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Plant Resources of Tropical Africa 7(I)

Timbers I

PROTA is an international Foundation involving the following participating institu-tLons:

Wageningen University GNU), Plant Sciences Group (PSG), Haarweg 333, P. 0.Box 341, 6700 AH Wageningen, NetherlandsAgropolislnternational(AGROPOLIS), Avenue Agropolis, F-34394 MontpellierCedex 5, FranceRoyal Botanic Gardens Kew (RBGKEW), Centrefor Economic Botany, Rich-mond, Surrey TW9 3AB, United KingdomCentre Nationalde Semences Foresti6res (CNSF), 01 B. P. 2682, 0uagadougou01, Burkina FasoCentre National de to Recherche Scientifique at Technologique (CENAREST),B. P. 842, Libreville, GabonForestry Research Institute of Ghana (FORTG), KNUST, University P. 0. Box63, Kumasi, GhanaParc Botanique at Zoologique de Tsimbazaza (PBZT), B. P. 4096, Tsimbazaza,Antananariv0 101, MadagascarNational Herbarium and Botanic GardensofMalawi(NHBGM), P. 0. Box 528,Zomba, MalawiMakerere University 011U), Department of Botany, P. 0. Box 7062, Kampala,UgandaWorld Agroforestry Centre (ICRAF), P. 0. Box 30677, Nairobi, KenyaProsea Foundation (PROSEA), P. 0. Box 332, Bogor 16/22, Indonesia

This publication has been made possible through the financial support by:Netherlands Ministry of Agriculture, Nature and Food QualityNetherlands Ministry of Foreign Affairs, Directorate-General for InternationalCooperation (DGIS)Netherlands Ministry of Spatial Planning, Housing and the EnvironmentNetherlands Organization for Scientific Research (NWO)Wageningen University, Netherlands

This work was carried out with the aid of a specific grant from:International Tropical Timber Organization (ITTO), Yokohama, Japan(Project PD 264104 Rev. 3 on, I) 2006-2008)

ITTO

,

Plant Resources of Tropical Africa 7(I)

Timbers I

Editors:

General editors:

D. LouppeA. A. Oteng-AmoakoM. Brink

R. H. M. J. Lemmens

L. P. A. OyenJR. Cobbinah

PROTA Foundation I Backhuys Publishers/ CTAWageningen, Netherlands, 2008

.

Correct citation of this publication:Louppe, D. , Oteng-Amoako, A. A. & Brink, M. (Editors), 2008. Plant Resources ofTropical Africa 7(I). Timbers I. PROTA Foundation, Wageningen, Netherlands IBackhuys Publishers, Leiden, Netherlands/ CTA, Wageningen, Netherlands. 704 pp.

Correct citation of articles from this publication:[Author name, initials, 2008. Title of article]. In: Louppe, D. , Oteng-Amoako, A. A. &Brink, M. (Editors). Plant Resources of Tropical Africa 7(I). Timbers I. PROTAFoundation, Wageningen, Netherlands/ Backhuys Publishers, Leiden, NetherlandsI CTA, Wageningen, Netherlands. pp. ...-....

IsBN 978-90-5782-209-4 I 978-3-8236-1541-5 (book only)IsBN 978-90-5782-210-0 I 978-3-8236-1542-2 (book + CD-Rom)

O PROTA Foundation, Wageningen, Netherlands, 2008.

No part of this publication, apart from bibliographic data and brief quotations eru-bodied in critical reviews, may be reproduced, re-recorded or published in any formIncluding print, photocopy, microfilm, electric or electromagnetic record withoutwritten permission from the copyright holder: PROTA Foundation, P. 0. Box 341,6700 AH Wageningen, Netherlands.

Printed in the Netherlands by Ponsen & Loonen bv, Wageningen.Distributed forthe PROTA Foundation by Backhuys Publishers, P. 0. Box 321, 2300AH Leiden, Netherlands (worldwide), and CTA, P. 0. Box 380, 6700 AJ Wageningen,Netherlands (ACP countries).

