;
Dolichos sinensis L.; Dolichos sphaerospermus (L.) DC.; Dolichos subracemosus
Jacq.; Dolichos tranquebaricus Jacq.; Dolichos unguiculatus L.; Dolichos
unguiculatis Thunb.; Phaseolus cylindricus L.; Phaseolus ellipticus Schur; Scytalis
R. W. Bussmann (*)
e-mail: rbussmann@gmail.c; rainer.bussmann@iliauni.edu.ge
G. N. Njoroge

catjang (Burm. f.) E. Mey.; Vigna catjang (L.) Walp.; Vigna cylindrica (L.) Skeels;
Vigna sesquipedalis (L.) Fruwirth; Vigna sinensis (L.) Endl. ex Hassk.; Vigna
sinensis (L.) Savi ex Hassk.; Vigna unguiculata subsp. catjang (Burm. f.) Chiov.;
Vigna unguiculata subsp. cylindrica (L.) Verdc.; Vigna unguiculata var. catjang
(Burm. f.) Bertoni; Vigna unguiculata var. cylindrica (L.) H. Ohashi; Vigna
unguiculata var. spontanea (Schweinf.) Pasquet
Local Names
Vigna schimperi: Samburu: Loyieti; Luhya: Mulanda (Kokwaro 2009).

Vigna subterranea: English: Bambara groundnut.
Vigna unguiculata: Kikuyu: Thoroko; Nyankore: Akaihakuru; Luo: Alot-Bo;
Padhola: Bole (Beentje 1994; Gachati 1989; Kokwaro 2009); English: Cowpea.
Botany and Ecology
Vigna friesiorum: Erect or decumbent perennial herb, 7·6–30 cm tall, from a softly
woody carrot-like or fusiform rootstock up to 25 cm long. Stems with dense to
scattered spreading hairs. Leaflets 3, oblong or elliptic to linear-lanceolate, 0·8–7 cm
long, 0·3–3·2 cm wide, acute to rounded at the apex, rounded at the base, glabrous to
pubescent or pilose on both surfaces, often rather characteristically margined with
short hairs, 3-nerved from the base; venation raised and reticulate on both surfaces;
petiole 1·5–8 cm long; rhachis 2·5–9 mm long; petiolules 0.5–1.5 mm long; stipules
oblong-lanceolate to lanceolate, 4–7 mm long, somewhat bilobed at the base, veined.
Racemes axillary, (1-) 2-several-flowered, subumbellate; peduncle 3·5–20 cm long,
somewhat thickened towards the apex; pedicels 1–2 mm long; bracteoles sub-
persistent, minute, 1.5 mm long, 1-nerved. Calyx-tube glabrous or hairy, truncate
at the base in fruit, nerved, 2–3·5 mm long; lobes deltoid, 2–3·5 mm long, ciliate or
minutely ciliolate, sometimes slightly tuberculate, the upper pair joined for two-thirds
or more of their length to form a bifid lip. Standard pale creamy-yellow, green with or
without mauve basal stripes or entirely mauve, obovate, 1–1·4 cm long, 0·8–1·3 cm
wide, emarginate, glabrous; wings green with mauve margins; keel green, elongated,
shaped very much like the end of a canoe. Pods erect, linear-cylindrical, 4–6 cm
long, 2–3 mm wide, yellowish pubescent or covered with very minute scale-like hairs,
17–20-seeded. Seeds brown, sometimes faintly mottled with black, subcylindrical
or rectangular, longest dimension 1.8–2.5 mm, shorter dimensions 1.5–2 mm; hilum
minute, central or eccentric; aril not developed.
Vigna membranacea: Annual or perennial climbing or trailing herb, more rarely
suberect, 0.2–1.8 m long. Stems ridged, at first densely spreading pilose with usually
rather tubercular-based hairs, later glabrescent. Leaflets 3, sometimes with a pale
chevron mark, ovate or ovate-triangular in outline, 1.2–10 cm long, 1.1–6.5 cm wide,
rounded or subacute to acuminate at the apex, rounded, truncate or subhastate at the
base, sometimes distinctly 3-lobed, at first rather dense ad pressed silvery pilose, later
Vigna friesiorum Harms. . . 1081
more sparsely pilose; petiole 1–8.5(-14) cm long; rhachis 0.5–1.4(1.9) cm long;

petiolules 1–4 mm long; stipules lanceolate, 3–12 mm long, bilobed and unequally
prolonged at the base. Inflorescences axillary, subumbellate, few–several-flowered;
rhachis 0·3–2 cm long, with conspicuous glandular nodes; peduncle 1–45 cm long,
often uncinulate towards the apex; pedicels 2–4 mm long; bracteoles linear, 1.5–5 mm
long, deciduous. Calyx glabrous or pubescent or with longer tubercular based hairs;
tube 2–3 mm long; lobes deltoid to linear, 0.15–1.7 cm long, often drawn out at the
tips, the upper pair practically free so that the calyx appears equally 5-fid, the margins
often conspicuously ciliate with long hairs. Standard mauve, pink-magenta, or blue;
yellow inside at base with darker purple guide lines, sometimes greenish outside,
oblate, 1–2.3 cm long and wide, glabrous; wings blue, mauve or pink; keel pale, tinged
blue, sometimes twisted but beak not incurved. Pods erect, linear-cylindrical, 3–9.5 cm
long, 2–3 mm wide, covered with very short, curved, tubercular-based hairs or small
scale-like hairs, 14–25-seeded. Seeds dark red or grey with dense black mottling,
narrowly oblong, angular, longest dimension 2.5–4.5 mm, shorter dimension
1.5–2.8 mm, 1.3–2.5 mm thick; hilum small, subcentral; aril not developed.
Vigna subterranea: Annual herb with short compact to long loose sparsely pubes-
cent densely leafy creeping stems. Leaves held erect; leaflets elliptic, obovate, or
oblanceolate, 3–8 cm long, 0·8–4 cm wide, rounded or emarginate at the apex,
cuneate at the base; petiole 2–30 cm long; rhachis (0.1-)1–2.5 cm long; petiolules
1–3 mm long; stipules ovate, 3 mm long. Racemes 1–3-flowered; peduncle 0.5–2 cm
long. Calyx glabrous; tube 1 mm. long; lobes 1 mm long. Standard yellow,
obovate, 4–7 mm long. Pods 1–2.2 cm long, 1.2–1.8 cm wide, glabrous. Seeds
variously colored white, yellow, red, or blackish or variously mottled, longest
dimension 8·5–15 mm, shorter dimension 6·5–10 mm.
Vigna ungiculata: Annual or perennial erect trailing or climbing herb, 1–3 m long.
Stems striate, glabrous, or slightly hairy particularly near the petiole-bases, sometimes
with minute stiff hook-like hairs. Leaflets 3, ovate, rhomboid, or lanceolate, the laterals
oblique, 1.5–16.5 cm. long, 1–12.5 cm wide, all entire or terminal leaflets sub-hastate
to 3-lobed at the base and laterals lobed on the outer margin, shortly acuminate or
subacute and mucronulate, acute to rounded at the base, glabrous or sparsely pubes-
cent on both surfaces; petioles 1.5–13 cm long; rhachis 0.6–2.5 cm long; petiolules 2–
5 mm long; stipules sub-medifixed, constricted at the point of attachment, upper part
lanceolate, 0.6–2 cm long, spur narrower, 2–6 mm long. Inflorescences axillary,
few-several-flowered; rachis 0.5–4(–5) cm long; peduncle 2–36 cm long, usually
minutely uncinulate pubescent towards the apex; pedicels 1–2(–4) mm long; bracts
deciduous, lanceolate, 3–5 m long; bracteoles persistent, spathulate, 3–5 mm long.
Calyx glabrous; tube 3–5.5 mm long; lobes triangular-lanceolate, 2.5–14 mm long,
acuminate, ciliolate; upper pair joined for about half their length or only at the base.
Standard white, greenish, yellow or lilac-purple, paler outside, round, 1.2–3 cm long,
1–3.2(–4) cm. wide, rounded or emarginate, glabrous; wings blue to purple; keel
usually white or pale, not twisted. Pods erect or eventually hanging in cultivated forms,
linear-cylindrical, 5·5-10(-90 in some cultivated races) cm. long, 3–11 mm wide,
glabrous or minutely verruculose. Seeds white to dark red or black, often mottled
with black or brown, oblong or reniform, longest dimension 3.5–5(–12) mm, shorter
dimension 2–3.5(–6.5) mm, 2.2(–4.5) mm thick; hilum one-third to half the longest
dimension, eccentric; rim-aril slightly developed. Throughout tropical Africa, where
doubtless it originated, but now widely cultivated in some form or other throughout
warmer parts of the world.
The roots of a variety of Vigna sp. are ground and drunk with water in Ethiopia as
cancer treatment (Flatie et al. 2009).
Vigna schimperi: Leaves chewed for tonsillitis (Kokwaro 2009).
Vigna subterranea: In Senegal leaf preparations are applied as a poultice for
infected wounds and abscesses; leaf sap is applied to the eyes as a treatment for
epilepsy; pounded seeds mixed with water are used to treat cataracts; the roots of the
plant can be taken as an aphrodisiac.
Vigna ungiculata: Various parts of the cowpea are used medicinally. The leaves and
seeds are applied to skin infections as a poultice, the leaves are chewed to relieve
toothache and powdered carbonized seeds are applied to insect stings. The roots are
used as treatment for epilepsy, chest pain, constipation, and as an antidote to
snakebites. Roots used to expel the placenta. Leaves applied to burns and dried
leaves crushed, and powder snuffed for headaches (Kokwaro 2009).
Local Food Uses
Vigna schimperi: The root is eaten (Bussmann 2006).