Contents

Contributors 6

PROTA Board of Trustees and Personnel 12

Introduction 14

Alphabetical treatment of timbers 19

Literature 602

Index of scientific plant names 687

Index ofvernacular plant names 697

PROTA in short 702

CTAin short 703

Map of Tropical Africa for PROTA 704

5

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6 TIMBERSl

Contributors

K. A. Adam, Forestry Research Institute of Ghana (FORIG), University P. 0.Box 63, KNUST, Kumasi, Ghana (Gineiino orboreo, Heritiero utilis)R. Aerts, Division Forest, Nature and Landscape, Katholieke Universiteit Leu-ven, Celestijnenlaan 200E, box 2411, BE-3001, Leuven, Belgium (Afrocoipusfolcotz, s, Erythrina o6yssinico)N. S. A1varez Cruz, Unidad de Medio Ambiente, Delegaci6n del CITMA, Cor.Legon 268 I Henry Reeve y Carlos Roloff, Sancti Spiritus C. P. 60100, Cuba(Autoenniogerminons, Antcennio inuring)L. C. N. Anglaaere, Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, KNUST, Kumasi, Ghana (Pertcopsiseloto)MM. Apetorgbor, Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, KNUST, Kumasi, Ghana (A1bizio zygio)J. Ayarkwa, Forestry Research Institute of Ghana (FORIG), University P. 0.Box 63, KNUST, Kumasi, Ghana (Cylicodiscus gobunensis)P. Baas, NationaalHerbarium Nederland (NHN), University of Leiden branch,P. 0. Box 9514, 2300 RA Leiden, Netherlands (wood anatomy)H. Bakamwesiga, Institute of Environment & Natural Resources, MakerereUniversity, P. 0. Box 7298, Kampala, Uganda (Juniperus procero)L. N. Banak, Institut de Recherches en ECologie Tropicale (IRET), Centre Na-tional de in Recherche Scientifique at Technologique (CENAREST), B. P. 842,Libreville, Gabon (wood anatomy)H. Beeckman, Laboratory for Wood Biology andXylarium, Royal Museum forCentral Africa, Leuvense steenweg 13, B-3080 Tervuren, Belgium (wood anat-omy)E. J. Bertrums, PROTA Network Office Europe, Wageningen University, P. 0.Box 341, 6700 AH Wageningen, Netherlands (layout)L. Bonn6hin, Projet autonome pour in conservation du Parc national de Tai(FACPNT), 01 B. P. 693, San Fedro, C6te d'TVoire (neghemello hechatii)C. H. BOSch, PROTA Network Office Europe, Wageningen University, P. 0. Box341, 6700 AH Wageningen, Netherlands (Stercttlio appendiculoto, Sterculi0 o6-longo, Sterculio quinquelobo, Sterculio rhinopetolo)P. P. BOSu, Forestry Research Institute of Ghana (FORTG), University P. 0. Box63, KNUST, Kuruasi, Ghana (Antioris toxicorio, Triplochiton sclero, cylon)M. Brink, PROTA Network Office Europe, Wageningen University, P. 0. Box341, 6700 AH Wageningen, Netherlands (Bombuso uulgoris, Benschmiediocorbisieri, Betlschmiedio dirersiftoro, Betlschmiedio fouleo, Betlschmtedio IOU-isit, Benschmiedio oblongifolio, Betlschmiedi0 14gondensts, Betlschmiedio ue-jutino, Berryo cordifolio, Curiorium mudogoscoriense, Corpodjptero of ticono,Gossipoure0 o1/2e!it, Gossipoureo euryoides, Gossipoureogummiftuo, Cussi-poureo inglosono, Gossipoureo ruu)ensoriensis, Cothoriostochys inodogos-conensts, Chlomydocolo chiomydontho, Christiono ofricon0, 0010 coulifloro,Cold CIOuoto, Colo greener;oyi, Golo latentio, Cola IOUrifolio, Comintphoropterocorpo, Cupressus instronico, Doeryodes igogongo, Doeryodes Morneono,