Vigna subterranea: Bambara groundnut is cultivated mainly for its seeds which can
be boiled, roasted or fried to make a delicious snack or mixed in with maize or
plantains to serve as a meal. The seeds can be ground into flour after roasting and
used to prepare porridge or they can be soaked, boiled, and ground into a paste and
used in fried or steamed dishes popularly eaten in Nigeria. The flour can also be used
as a thickener in soups and stews and in Zambia it is commonly made into bread.
Milk can be made from the seeds and fermented products similar to tempeh and
dawadawa can be prepared. Vigna ungiculata: Cultivated mainly for its seeds,
which can be cooked and eaten alone or mixed in with other vegetables and spices
to produce a nutritious bean soup. Cowpea is eaten in large parts of Africa and in
many areas, it is the preferred pulse. In West Africa, cowpea seeds are ground into
flour and mixed with onions and spices to make cakes which are either deep dried
(“akara balls”) or steamed (“moin moin”) for a delicious snack. Cowpea flour can
also be processed into crackers and baby foods, as is the case in Senegal, Ghana, and
Benin. In the United States, green seeds are sometimes roasted like peanuts. The
leaves and pods of cowpea are eaten as vegetables which can be served boiled or
fried. Many people dry the leaves in the sun to preserve them so that they can be
eaten during the dry season when food is scarce. The best leaves to use are those
picked towards the end of the season, because these tend to grow under conditions of
Vigna friesiorum Harms. . . 1083
stress and are generally tastier. In some parts of Africa, such as Ethiopia and Sudan,
the roots are occasionally eaten.
Signa schimperi: Browsed by lifestock (Bussmann 2006).

Vigna subterranea: Besides being a food crop, the seeds and leafy shoots of
bambara groundnut, which are rich in protein and phosphorus, make good fodder
for pigs and poultry.
Vigna ungiculata: Its ability to fix nitrogen from the air makes cowpea a good
fertilizer for the soil. In rice farming, farmers grow cowpea either before or after the
rice crop, as this is found to increase the food production from the land area. In
folklore, cowpea is used in sacrifices by the Hausa and Yoruba tribes to drive away
evil and to pacify the spirits of sickly children. In Nigeria some varieties of cowpea
are cultivated for their strong fiber which is used to make fishing gear as well as to
produce good quality paper. The dry seeds also make a good alternative to coffee.
The high protein content of cowpea makes it an excellent fodder crop. Cowpea is
cultivated for fodder in West Africa, Asia, and Australia; it is used for grazing or
mixed in with dry cereals for animal feed.
The roots of Vigna schimperi are used as livestock fodder and boiled as food in
Northern Kenya (Bussmann 2006).
References
2009;5:14. https://doi.org/10.1186/1746-4269-5-14.
Vitex keniensis Turrill
LAMIACEAE

Grace N. Njoroge
Local Names
Vitex keniensis: Kikuyu: Muhuru; Meru: Muuru, Moru; English: Meru oak
Botany and Ecology
Vitex keniensis Turrill: Tree 12–30 m. tall, up to 1.8( 2.3) m in diameter, bole 12–
18 m tall, bark very thin, rough, and slightly fissured, slash creamy yellow turning
dirty green, stems, petioles and leaf-venation beneath with longer more shaggy
indumentum. Cymes ochraceous tomentose, somewhat more lax, forming axillary
panicles up to 12 cm long, 24 cm wide, peduncles 6.3–13 cm long, secondary
branches to 4.5 cm long, pedicels 1–4 mm. long, densely tomentose, bracts 0.5–
1 cm long, 1–4 mm. wide. Leaves 5-foliolate, leaflets obovate, 5.5–17 cm long, 3.2–
R. W. Bussmann (*)
G. N. Njoroge

8.5 cm wide, broadly rounded to obtusely acuminate at the apex, cuneate to rounded
at the base, coriaceous, usually drying darker than in V. fischeri, sparsely puberulous
above, paler beneath and completely covered with soft ochraceous tomentum and
glands but becoming sparser, petiole 13.5–17 cm long, petiolules absent or
intermediate one 0.6–4 cm long. Ovary globose, with simple hairs and sparse glands,
style filiform, 6 mm long, stigma-lobes linear. Corolla white or white tinged blue
with largest lobe mauve, tube curved, 5–6 mm long, ochraceous tomentose above
outside, limb 8–10 mm. wide, the largest lobe round or ovate, 3–5 mm long, 3.5 mm
wide, entire, the others 2.5–3 mm long, 2 mm wide, slightly emarginate, densely
ochraceous tomentose outside. Calyx densely ochraceous tomentose, rather more
distinctly toothed than in V. fischeri, tube 2 mm long, lengthening to 5–6 mm in fruit,
teeth broadly triangular, 0.5 mm long, lengthening to 3 mm. Drupes black with white
spots, obovoid, (1.1–)2.2–2.4 cm long, (0.8–)1.3–1.5 cm wide, shiny, glabrous,
rounded at the apex, contracted to the base, mesocarp fleshy, endocarp very
woody, the putamen 2 cm long, 1.1–1.2 cm wide, 6–7 mm diameter at the base,
obtusely 4-angled, calycine cup 1.1 cm wide (Verdcourt 1992). Vitex keniensis is an
important species in lower Afromontane forests dominated by Ocotea usambarensis
(Bussmann 2002a, b). It is one of the fastest growing primary forests species
(Bussmann 1999), but rather problematic to germinate (Bussmann and Lange 2000).
Vitex keniensis: Fruits are eaten for stomach problems (Kokwaro 2009).
Vitex negundo: Plant juice is used for headache. Leaf juice is useful for gastric
troubles and used for common cold, fever, and dermatitis (Kunwar et al. 2010), for
colds and as analgesic (Mohagheghzadeh and Faridi 2006). It is also used for
toothache (Kunwar et al. 2019), as well as indigestion, gas problems, and stomach-
ache (Raj et al. 2018; Ur-Rahman et al. 2018). In Pakistan the species is also used for
jaundice and kidney problems (Muhammad et al. 2019). Vitex agnus-castus is a
very well-known medicinal species employed for gynecological problems
(Mohagheghzadeh and Faridi 2006). Vitex lanigera is used in Madagascar to treat
diarrhea (Randrianarivony et al. 2016a, b, 2017), intestinal colics, epilepsy, gonor-
rhea, premature fontanelle closure (Randrianarivony et al. 2016c). Vitex beraviensis
serves to treat premature fontanelle closure (Randrianarivony et al. 2016c).
Vitex keniensis: Great timber (Beentje 1994; Gachati 1989; Kokwaro 2009).
In Madagascar, Vitex coursii is used for ceremonial purposes (Rakotoarivelo
et al. 2013). Vitex beraviensis is used for construction, as firewood, and for cultural
purposes (Randrianarivony et al. 2016c); Vitex lanigera is used for construction
(Randrianarivony et al. 2016c).
Vitex keniensis Turrill 1087
References
Nat Hist. 2000;891–2:101–11.
Kunwar RM, Shrestha K, Malla S, Acharya T, Sementelli A, Kutal D. Relation of medicinal plants,
their use patterns and availability in the lower Kailash Sacred Landscape, Nepal. Ethnobot Res
Appl 2019;187. https://doi.org/10.32859/era.18.6.1-14.
018-0208-9.
Randrianasolo A, Bussmann RW. Guide de plantes d’Ambalabe, vol. 1. St. Louis: William L.
Brown Center, MBG; 2013. ISBN 978-0-9848415-7-8.
vol. 1. St. Louis: William L. Brown Center, MBG; 2016c. ISBN 978-0-9960231-4-6.
2017. https://doi.org/10.1186/s13002-017-0147-x.
Verdcourt B. Flora of tropical East Africa: Verbenaceae. Rotterdam: Balkema; 1992. 155 pp. ISBN
90-6191-358-6 [Fl. Trop. E. Africa, Verben.].
Warburgia salutaris (Bertol.) Chiov.
CANELLACEAE

Grace N. Njoroge
Synonyms
Warburgia salutaris (Bertol.) Chiov.: Chibaca salutaris G. Bertol., Warburgia

breyeri R. Pott., Warburgia ugandensis Sprague
Local Names
Warburgia salutaris: Kikuyu: Munderendu, Muthiga, Muthaiga; Kisii:

Omenyakige; Luhya: Apacha; Luo: Sogo-Maitha, Abaki; Maa: Osokoni,
Ol-sogoni; Marakwet: Sekwan; Meru: Musunui; Kipsigis: Sogoet, Soget;
Tugen: Soke; Ndorobo: Marut; Luganda: Mukuzanume; Iraqw: Sakwenamo;
English: East African greenheart (Beentje 1994; Gachati 1989; Kokwaro 2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Warburgia salutaris (Bertol.) Chiov.: Tree or shrub 4–10(18) m tall with rough
mottled grey bark, red on inner side, branches striate and lenticellate. Leaves 4.5–
11 1.5–2.5 cm, oblong to oblong-lanceolate or elliptic, acute at the apex, cuneate
at the base, shiny above, paler beneath, densely pellucid-dotted, petiole 2–5 mm
long. Flowers solitary or in short 3-flowered cymes, peduncle 2–3 mm long, bracts
0.5 mm long, deciduous leaving conspicuous scars, pedicels c. 1.5 mm long. Sepals
2 3 mm, minutely ciliate. Petals: outer 4–5 3 mm, obovate, concave, sub-
coriaceous, inner yellow, 4 1.5 mm spathulate, thinner. Staminal tube 3–4 mm
long, anthers 1.5 mm long. Ovary 3 mm long, ovules 15–20. Fruits reddish-blue,
2–3 1.5–2.5 cm, pericarp coriaceous, wrinkled, seeds several, flattened. A com-
mon component in drier East African Mountain forests (Bussmann 2002a, b). The
species is highly overharvested (Bekalo et al. 2009; Hamilton 2004) (Figs. 1 and 2).
In Kenya used to remedy malaria and stomach problems (Maundu et al. 2001;
Njoroge and Bussmann 2006a; Bussmann et al. 2006, 2018), common cold and
chest pain/pneumonia, malaria, and toothache (Maundu et al. 2001; Njoroge and
Bussmann 2006b; Bussmann et al. 2006; Beentje 1994; Gachati 1989; Kokwaro
2009), as well as to treat back pain in women (Njoroge and Bussmann 2009). The
ground bark is used as emetic, bark with fat given to mothers for easy delivery
(Maundu et al. 2001). Dried bark chewed for stomachache, constipation, cough,
fever, toothache, muscle pain, weak joints, malaria, and body pains. Bark powder
can also be used. The roots are boiled as soup to prevent diarrhea (Kikuyu). The
leaves are used in baths for skin problems (Kokwaro 2009). Used for cleaning the
uterus, common cold (Nankaya et al. 2019), gastrointestinal problems, respiratory
disorders, musculo-skelettal problems, and malaria (Lulekal et al. 2008; Nankaya
Fig. 1 Warburgia salutaris

(Canellaceae), Sekenani
Warburgia salutaris (Bertol.) Chiov. 1091
Fig. 2 Warburgia salutaris

(Canellaceae), fruits,
Sekenani Camp, Maasai
Mara, Kenya. (Photo
R.W. Bussmann)
et al. 2020). Sometimes used as anthelminthic (Gafna et al. 2017). Smoke used for
cough and cold relief (Mohagheghzadeh and Faridi 2006). The plant extracts serve
also to remedy pneumonia, tonsillitis, constipation, and fever (Jeruto et al. 2008).
Local Food Uses
The fruits are dibble (Maundu et al. 2001; Beentje 1994; Gachati 1989; Kokwaro
2009).
Interestingly, in veterinary medicine the plant is used to treat ectoparasites, and also
as dietary supplement for cattle (Njoroge and Bussmann 2006c). The species is used
as timber, but never as firewood (Maundu et al. 2001). Wood used for construction
and furniture, resin used as glue (Beentje 1994; Gachati 1989; Kokwaro 2009).
References
doi.org/10.1186/1746-4269-5-26.
Imenti, Ngaia, Nyambeni, Loita, Nguruman, Nairobi). J East Afr Nat Hist 2002a;91 (1/2):27–79,
Hamilton AC. Medicinal plants, conservation and livelihoods. Biodivers Conserv. 2004;13:1477–
517.
10.1186/1746-4269-4-10.
R. Ethnobotany of the Loita Maasai people and plants working paper 8. Paris: UNESCO; 2001.
larial ethnophytotherapeutic remedies among the Kikuyus (Central Kenya). J Ethnobiol
of cattle diseases among the Kikuyus (Central Kenya). J Ethnopharmacol. 2006c;108:332–9.
Knowl. 2009;8(2):262–9.
Withania somnifera (L.) Dunal
SOLANACEAE

Grace N. Njoroge
Synonyms
Withania somnifera (L.) Dunal: Physalis somnifera L., Withania kansuensis Kunag
& A.M. Lu, Withania microphysalis Suess.
Local Names
Withania somnifera: Kikuyu: Murumbae; Borana: Hidigaga, Idigaga; Gabbra:

Idi; Iraqw: Xaxardu; Sandawi: Xoxorik’o; Kamba: Mwanzo; Kipsigis:
Chepterekiat, Chesamiset; Luo: Ofuyaendwa; Maa: Ol-asaiyet; Marakwet:
Kipkogai; Pokot: Lopotwo; Samburu: Leekurun, Lesayet; Sukuma: Mtemua,
Shimba; Tugut: Kabarra; Turkana: Emotoe (Beentje 1994; Gachati 1989; Kokwaro
2009).
R. W. Bussmann (*)
G. N. Njoroge

Botany and Ecology
Withania somnifera (L.) Dunal: Herbs perennial, 30–150 cm tall, pubescent with
dendritic hairs. Stems woody proximally, erect or reclining, branched, tomentose.
Petiole 1–2 cm, leaf blade ovate, obovate, or oblong, 2.5–12 2–7 cm, glabrescent
adaxially except along midvein, pubescent abaxially, base cuneate, apex acute.
Inflorescences subsessile clusters of 4–6 flowers, peduncle obsolete. Pedicel
ca. 5 mm. Calyx campanulate, 3–5 mm, tomentose, lobes deltate, 1–2 mm. Corolla
yellowish green, narrowly campanulate, 5–8 mm, tomentose at throat, lobes ovate,
spreading or recurving, 2–2.5 mm. Filaments ca. 1.8 mm, anthers yellow, ovoid,
ca. 1 mm, minutely apiculate. Style exserted. Fruiting calyx becoming brown and
translucent, globose or ovoid, truncate at base, 1–2.2 cm, lobes short, somewhat
urceolate. Berry shiny, scarlet, globose, 5–8 mm. Seeds drying pale brown, reniform
discoid, 2–2.5 2 mm. (Wu et al. 1994–2013) (Figs. 1, 2, and 3).
Withania somnifera: Root decoction used against gonorrhea (Maasai), as child tonic
(Pokot) and for eye complaints (Kikuyu), for colds, skin rashes, excess bile, labor
pains and to improve health, for jaundice, rheumatism, and gonorrhea (Beentje 1994;
Gachati 1989; Kokwaro 2009; Lulekal et al. 2008; Muthee et al. 2011; Wondimu
et al. 2007). Pounded root sap for ulcers and stomachache. Heated leaves applied for
sore joints (Kokwaro 2009). Used to treat tumors (Yineger et al. 2007; Yineger and
Yewhalaw 2007). For amoebic diarrhea, stomachache, boils, abcesses, and swellings
(Teklehaymanot and Giday 2010). Sometimes used to treat excessive menstrual flow
(Njoroge and Bussmann 2009), as antimalarial (Njoroge and Bussmann 2006a), also
for asthma and common cold (Njoroge and Bussmann 2006b). Sometimes used as
Fig. 1 Withania coagulans

(Solanaceae), Pakistan. (Photo
W. Hussain)
Withania somnifera (L.) Dunal 1095

(Solanaceae), Pakistan.
(H. Ahmad Jan)
abortifacient, for tuberculosis, smallpox, measles, skin diseases, rheumatism, cardiac

problems and as anti-fungal (Mohagheghzadeh and Faridi 2006). Applied some-
times as diuretic (Mekonnen et al. 2010, 2015). Roots are smashed or chewed like a
carrot to treat stomach problems (Bussmann et al. 2011). For chest pain and typhoid
(Giday et al. 2003, 2010). Applied to treat malaria, stomachache, nightmares,
hyperglycemia, asthma, irregular menstruation, breast cancer, and wounds (Umair
et al. 2019). Used also to treat infertility, as tonic and aphrodisiac, for general debility
and rheumatic swellings (Sher et al. 2016). Serves as coagulant, for colds and flu,
and postpartum pains (Njoroge et al. 2004). Used for constipation, together with
Withania coagulans (Ali et al. 2019; Ahmad Jan et al. 2019).
Withania coagulans is used to treat diabetes (Ullah et al. 2019) and for liver
complaints (Muhammad et al. 2019).
Withania somnifera: The leaves are burnt as incense as spiritual medicine. When the
yoke hurts the back of the oxen and produces a swelling, crush the leaves and put on
the swelling (Bussmann et al. 2011). To expel evil spirits affecting livestock
(Yineger et al. 2007). Also used to treat “devils’ disease” in babies (Teklehaymanot