Docryodes normondii, Docryodespz, bescens, Dendroco!Qinus OSper, Dendro-coldmus giganteus, Doin beyo rotundifolio, Doin beyo torndo, Duboscio inocro-corpo, EUColyptus CIOeziono, EUColypttts tereticornis, Eucalyptus viminolis, Fi-cus uoriifolio, Ficus Dogeliono, Grett!to bicolor, Grewio mollis, Hibiscus lostococ-ci4s, Hildegordio erythrosiphon, Hypodophnis zenkeri, Juniperus bermz4dtono,Kirhio ocuminoto, Letestuo dunssimo, Neolemonniero clitondrifolio, Not-hospondios stoudtii, Ochromopyromidole, Octolobus spectobtlis, Odyendyeogobonensis, Pinus cartboeo, Pinus elltottii, Pinus o0corpo, Pinus rodioto,Premno origolensis, Premno muximo, Rhodognopholon schumonnionum,Sidero, cylon titerme, Streblus dimepote, Syzygium cordotum, Voltho diffuso,Widdrtngtonio whytei, editor)J. R. Cobbinah, Forestry Research Institute of Ghana (FORIG), University P. 0.Box 63, KNUST, Kumasi, Ghana (Quossio unduloto, generaleditor)C. Couralet, Royal Museum for Central Africa, Leuvensesteenweg 13, 3080Tervuren, Belgium (Juniperus procero)P. Derienne, CTRAD, Departement Persyst, UpR Bois tropicaux, 73 rue Jean-Francois Breton, TA B-40 I 16 (Bat. 16, Bur. 106), 34398 Montpellier Cedex 5,France (wood anatomy)D. Dongock Nguemo, Departement des Sciences Biologiques, Facult6 des Sci-ences, Universit6 de Ngaound6r6, BP 454, Ngaound6r6, Cameroon (Cedrelopsisgreuei)J. C. Doran, Forestry and Forest Products, CSIRO, Queen Victoria Terrace, P. 0.Box 4008, Kingston, ACT 2604, Australia (EUColyptus coinoldulensis)C. Doumenge, Cirad, Campus International de Baillarguet TA-C-3610, F-34398Montpellier cedex 5, France (Doeryodes buettneri)CS. Duvall, Department of Geography, Bandelier West, Room 121, Universityof New Mexico, A1buquerque, NM, 87131, United States (Getbopentondro,Pterocorpus erinoceus)E. Ebanyenle, Forestry Research Institute of Ghana (FORTG), University P. 0.Box 63, KNUST, Kumasi, Ghana (wood anatomy)P. E. Gasson, JodrellLaboratory, Royal Botanic Gardens, Kew, Richmond, Sur-reyTW9 30S, United Kingdom (wood anatomy)J. N. Gyakari, Forestry Commission, Box 1457, Kumasi, Ghana (Quossio undu-loto)J. B. Hall, School of Agricultural and Forest Sciences, University of Wales, Ban-gor, Gwynedd LL57 2UW, United Kingdom (Orytenonther0 o6yssinico, Sino-rundinorio ulpino)I. Hemz, Thadenstrasse 110B, 22767 Hamburg, Germany (wood anatomy)T. Triada, 3-21-12, Toyotamanaka, Nerimaku, Tokyo, 176-0013, Japan(03cytenonther0 o6yssinico, Sinorz, ridinori0 o1pino)P. C. M. Jansen, PROTA Network Office Europe, Wageningen University, P. 0.Box 341, 6700 AH Wageningen, Netherlands (Pterocorpus soyou, cii)R. B. Jiofack Tafokou, ECologic Museum of Cameroon, P. 0. Box 8038, Yaound6,Cameroon (EUColyptus robusto, Guoreo cedroto, Piptodeniostrum @1nconum,Zanthoxylum heitati)V. A. Kerneuz6, Millennium ECologic Museum, BP 8038, Yaound6, Cameroon(Britondrophrogmo cylindricum, Rhodognopholon breuicz, spe)