(Solanaceae), Pakistan.
(H. Ahmad Jan)
et al. 2007). FTo treat evil eye (Giday et al. 2003, 2010). Withania coagulans is used
as rennet replacement for making cheese (Ullah et al. 2019).
References
medicine research, prospects and limitations in Pakistan: a literature review. Acta Ecol Sin 2019.
https://doi.org/10.1016/j.chnaes.2019.12.005.
org/10.32859/era.18.24.1-19.
Withania somnifera (L.) Dunal 1097

10.1186/1746-4269-4-10.
Mekonnen T, Urga K, Engidawork E. Evaluation of the diuretic and analgesic activities of the
rhizomes of Rumex abyssinicus Jacq in mice. J Ethnopharmacol. 2010;127:433–9.
2009;82:262–9.
2019;188. https://doi.org/10.32859/era.18.8.1-20.
10.1186/s13002-019-0285-4.
District, Jimma zone, southwestern Ethiopia. J Ethnobiol Ethnomed. 2007;3:24.
Ximenia americana L.
XIMENIACEAE

Grace N. Njoroge
Synonyms
Ximenia americana L.: Amyris arborescens P. Browne; Heymassoli inermis Aubl.;

Heymassoli spinosa Aubl.; Primecaria odorata Raf.; Ximenia aculeata Crantz;
Ximenia americana fo. inermis (Aubl.) Engl.; Ximenia americana fo. inermis
S. Moore; Ximenia arborescens Tussac ex Walp.; Ximenia elliptica G. Forst.;
Ximenia fluminensis M. Roem.; Ximenia inermis L.; Ximenia laurina Delile;
Ximenia loranthifolia Span.; Ximenia montana Macfad.; Ximenia multiflora Jacq.;
Ximenia oblonga Lam. ex Hemsl.; Ximenia spinosa Salisb.; Ximenia verrucosa
M. Roem.; Ziziphus littorea Teijsm. ex Hassk.
Local Names
Ximenia americana: Swahili / Digo / Giriama: Mtundakula; Bajun: Mchunda-

Kula; Ilwana: Huda-Hudo; Kamba: Kitula, Mutula; Kikuyu: Mutura; Luo:
Olemo; Maa: Ol-Amai, Engamai; Marakwet: Timyotwa; Nyankore: Omuseka;
R. W. Bussmann (*)
G. N. Njoroge

Oromo: Huda-Hodo-Baddah; Pokot: Kinyotwo; Samburu: Lamaa, Lamai; Sebei:

Munyotwo, Mutenywa; Tugut: Myengwo; Turkana: Elamai; Borana: Dahbabes,
Huda; Somali: Madarau, Madarud; English: Sour plum, Tallow nut (Beentje 1994;
Botany and Ecology
Small tree or shrub up to 7 m tall, usually armed with axillary spines; bark greyish-
brown to black; young stems never tomentose. Petiole up to 6 mm long, canaliculate,
usually pubescent on the inner surface; leaf-blade coriaceous, glabrous, lanceolate to
oblong-lanceolate or elliptic, 3–8 cm long, 1.5–4 cm wide, apex obtuse to emargin-
ate, base usually cuneate; midrib prominent beneath, impressed above; lateral veins
3–7 pairs, inconspicuous. Inflorescence a shortly pedunculate raceme or umbel.
Flowers white to greenish, fragrant; pedicels 3–7 mm long, subequal to or shorter
than the newly opened flowers; both peduncles and pedicels glabrous. Calyx small,
with 4( 5) deltoid lobes, slightly ciliate at the margin. Petals 4( 5), 5–10 mm long,
1–2.5 mm wide, densely pilose on their inner surface, apiculate, recurved when
mature. Stamens 8; filaments 2–4 mm long; anthers 2–4 mm long and up to 0.8 mm
wide. Ovary superior, 3–4-partite almost to its apex; style up to 3 mm long. Fruit a
globose to ellipsoidal drupe, 2–2.5 cm in diameter, glabrous, yellow (or rarely
orange-red) at maturity; mesocarp pulpy. Seed woody, up to 1.5 cm long and
1.2 cm wide.
In South America used to treat stomach problems, nervous system problems and to
regulate menstruation (Bussmann and Sharon 2006, 2015a, b). The fruits can be used
as vermifuge, for tonsillitis, and mouth sores (Teklehaymanot and Giday 2007). A
seed paste is applied to cracked feet. The filtered extract of leaves is used to treat
trachoma. A root decoction serves to treat syphilis, hookworm, and chest pain. Fresh
leaves chewed for stomach pain. The bark decoction is used for venereal diseases
and diarrhea, as well as impotence and sterility (Kokwaro 2009; Lulekal et al. 2008).
Also for uterine bleeding (Muthee et al. 2011) and gastrointestinal problems
(Nankaya et al. 2019). Used for external injuries, eye problems, and flu
(Teklehaymanot et al. 2010). Commonly used in Ethiopia (Wondimu et al. 2007),
as well as in Latin America to treat nerves, stomach problems, and for menstrual
regulation (Bussmann and Sharon 2006; Bussmann and Glenn 2010). It is often part
of herbal mixtures (Bussmann et al. 2010a). The antibacterial properties of the
species have been shown in a variety of studies (Bussmann et al. 2009, 2010b, c,
2011; Geyid et al. 2005).
Ximenia americana L. 1101
Local Food Uses
Fruits edible (Balemie and Kebebew 2006; Beentje 1994; Gachati 1989; Kokwaro
2009).
Has hard, scented timber (Beentje 1994; Gachati 1989; Kokwaro 2009). Ponded
leaves are applied for conjunctivitis in cattle (Kokwaro 2009). In veterinary medi-
cine used against rabies (Teklehaymanot 2009).
References
Bussmann RW, Glenn A. Medicinal plants used in Northern Peru for reproductive problems and
female health. J Ethnobiol Ethnomed. 2010;6:30.
Bussmann RW, Sharon D, Diaz D, Cardenas R, Chait G, Castro M, Regalado S, Del Toro CR,
Malca GG, Perez AF, Glenn A. Antibacterial activity of medicinal plant species in Northern
Peru. Arnaldoa. 2009;16(1):93–103.
Bussmann RW, Glenn A, Meyer K, Rothrock A, Townesmith A. Herbal mixtures in traditional
medicine in northern Peru. J Ethnobiol Ethnomed. 2010a;6:10.
Bussmann RW, Glenn A, Sharon D. Antibacterial activity of medicinal plants of northern Peru – can
traditional applications provide leads for modern science? Indian J Tradit Med. 2010b;9(4):742–53.
Bussmann RW, Malca G, Glenn A, Sharon D, Chait G, Díaz D, Pourmand K, Jonat B, Somogy S,
Guardado G, Aguirre C, Meyer K, Rothrock A, Townesmith A, Effio-Carbajal J, Frías-
Fernandez F, Benito M. Minimum inhibitory concentration of medicinal plants used in Northern
Peru as antibacterial remedies. J Ethnopharmacol. 2010c;132:101–8.
Guardado G, Aguirre C, Meyer K, Rothrock A, Townesmith A. Proving that traditional
knowledge works – the antibacterial activity of Northern Peruvian medicinal plants. Ethnobot
Res Appl. 2011;9:67–96.
medicinal flora of Northern Peru. William L. Brown Center, MBG, St. Louis. 2015a. (ISBN
978-0-9960231-2-2).
medicinal del Norte de Peru. William L. Brown Center, MBG, St. Louis. 2015b. (ISBN 978-0-
9960231-3-9).
doi.org/10.1186/1746-4269-5-34.
4269-4-10.
1746-4269-3-12.
Zanthoxylum chalybeum Engl.
Zanthoxylum gilletii (De Wild) Waterm.
Zanthoxylum usambarense (Engl.) Kokwaro
RUTACEAE

Grace N. Njoroge
Synonyms
Zanthoxylum gilletii (De Wild) Waterm.: Zanthoxylum macrophyllum Oliv.
Local Names
Zanthoxylum chalybeum: Maa: Cloisuki; Ndorobo: Loisuki; Sonjo: Mdongo;

Shambaa: Mfuakumbi; Hehe: Mulungulungu; Digo: Mundungu, Ndumgu; Swa-
hili: Majafari; Borana: Gadah; Gabbra: Gaddaa; Giriama: Mdungu; Kamba:
Mukenea; Luo: Roko; Marakwet: Songoiywa, Songorurwa; Pokot: Loisugi,
Songowowo; Turgut: Koktin, Kokin (Beentje 1994; Gachati 1989; Kokwaro 2009)
Zanthoxylum gillettii: Kikuyu: Mucagatha; Kipsigis / Nandi: Sagawoita; Luhya:
Shihkoma, Shikhuma; Sukuma: Mfwakumbi; Luganda: Munyenye; Ndorobo:
Kikomit; English: African satinwood (Beentje 1994; Gachati 1989; Kokwaro 2009)
R. W. Bussmann (*)
G. N. Njoroge

Zanthoxylum usambarense: Kikuyu: Mugucua, Muguchwa; Kamba: Muvu,

Mulasi; Kipsigis: Sagawaita; Maa: Ol-Oisugi; Samburu: Loisugi (Beentje 1994;
Botany and Ecology
Zanthoxylum chalybeum Engl.: A deciduous shrub or tree to 10 m high, stem armed

with large woody spines, branches glabrous, with terminal buds protected by dark
scales, armed with black or reddish recurved usually paired prickles 5–17 mm long.
Female flowers subsessile, staminodes reduced to the aborted anthers, ovary
1.1–1.6 mm long. Fruit an oblique subglobose follicle, 5–8 mm in diameter,
glandular-bullate, stipe up to 1.5 mm long. Seed black and shiny, 5–7 mm in diameter.
Male flowers: pedicels slender, 1.5–2 mm long, sepals united at the base, 0.4 mm long,
petals elliptic, 2.5 mm long, 1 mm broad, stamens 4–5, filaments as long as the petals,
gynophore very short, vestigial ovary ellipsoid, 1.5 mm long. Flowers usually clus-
tered, yellowish green, 4–5-merous. Inflorescence of racemes or panicles up to 9 cm
long, borne at the base of the new branches below the first leaves (rarely in the axils).
Leaves 6–22 cm long, glabrous to densely hairy, rhachis armed below with a few small
scattered prickles, subterete, slightly grooved above, leaflets (1–)2–5-paired, opposite
to subopposite, sessile or the terminal one on a petiolule to 15 mm, ovate-oblong to
elliptic, 2.5–9.5 cm long, (1–)2–4.3 cm broad, acute to obtuse at the apex, cuneate at
the base, slightly crenulate to subentire, chartaceous to subchartaceous, sparsely dotted
with pellucid glands, lateral nerves 6–9 pairs (Kokwaro 1982) (Figs. 1 and 2).
Zanthoxylum gilletii (De Wild) Waterm.: Deciduous tree 10–35 m high, trunk
usually straight and without branches for several m., 30–90 cm in diameter at the
base, armed with conical woody prickle-bearing straight protuberances 1–3 cm long,
bark grey and smooth, branches glabrous, armed with conical reddish straight or
slightly recurved prickles 2–8 mm long. Leaves in terminal clusters, closely
Fig. 1 Zanthoxylum
armatum (Rutaceae), plant
with ripe fruits in Dadeldhura
district, Nepal. (Photo
R.M. Kunwar)
Zanthoxylum chalybeum Engl. . . . 1105
Fig. 2 Zanthoxylum
armatum (Rutaceae), plant
with ripe fruits in Dadeldhura
district, Nepal. (Photo
R.M. Kunwar)
alternate, 25–120( 150) cm long, rhachis terete at the base, flattened above, some-
times with straight sharp prickles 2–7 mm long, glabrous, petiolules 5–10 mm long,
leaflets 13–27, alternate or subopposite, elliptic-oblong to elliptic, (8–)14–30 cm long,
3.5–10 cm broad, attenuate or abruptly acuminate to the obtuse apex, asymmetrical
and cuneate or rounded at the base (sometimes very asymmetrical), entire or slightly
crenulate, coriaceous, glabrous, gland-dots numerous but small and almost inconspic-
uous, midrib sometimes with sparse prickles, prominent beneath, lateral nerves 8–14
pairs. Sepals 5, united halfway, ovate to subcircular, 0.5–0.8 mm long. Inflorescence of
terminal and axillary pyramidal panicles, 20–34 cm long, sometimes aculeate at the
base, flowers clustered, sessile or very shortly pedicellate. Male flowers: stamens 5, all
varying in length, filaments 1.5–2.8 mm long, disk dome-shaped and lobed, vestigial
ovary obsoletely lobed. Petals pandurate to obovate, 1–2.5 mm long, 1–1.5 mm broad,
white, turning brown. Fruit reddish, subglobose, 3.5–6 mm in diameter, glandular-
foveolate, sessile or with a short stipe and a persistent calyx. Female flowers:
staminodes 5, reduced to aborted anthers, gynophore 0.2–0.3 mm long, ovary ovoid,
1–1.4 mm long, foveolate (Kokwaro 1982) (Figs. 3, 4, and 5).
Zanthoxylum usambarense (Engl.) Kokwaro: A much-branched tree 8–14( 16) m
high, trunk without branches for several m, 22–35 cm in diameter at the base, armed
with conical woody prickle-bearing outgrowths 2–3 cm long ending in sharp straight
thorns 5–9 mm long, bark conspicuously rough, with longitudinal ridges and
furrows (as deep as 5 cm), greyish brown but peeling yellow underneath, branches
with sharp straight or slightly upcurved dark red prickles 6–12 mm long, roots with
yellow bark which (together with the bark) are heavily exploited for medicinal
purposes. Leaves 10–24 cm long, rhachis unarmed or with small straight or
upcurved prickles, petiolule (0–)0.5–2 mm long except on the terminal leaflets
where it is up to 5 mm, leaflets 2–8 pairs, usually subopposite to opposite, elliptic
to elliptic-oblong, 2.2–8 cm long, 1.5–3 cm broad, acuminate or apiculate to almost
obtuse at the apex, obtuse and slightly oblique to cuneate at the base, crenate, with
distinct pellucid glands along the margin, coriaceous, glabrous, midrib occasionally
aculeate beneath, lateral nerves 8–16 pairs. Inflorescence terminal much-branched
corymbose panicles, 10–15 cm long, 7–19 cm broad, glabrous, with cymose clusters
Fig. 3 Zanthoxylum
armatum (Rutaceae),
Pakistan. (Photo H., Ahmad
Jan)
Fig. 4 Zanthoxylum
armatum (Rutaceae),
Jan)
of flowers on pedicels 0–3 mm long. Sepals 4, united at least at the base, ovate to
elliptic, 1–1.5 mm long, 1 mm broad. Petals 4, elliptic to elliptic-oblong, 2–4.5 mm
long, 1.5–2.2 mm broad, apiculate or obtuse, white to cream or greenish yellow.
Male flowers: stamens 4, filaments 2–6 mm long and variable in length in the same
flower, anthers 1–2 mm long, gynophore short, glabrous, vestigial pistils 2, up to
1.4 mm long. Female flowers: staminodes absent, gynophore short, ovaries 2, par-
tially united, 1–1.5 mm long, styles 2, 0.8–1 mm long, stigmas 2, often partially
united. Fruit of paired subglobose follicles 8–10 mm long, 6–7.5 mm in diameter,
mucronate, usually on a common stipe but individually sessile, one of the pair
sometimes rudimentary, reddish green, glandular-foveolate, Seeds 5–7 mm long,
4.5–5.5 mm in diameter, dark blue-black and glossy (Kokwaro 1982). In dry,
Elaeodendron and Brachylaena dominated Afromontane forests (Bussmann 2002).
Fig. 5 Zanthoxylum
armatum (Rutaceae),
Jan)
Zanthoxylum chalybeum: Bark used as emetic, to treat cough, sore throat, and
malaria (Beentje 1994). A decoction from boiled leaves as well as root bark is used
to treat kwashiorkor edema. The boiled root decoction is drunk for the chest pain and
body pain, malaria, colds, and dizziness. The root is chewed for toothache. In milk
the bark is given to children for better appetite (Maasai, Sonyo). The fruits are
chewed for sore throat and chest pain (Kokwaro 2009).
Zanthoxylum gilletii: Bark decoction to treat cough (Beentje 1994). The bark is
chewed for stomach problems, and for toothache. A bark decoction serves to remedy
joint pains, fever, rheumatism, venereal diseases, and is applied to wounds
(Kokwaro 2009).
Zanthoxylum usambarense: Bark decoction used as emetic and against malaria, for
cough, and rheumatism. The fruits in milk are used to treat fever, sore throat,
tonsillitis, cough, and chest pains (Beentje 1994; Gachati 1989; Kokwaro 2009).
Used for throat infections (Delbanco et al. 2017). Used also as antimalarial (Njoroge
and Bussmann 2006), and for postpartum weakness (Njoroge and Bussmann 2007).
Employed to treat rheumatic pain and joint pain (Nankaya et al. 2019).
Zanthoxylum alatum is used for hypertension (Ahmad Jan et al. 2019) and has
antidiabetic properties (Ullah et al. 2019).
Zanthoxylum armatum is used as mouthwash and for toothache (Bhat et al. 2013;
Kunwar et al. 2013, 2015, 2019; Malik et al. 2015; Singh et al. 2017, 2019).
Essential oil from the fruit has deodorant and antiseptic properties. The plant is
used in toothache, headache, skin diseases, indigestion, and diarrhea. It is employed
as a tonic, analgesic, carminative, expectorant, diuretic and used in gastritis, liver
problems, blood disorders in unani system (Kunwar and Bussmann 2009; Kunwar
et al. 2009). It is also used for epilepsy and to treat cough (Sher et al. 2016).
Zanthoxylum capense is used for mood disorders, and Zanthoxylum