CONTRIBUTORS 7

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8 TIMBERSl

E. Krampah (+), Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, KNUST, Kumasi, Ghana (Antioris toxicorio, Gineiino orboreo,Triplochtton sclero, cylon)K. J. M. Ky, Centre National de Semences Forestfores, 07 B. P. 5252, 0uagadou-gou 07, Burkina Faso (Vite, c doniono)R. H. M. J. Lemmens, PROTA Network Office Europe, Wageningen University,P. 0. Box 341, 6700 AH Wageningen, Netherlands (ACUcio coffro, ACUciogo!pinti, ACUcio heterophyl!0, ACocio melono, cylon, ACUcio wigrescens, ACUciorobusto, ACUcto rouumoe, ACUcto ranthophloeo, Adenonthero montoroo, Aes-chynomene elophro, cylon, Alofttsilodendron uillosum, A1bizio odionthifolio, A1-bigt0 o1tissimo, A1bizio gritunesiono, A1btzio orenicolo, A1bizio orylmert, A1biziobemien, A1bizio botuinit, A1bizio globerrimo, A1bizio schimperto, IQ, A1bizio uer-siColor, AIStonto congensis, Amblygonocorpus undonger, sis, Amphimos Ierrugi-iteus, Aroz, cono cunninghornii, Aubreuilleoplotycorpo, AMtronello congolensis,Bophio kithii, Bornsonthus spedosus, 0010dendrz, in copense, 0010phyl!urn trio-phyllum, Golpoco!y, c breuibrocteotus, CUIpocoly, c heirzii, Coraljin brochioto,Cedrelo odoroto, Chrysophyllum o17iconz, in, Chrysophyllttm botuinionum,Chrysophyllttm gigonteum, Chrysophyllum IOCourtionum, Chrysophyllumper-PMlchrum, Chrysophyllumpruniforme, Chrysophyllttm subiLudum, Chukrosiotubularts, Do16ergto boronii, Do16ergio chopeliert, Do16ergto chiorocorpo, DoI-bergio greueono, Do16ergto hitdebrondtii, Do16ergio lotifo!to, Do16ergio IOU-uelii, Do16ergio ingdogoscoriensis, Do16ergio melonoxylon, Do16ergio mollis,Do16ergio monticolo, Do16ergio nitidulo, Do!bergiopt, rpuroscens, Do16ergiosissoo, Do16ergio trichocorpo, Dichrostochys myriophyllo, Dicroeopetoluminghofoliense, Dypsts mudogoscoriensis, ETLtodoperuillei, Britondrophrogmoexcelsum, Erythrino excelso, Erythrino Doge!it, Pogoropsis origolensis, Filloeop-sts discophoro, Guoreo thornpsonii, HOPIormosio monophyllo, Heritiero denst-noro, Heritiero littorolis, Khoyo toorensis, Lecomted03co hloineono, Lepidotri-chino o01hensii, Mornme0 @1ncono, Milletttogrondis, Millettio rhodontho, Mil-lettio richerrdiono, Millettio sti, himunitii, Millettio Dersicolor, Mimusops on-dongensis, Mimusopscoffro, Mtmusopselengi, Mimusops hummel, Mimusopsmuximo, Mimusops zeyheri, Neohormsio boronii, Newtonio let4cocorpo, Newto-ittopoucjjugo, Ocoteo cymoso, Ocoteo henyensis, Ompholocorpum elotum,Oreobombos buchu)oldii, Ormocorpopsis itreinoensis, Porkio flitcoideo, Phyl-109:ylon perrieri, Phyllo, cylon icylophylloides, Plotysepolum cheuolieri, Plotyse-perlum uioloceum, Platocorpopycnontho, Poitgomtopsisperutlleono, Pouterioudoni-friedendi, Pouterio Qinifolio, Pouteri0 o1tissimo, Pouterto oningeri,Pseudobersomo mossombicensts, Pseudocedrelo hotschyi, Ptoero, cylon o61tquum,Pterocorpus IIJcens, Pterocorpus OSun, Pterocorpus rotundifolius, Pterocorpussontolinoides, Pterocorpus tinctoriz, s, Pyronthus o10800, Quiuisionthepopinoe,Sohoonolo inodogoscoriensis, Suitetento macrophyl!0, Suite terrier inghogoni,Synsepo!urn o1^:elii, Synsepolum breuipes, Tobernoemonto?, o stopfiono,Tieghemello of neono, neghemello hechem, Toono echoto, Trichilio gilgiono,Trichilio monodelpho, Trichtlioprieureono, Voughonio dionoeifolio, Veprtsloneeoloto, Vepris nobilis, Viguieronthus kony, Vite, c grondifolio, Vite, c micron-tho, Xonthocercis inodogoscoriensts, Xeroderris stuhlmonnii, Xylio euonsii,Zoritho, cylum douyi, Zorithoxylz, in tsthonimposo, general editor, illustrations)