zanthoxyloides as febrifuge (Mohagheghzadeh and Faridi 2006). Zanthoxyluim
tsihanimposa is used for postpartum recovery (Randrianarivony et al. 2016a, b).
Local Food Uses
The fruits of Zanthoxylum armatum are used as spice.
Zanthoxylum chalybeum: Bark decoction to treat sick camels (Beentje 1994). The
root and bark decoction is used to cure diarrhea and pneumonia in goats, and
septicemia and helminthiasis in livestock (Balemie and Kebebew 2006; Kokwaro
2009).
Zanthoxylum gilletii: Used for boats and carpentry (Beentje 1994). Sometimes used
as firewood (Kiefer and Bussmann 2008). Zanthoxylum angolense, Zanthoxylum
gilletii, Zanthoxylum heitzii, Zanthoxylum lemairei, Zanthoxylum leprieurii,
Zanthoxylum viridis, and Zanthoxylum xanthoxyloides are used as fishing poisons
(Neuwinger 2004).
Zanthoxylum usambarense: The wood is used for construction, furniture, and bows
(Beentje 1994).
Zanthoxylum armatum is an important medicinal plant for export in Nepal (Kunwar
et al. 2006, 2008, 2018).
References
2019. https://doi.org/10.1093/ecam/neh088.
Bussmann RW. Islands in the desert – forest vege.tation of Kenya’s smaller mountains and highland
Delbanco AS, Burgess ND, Cuni-Sanchez A. Medicinal plant trade in Northern Kenya: economic
Biodiversität, Naturausstattung im Himalaya, vol III. Erfurt; 2009. p. 475–89
study from Dolpa, Humla, Jumla and mustang districts of Nepal. J Ethnobiol Ethnomed.
2006;2:27.
Kunwar RM, Chowdhary CL, Bussmann RW. Diversity, utilization and management of medicinal
medicines in Farwest Nepal. J Ethnopharmacol. 2015;163:210–9.
P. Cross-cultural comparison of plant use knowledge in Baitadi and Darchula Districts, Nepal
Kunwar RM, Shrestha K, Malla S, Acharya T, Sementelli A, Kutal D. Relation of medicinal plants,
their use patterns and availability in the lower Kailash Sacred Landscape, Nepal. Ethnobot Res
Appl 2019;187. https://doi.org/10.32859/era.18.6.1-14.
Nankaya J, Gichuki N, Lukhoba C, Balslev H. Sustainability of the Loita Maasai children’s
Ethnomed. 2006;2:8.
Randrianarivony TN, Randrianasolo A, Andriamihajarivo T, Ramarosandratana AV, Jeannoda VH,
2017;13:49. https://doi.org/10.1186/s13002-017-0178-3.
https://doi.org/10.32859/era.18.14.1-11.
2019;188. https://doi.org/10.32859/era.18.8.1-20.
Zehneria scabra (L.f.) Sond.
CUCURBITACEAE

Grace N. Njoroge
Local Names
Zehneria scabra: Kikuyu: Rwegethia; Shambaa: Fsuiza (Beentje 1994; Gachati

1989; Kokwaro 2009).
Botany and Ecology
Zehneria scabra (L.f.) Sond.: Perennial herb climbing or trailing to 6 m, young

stems rather shortly and closely crisped-pubescent, sometimes densely so, but
sometimes almost glabrous except at the nodes, less often sparsely spreading-hairy
or completely glabrous, older stems thickened, woody, up to 15 mm across, with
greyish-brown slightly fissured bark. Male flowers 2–60 in sessile or up to 85 mm
pedunculate umbelliform clusters, often with a coaxillary long-pedicellate solitary
flower, rarely the inflorescence compound with more than one umbelliform cluster
borne at intervals along the peduncle, bracts usually absent, receptacle-tube
R. W. Bussmann (*)
G. N. Njoroge

2,5–4 mm long, hairy within, lobes more or less subulate, 0,5–0,8 mm long, petals
white, becoming creamy-yellow when old, 1.8–3 mm long, 1.3–2 mm broad,
stamens 3, on fairly long filaments. Groups, often with a coaxillary solitary flower,
pedicels 1–10 mm. long, ovary sub-globose to fusiform, sometimes beaked, 3–6 mm
long, 1.5–2.3 mm across, receptacle-tube campanulate, hairy within, 2–4 mm long,
lobes subulate, more or less 0.5 mm long, petals 2–2.5 mm long, 1–1.5 mm broad.
Leaf-blade more or less ovate-cordate or ovate-subtruncate in outline, more or less
scabrid-punctate above, paler green and subglabrous or sparsely scabrid-hairy to
densely cinereous-tomentose beneath, sinuate-toothed, acute to rounded, apiculate
and often more or less acuminate, 25–110 mm long, 20–110 mm broad, sometimes
shallowly 3-lobed with large middle lobe and small triangular lateral lobes, rarely
palmately 5-lobed with the lobes triangular, or obovate and distinctly narrowed
below, sometimes heterophyllous, petiole usually rather shortly and sparsely hairy,
7–85 mm long. Dioecious. Fruits solitary or more often in sessile or pedunculate
axillary clusters of up to 10 (rarely more), with 2–20 mm long pedicels, baccate,
sphaeroid, rounded or more rarely slightly apiculate, 6–12 mm long, 5–11 mm
across, scarlet, more or less scrobiculate when dry. Seeds ovate in outline, smooth,
compressed, lenticular, 2.2–4 1,3–2.5 0.5–0.8 mm (Jeffrey 1967). Commonly
found in drier Afromontane forests (Bussmann 2002) (Figs. 1, 2, and 3).
Root used for abdominal pains. Burned plant parts applied to burns, and a wash from
pounded leaves for skin rashes (Kokwaro 2009). The root decoction is used to treat
malaria (Giday et al. 2010). Serves also as febrifuge and for gynecological problems
(Mohagheghzadeh and Faridi 2006), and for nose, ear, and throat diseases (Njoroge
and Bussmann 2006). The crushed leaves are applied to swellings (Teklehaymanot
et al. 2007), and the steam is inhaled for febrile illnesses (Teklehaymant 2009),
anemia, and malaria (Gedif and Hahn 2003; Giday et al. 2007; Muthee et al. 2011),
and to treat diarrhea (Gedif and Hahn 2003), as well as flu and hypertension
(Njoroge et al. 2004).
Local Food Uses
The leaf juice is drunk (Gedif and Hahn 2003).
Wash from pounded leaves to treat calves for lice (Kokwaro 2009). Browsed by
livestock (Bussmann et al. 2006, 2011), and formerly burnt and mixed with charcoal
to make ink (Bussmann et al. 2011). Applied nasally for veterinary purposes
Zehneria scabra (L.f.) Sond. 1113
Fig. 1 Zehneria scabra

(Cucurbitaceae), Mt. Kenya
R.W. Bussmann)

R.W. Bussmann)

R.W. Bussmann)
(Yineger and Yewhalaw 2007). Leaves used as soap. When giving birth, the deliv-
ering mother’s body is washed with this plant (Luizza et al. 2013).
References
Res Appl. 2013;11:315–39.
Zehneria scabra (L.f.) Sond. 1115
Ziziphus jujuba Mill.
Ziziphus mauritiana Lam.
Ziziphus mucronata Willd.
RHAMNACEAE

Grace N. Njoroge, Maroof Ali Turi, and Hammad Ahmad Jan
Synonyms
Ziziphus jujuba Mill.: Ziziphus sativa Gaertn; Ziziphus vulgaris Lam, Rhamnus
zizyphus L.
Ziziphus mauritiana Lam.: Paliurus mairei H. Lév.; Rhamnus jujuba L.; Ziziphus
abyssinica Hochst.; Ziziphus jujuba (L.) Gaertn.; Ziziphus jujuba (L.) Lam.;
Ziziphus mairei (H. Lév.) Browicz & Lauerer; Ziziphus muratiana Maire; Ziziphus
orthacantha DC.; Ziziphus rotundata DC.; Ziziphus tomentosa Poir.
Ziziphus mucronata Willd.: Ziziphus adelensis Delile; Ziziphus baclei DC.;
Ziziphus bubalina Licht. ex Schult.; Ziziphus madecassus H. Perrier; Ziziphus
mitis A. Rich.
R. W. Bussmann (*)
G. N. Njoroge
M. A. Turi
College of Life Science, Anhui Normal University, Wuhu, China
H. A. Jan
Islamia College Peshawar Jamrod Road, University Campus Peshawar, Peshawar, Pakistan

Local Names
Ziziphus abyssinica: Kamba: Kiae, Muae; Lugbara: Lu; Hehe: Mutanula; Acholi:
Olango (Kokwaro 2009).
Ziziphus mauritiana: Amharic: Kurkura; Borana: Qurgura; Marakwet: Tilomwa,
Yilomwa; Pokot: Tlomwo, Tlomwa; Rendile: Qasa; Somali: Geb, Gub; Tugut:
Tilolwo; Turkana: Enkalale; Swahili: Mkunazi; Acholi: Lango; Hehe: Mtyanula;
English: Dunks, Jujube, Indian cherry, Indian jujube, Indian plum, Chinese date,
Chinese apple, Bear tree, Desert apple (Beentje 1994; Gachati 1989; Kokwaro
2009).
Ziziphus mucronata: Digo: Mungugune; Girama: Mguguna; Kipsigit:
Mimanbiliot; Luo: Lang’o, Longo; Maa: Ol-Oilailke; Marakwet: Nonoiywa;
Pokot: Tirokwo; Samburu: Loilalei; Tugut: Ninoiwa; Noiwa; Turkana: Esilang,
Amaleri; Sukuma: Ilegero; Swahili: Mkunazi; Sandawi: Tsindimak’o; English:
Buffalo thorn, Cape thorn. (Beentje 1994; Gachati 1989; Kokwaro 2009).
Botany and Ecology
Ziziphus jujuba: Shrub or mall tree, 3–8 m tall. Spreading-branching spinose plants,
branches angularly flexuose, glabrous, red-brown, with paired large acute prickles
up to 3 cm long, and with thin erect greenish biseriately, leafy shoots (1–3 on each)
like a compound leaf; leaves coriaceous, glabrous, dark green above, shiny,
obliquely oblong- ovate to broadly lanceolate usually obtuse, rounded or slightly
cordate at base, obtusely and finely dentate, short -petiolate or subsessile, with small
stipules at base. Flowers stellate, 3–4 mm in diameter, in dense glomerules on very
short peduncles, peduncles, sepals 5, ovate-triangular; petals 5, greenish-yellow,
obovate, clawed at base. Fruit globulose, 1–1.5 cm long (in cultivated forms up to
3 cm long, often oblong-oval), to dark cinnamon, reddish-orange to red-purple shiny.
Flowering from. June–July, fruiting from August. Caucasus, Middle Asia, on dry,
gravelly, stony slopes, in open forests, up to 1500 m. Uncommon, occurs in small
populations and as solitary individuals (Shishkin and Boborov 1949) (Figs. 1, 2, 3, 4,
5, 6, and 7).
Ziziphus mauritiana Lam.: Spiny, evergreen shrub or small tree up to 3–15 m tall,
with trunk 40 cm or more in diameter. Crown spreading; branches with stipular
spines, drooping. Bark dark grey or dull black, irregularly fissured. Leaves variable,
alternate, in 2 rows, oblong-elliptic, 2.5–6 1.5–5 cm, with tip rounded or slightly
notched base; finely wavy-toothed on edges, shiny green and hairless above; dense,
whitish, soft hairs underneath. Inflorescence axillary cymes, 1–2 cm long, with 7–20
flowers; peduncles 2–3 mm long; flowers 2–3 mm across, greenish-yellow, faintly
fragrant; pedicels 3–8 mm long; calyx with 5 deltoid lobes, hairy outside, glabrous
within; petals 5, sub-spathulate, concave, reflexed. Fruit a drupe, globose to ovoid,
up to 6 4 cm in cultivation, usually much smaller when wild; skin smooth or
rough, glossy, thin but tough, yellowish to reddish or blackish; flesh white, crisp,
juicy, acidic to sweet, becoming mealy in fully ripe fruits. Seed a tuberculate and
Ziziphus jujuba Mill. . . . 1119
Fig. 1 Ziziphus jujuba

(Rhamnaceae), Lagodekhi,
Georgia. (Photo R. W.
Bussmann and N. Y.
Paniagua-Zambrana)
irregularly furrowed stone, containing 1–2 elliptic brown kernels each 6 mm long
(Johnston 1972) (Fig. 8).
Ziziphus mucronata Willd.: Shrubs or small trees to 15(–30) m tall, armed with
spinous stipules. Bark of bole dark grey, smooth to rough to corrugated; slash-wood
crimson, soft. Branches often zig-zag. Leaf-blades ovate, (2–)3–6(–8) cm long,
(1.3–)2–3.5(–4.7) cm wide, at base often markedly asymmetrical, rounded or rarely
very shallowly cordate, acute, often acuminate or rarely obtuse, serrulate, 3-nerved
from base, the nerves of the dull upper surface only slightly if at all impressed;
petioles 2–7 mm long. Cymes (5–)10–15 mm long, and about as thick, (3–)7–25
flowered; peduncles 1–3 mm. long; pedicels 1–3 mm. long in flower, 3–5 mm long
in fruit. Sepals 1.5–2 mm. long, spreading in flower. Petals 1–1.5 mm long, spread-
ing in flower. Ovary 2-celled, stigmas 2. Fruit globose, 12–20 mm thick, reddish or
reddish brown at maturity (Johnston 1972).
Ziziphus abyssinica: Root decoction used as abortificant. Dried and powdered root
rubbed in chest incision to treat pneumonia. Root decoction also to treat asthma,
venereal diseases, stomachache, and snake bites. Burnt leaves for tonsilitis
(Kokwaro 2009).

Bussmann and N. Y.
Paniagua-Zambrana)
Ziziohys jujuba: Up to the end of the nineteenth century it was highly valued in
Europe in medicine where the fruits (Baccae jujubae, jujube, Brustbeere, breast
berry) and preparations made from them were used as emollients in the treatment
of throat and catarrhal infections (Sokolov 1988). In Middle Asia used as a laxative,
sudorific, and anti-inflammatory, hypotensive, expectorant, emollient, anti-
asthmatic, as hemostatic, and cystitis. Topically the extract is used for stomatitis.
The fruits are used as sedative. The fruits are used for catarrh of the upper airways,
fevers, and to treat intestinal infections. The root bark is used as a stimulant and the
fruits have antibacterial action. In Middle Asia, a decoction of the fruit is used for
anemia, chest pains, asthma, coughs, smallpox, diarrhea, and as an analgesic for
diseases of the liver, kidneys, and intestines and also as hypotensive (Sokolov 1988).
Has antibacterial properties (Ahmad Jan et al. 2019).
Ziziphus mauritiana: Leaves, fruits, and bark are used medicinally. Pounded roots
are added to drinking water and given to poultry suffering from diarrhea and to
humans for indigestion. The root decoction is used as abortificant, the infusion for
dysentery, tuberculosis, and indigestion. Dried and powdered root rubbed in chest
incision to treat pneumonia. Root decoction also as prophylactioc to avoid swollen
legs in menstruating women (Kokwaro 2009). Serves to treat chicken pox, ulcers,
diarrhea, asthma, toothaches, and jaundice (Umair et al. 2019). In India, the species
is used to remedy blood purifier, cholera, colic, diarrhea, digestion, dysentery, eye
diseases (conjunctivitis), hair loss, fever, headache, edema, rheumatism, scorpion

Bussmann and N. Y.
Paniagua-Zambrana)

Bussmann and N. Y.
Paniagua-Zambrana)
sting, sores, spleen complaints, stomach ache, ulcer, wound, and whooping cough
(Verma et al. 2007). In Nepal it is used as Root extract is antimycobacterial,
anticonceptive, analgesic, and antidiabetic (Kunwar et al. 2009).
Ziziphus mucronata Roots are used as a remedy for pain and to treat snakebite.
Boils and other skin infections are treated with leaf paste, and this, together with an
infusion of the roots, is a treatment for tubercular gland swellings, measles,

(Rhamnaceae), market,
Tbilisi, Georgia. (Photo R. W.
Bussmann and N. Y.
Paniagua-Zambrana)

(Rhamnaceae), flowering
branch. (Photo Maroof Ali
Turi)