D. Louppe, CTRAD, Departement Environnements et Societes, Cirad es-dir,Campus international de Baillarguet, TA C-DIR IB (Bat. C, Bur. 1/3), 34398Montpellier Cedex 5, France (Boillonello tortspermo, Broussonetio greueono,Sterculio OPPendiculoto, Sterculio oblongo, Sterculio quinquelobo, Stercz, flurhinopetolo, rectono grondis, wood anatomy, editor)A. U. Lumbile, Botswana College of Agriculture, Private Bag 0027, Gaborone,Botswana (Ficus sur, Pertcopsis origolensis)F. S. Mairura, Tropical SoilBiology and Fertility Institute of CIAT, P. 0. Box30677, Nairobi, Kenya (Ekebergio copensis, NewtorLi0 614chononii, Ocoteo hen-yensis)A. Maroyi, Department of Biological Sciences, Bindura University of ScienceEducation, P. B. 1020, Bindura, Zimbabwe (A1biziogz, minifero, Khoyo on-thotheco, Syzygium gutneense)K. K. Mogotsi, Botswana College of Agriculture, Private Bag 0027, Gaborone,Botswana (Ficus sur)N. P. Monel, Tropical Pesticides Research Institute, National Herbarium of Tan-zania, P. 0. Box 3024, Arusha, Tanzania (wood anatomy)P. Mugabi, Forest and Wood Science Department, Stellenbosch University,BOSman Street, Stellenbosch 7600, South Africa (wood anatomy)DB. Mujuni, P. 0. Box 1752, Kampala, Uganda (Enterridrophrogmo utile)P. Ng'andwe, Department of Wood Science and Technology, SchoolofNaturalResources, The Copperbelt University, P. 0. Box 21692, Kitwe, Zambia (woodanatomy)S. A. Nigro, Research Centre for Plant Growth and Development, Schoolof Bio-logical and Conservation Sciences, University ofKwaZulu-Natal, Private BagXOl, Scottsville 3209, South Africa (Pinuspotulo)A. Nikiema, ICRISAT Sahelian Centre, B. P. 12404, Niamey, Niger (Khoyoservegolensis)AChmad Satin Nurhaman, Southeast Asian Regional Centre for Tropical Biol-ogy (SEAMEO BIOTROP), P. 0. Box 17, Bogor, Indonesia (illustrations)NyunaiNyemb, Institut de Recherches Medicales at d'Etudes des PlantesMedicinales, B. P. 3805, Yaound6, Cameroon (Betlschmiedio monnii, Errtondro-phrogmo condollei, Eucalyptus grondis, Louoo trichilioides, Pinus foesiyo)O. Oagile, Botswana College of Agriculture, Private Bag 0027, Gaborone, Bot-swana (Pertcopsis origolensis)D. A. Of on, Forestry Research Institute of Ghana (FORIG), University P. 0. Box63, KNUST, Kumasi, Ghana (Mindo excelso, Mindo regio)F. Ohene-Coffie, Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, KNUST, Kumasi, Ghana (Monsoni0 o1tissimo)J. M. Okeyo, TSBF-CIAT, World Agroforestry Centre (ICRAF), P. 0. Box 30677,Gigiri, Nairobi, Kenya (Ocoteo usumborensis, Podocarpus lotifolius)M. M. Okeyo, LondianiRegional Research Centre, P. 0. Box 382 - 20203,Londiani, Kenya (Zoritho, cy!urn gilletii)E. Opuni-Frimpong, Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, ERUST, Kumasi, Ghana (Khoyogrondifoliolo)A. A. Oteng-Amoako, Forestry Research Institute of Ghana (FORTG), UniversityP. 0. Box 63, KNUST, Kumasi, Ghana (Pinus conboeo, wood anatomy, editor)B. Ouattara, Rue de I'Et6 4, 1050 Brussels, Belgium (Morus meSO3ygio)