(Rhamnaceae), flowers,
Pakistan. (Photo Maroof Ali
Turi)
Fig. 8 Ziziphus mauritiana

(Rhamnaceae), Pakistan.
(Photo Hamand Ahmad Jan)
dysentery, lumbago and chest complaints. The bark is used as an emetic; bark
decoction is used for rheumatism and stomach troubles, bark infusion is used to
treat coughs, and bark is used in a steam bath to purify the complexion. The bark
decoction is used for snakebites, rheumatism, and stomach problems. Pounded
leaves and roots are applied as poultice to cure boils and skin infections (Kokwaro
2009).
Ziziphus nummularia is used in Pakistan as tonic, for hyperglycemia, constipation,
throat problems, colds, and scabies (Umair et al. 2019), and in India, the species is
used to remedy boils, colds, diarrhea, gum inflammation, and tonsilitis (Verma et al.
2007).
Local Food Uses
Ziziphus jujuba: The fruits are eaten, mostly dried. The fruits contain sugar that is
both nutritious and tasty. Since ancient times the plant has been cultivated in gardens
throughout its entire distribution area, especially in the East where there are several
cultivated strains (Bussmann et al. 2016, 2018; Sokolov 1988).
Ziziphus mauritiana: Fruit is eaten fresh or dried and can be made into a floury
meal, butter, or a cheeselike paste, used as a condiment. Also used for candy making
and pickling. The fruit is a good source of carotene, vitamins A and C, and fatty oils.
A refreshing drink is prepared by macerating fruits in water. In Indonesia, young
leaves are cooked as a vegetable.
Ziziphus mucronata: The fruit is eaten fresh or dried, in meal or porridge. The
young leaves can be cooked and eaten as spinach. Seeds are roasted, crushed, and
used as a coffee substitute.
Ziziphus abyssinica: Root decoction given to livestock as anthelminthic (Kokwaro

2009).
Ziziphus jujuba: Used to tan leather and to dye wool and silk yellow. The wood is
used for tool handles. The leaves can be used to feed silkworms. Planted as
ornamental (Sokolov 1988).
Ziziphus mauritiana: In parts of North Africa, the leaves are used as fodder for
sheep and goats. Ziziphus mauritiana produces excellent firewood and good char-
coal. When in bloom it is occasionally a source of pollen. Ziziphus mauritiana yields
a medium-weight to heavy hardwood. The wood is used for general construction,
furniture and cabinet work, tool handles, agricultural implements, tent pegs, golf
clubs, gun stocks, sandals, yokes, harrows, toys, turnery, household utensils, bowl-
ing pins, baseball bats, chisels and packaging. It is also suitable for the production of
veneer and plywood. The bark, including the root bark, is used for tanning; it also
yields brown and grey or reddish dyes. Ziziphus mauritiana is also used to stupefy
fish in Ethiopia. A suitable species to aid in fixation of coastal dune sand. The tree is
planted for shade and windbreaks and is well suited gardens. In Pakistan, the species
is used to treat chicken-pox, jaundice, asthma, ulcers, diarrhea and toothache (Umair
et al. 2019). Ponded roots given to poultry against dysentery (Kokwaro 2009).
Ziziphus mucronata Both leaves and fruits are browsed and used as fodder. The
flowers are a source of honey. The wood is hard and makes good firewood and
charcoal. The bark contains 12–15% tannin. It is planted as a living fence.
References
2019; https://doi.org/10.1016/j.chnaes.2019.12.005.
Caucasus. J Ethnobiol Ethnomed. 2016;12:43. https://doi.org/10.1186/s13002-016-0110-2.
Johnston MC. Flora of tropical East Africa: Rhamnaceae. London: Crown Agents for Overseas
Shishkin BK, Boborov EG. Flora of the USSR, Volume 14: Geraniales, Sapindales, Rhamnales.
Leningrad: Akademia Nauk; 1949, 616 pages, 39 b/w plates, 2 maps. (English 1974).
Sokolov PD, editor. Plant resources of the USSR: flowering plants, their chemical composition, use,
Volume 4. Families of Rutaceae-Elaeagnaceae. Leningrad: Akademia Nauk; 1988,
10.1186/s13002-019-0285-4.

Ethnobotany of The Mountain Regions of Africa 2021

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Ethnobotany of The Mountain Regions of Africa 2021

Uploaded by

Copyright:

Available Formats

Ethnobotany of Mountain Regions

Rainer W. Bussmann Editor

More information about this series at http://www.springer.com/series/15885

With 666 Figures

ISSN 2523-7489 ISSN 2523-7497 (electronic)

Part I Regions ........................................... 1

Part II Plants ............................................ 67

Abutilon sp. Abutilon angulatum (Guill. & Perr.) Mast. Abutilon

Aerva lanata (L.) Juss. ex Schult. AMARANTHACEAE . . . . . . . . . . . . . . . . . 119

Blighia unijugata Bak. SAPINDACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221

Clutia abyssinica Jaub. & Spach. PERACEAE . . . . . . . . . . . . . . . . . . . . . . 305

Cyphostemma maranguense (Gilg.) Desc. VITACEAE . . . . . . . . . . . . . . . . 399

Eragrostis tef (Zuccagni) Trotter POACEAE . . . . . . . . . . . . . . . . . . . . . . . 489

Grewia bicolor Juss. Grewia tembensis Fresen. Grewia tenax

Harungana madagascariensis Poir. HYPERICACEAE . . . . . . . . . . . . . . . . . 591

Mangifera indica L. ANACARDIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691

Phytolacca dodecandra L’Hér. PHYTOLACCACEAE . . . . . . . . . . . . . . . . . . . 807

Piper capense L. PIPERACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 825

Pistacia aethiopica Kokwaro ANACARDIACEAE . . . . . . . . . . . . . . . . . . . . . 831

Podocarpus milanjianus Rendle Afrocarpus falcatus (Thunb.)

Portulaca oleracea L. PORTULACACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 845

Pouteria adolﬁ-friederici (Engl.) A. Meeuse Pouteria altissima

Prunus africana (Hook. f.) Kalkman ROSACEAE . . . . . . . . . . . . . . . . . . . 857

Psidium guajava L. MYRTACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 863

Psychotria orophila E.A.A. Petit Psychotria sp. RUBIACEAE . . . . . . . . . . . 869

Pterolobium stellatum (Forssk.) Brenan FABACEAE . . . . . . . . . . . . . . . . . 873

Rapanea melanophloeos (L.) Mez. PRIMULACEAE . . . . . . . . . . . . . . . . . . . 879

Rhamnus prinoides L’Hér. Rhamnus staddo A. Rich. RHAMNACEAE . . . . 885

Rhus natalensis Bernh. ex C. Krauss Rhus ruspolii Engl. Rhus

Ricinus communis L. EUPHORBIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 903

Rubus apetalus Poir. Rubus rosifolius Sm. Rubus volkensii Engl.

Tabernaemontana stapﬁana Brizten APOCYNACEAE . . . . . . . . . . . . . . . . . 1005

Vitex keniensis Turrill LAMIACEAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1085

Dr. Rainer W. Bussmann earned his M.Sc. (Diploma)

Dr. Bussmann is a past President of the Society for

Rainer W. Bussmann Department of Ethnobotany, Institute of Botany and

Rainer W. Bussmann, Narel Y. Paniagua-Zambrana, and

Geology, Landforms and Soil Conditions

Africa is an old continent. Nowhere else is so much of the surface covered by

© Springer Nature Switzerland AG 2021 3

Fig. 3 Simien Mountains landscape, Ethiopia. (Photo R.W. Bussmann)

Fig. 4 Bale Mountains, Southern Ethiopia. (Photo R.W. Bussmann)

Vegetation Belts in African Mountains

vegetation display strong similarities to Mediterranean mountains like the Sierra

The Moroccan Atlas

The Rif and the Tell Atlas

The Saharan Atlas in Northern Algeria

Dry Mountains in Central Sudan: The Jebel Marra

Mount Kenya: A Typical Example for the Altitudinal Zonation

Supratropical Evergreen Montane Forests

Fig. 6 Dense crowns of

Orotropical Montane Forest: Evergreen Broad-Leaved and Evergreen

Fig. 8 Dry Juniperus-Olea

Orotropical Bamboo Forest

Fig. 9 Sinarundinaria alpina

Fig. 10 Mixed bamboo-

Orotropical Cloud Forest

Altotropical Grasslands and Moorlands

Fig. 14 Dendrosenecio keniodendron on Mt. Kenya, Kenya. (Photo R.W. Bussmann)

Fig. 15 Lobelia telekii on

Mt. Kenya’s Neighbors: Mt. Kilimanjaro, Mt. Meru, Aberdare Mts.,

Although they are separated by hundreds of kilometers of lowlands, the neighboring

Fig. 16 Vegetation zonation on Mt. Kilimanjaro (Bussmann 2006)