CONTRIBUTORS 9

10 TIMBERSl

P. Oudhia, SOFAA1, 28-A, Geeta Nagar, Raipur, 492001, C. G. , India (Thespesiopopulneo)F. W. Owusu, Forestry Research Institute of Ghana (FORIG), University P. 0.Box 63, KNUST, Kumasi, Ghana (Cyltcodiscus gobz, itensis, TMrroeonthus ofri-coitus)L. FA. Oyen, PROTA Network Office Europe, Wageningen University, P. 0. Box341, 6700 AH Wageningen, Netherlands (Nesogordonio holtzii, Nesogordor, tohubingoensis, Pterygoto mererocorpo, Pterygoto mildbroedii, Symphonio globu-filero, general editor)F. Falla, SIC Cellule de coordination du programma ECOFAC, B. P. 15, 1/5 Li-breville, Gabon (AIStonio boonei)D. Pasternak, ICRISAT Sahelian Centre, B. P. 12404, Niamey, Niger (Khoyosenegolensis)G. H. Schmelzer, PROTA Network Office Europe, Wageningen University, P. 0.Box 341, 6700 AH Wageningen, Netherlands (Chioroxylonfoho, Chioroxylonsuitetento)M. M. Spitteler, Het Hoge Stuk 19, 8431 KL 00sterwolde, Netherlands (Illustra-tions)Iskak Syamsudin, Herbarium Bogoriense, Research Centre for Biology - LIPl,Jalan Ir. H. Juanda 22, Bogor 16122, Indonesia (illustrations)R. Takawira-Nyenya, National Herbarium and Botanical Garden, P. 0. BoxA889, Avondale, Harare, Zimbabwe (Pterocorpus origolensis)P. Tan6, Departement de Chimie, Universit6 de DSChang, BP 67, DSChang, Ca-meToon (Amphimospterocorpoides)A. T. Tchinda, Institut de Recherches Medicales at d'Etudes des Plantes Medici-nales (IMPM), Ministere de to Recherche Scientifique at de I'Innovation, B. P.6163, Yaound6, Cameroun (Amphimospterocorpotdes, Britondrophrogmo origo-lense, Millettto IOUrentii, Porkio bicolor)M. Thiam, Unite de Formation at de Recherehes, Sciences Agronomiques at 06-veloppement Rural, Universit6 de Thi6s, B. P. A296, Thi6s, Senegal(wood anat-omy)H. J. C. Thijssen, Mirabelweg 16, 5632 PC Eindhoven, Netherlands (Vite, cfische-ri)G. Todoll, Herbier National du Cameroun, B. P. 1601, Yaound6, Cameroon (Do-cryodes buettneri)B. TDIrambe Bamoninga, Laboratoire de Biologie du bois at Xylarium, MuseeRoyal pour I'Afrique Centrale, Leuvensesteenweg 13, 3080 Tervuren, Belgium(Morus me802ygio)K. Twum-Ampofo, Institute of Renewable Natural Resources, Department ofAgroforestry, Kwame Nkrumah University of Science and Technology(KNUST), Kumasi, Ghana (A1bizioferrugineo)E. Uetimane, Departamento de Engenharia F10restal, Faculdade de Agronomiae Engenharia F10restal, Universidade EduardiMondlane (UEM), Maputo, Mo-zambique (wood anatomy)J. L. C. H. van Valkenburg, De Hoef45, 6708 DC Wageningen, Netherlands (Arycoumeo Morneono)G. Vaughan, 3620 N. Wilton, Chicago, IL 60613, United States (EUColyptttsglobulus)

W. Wessel-Brand, P. Potterhof 23, 4033 AN Lienden, Netherlands (illustra-tions)E. A. Wheeler, Department of Wood and Paper Science, College of Forest Re-sources, North Carolina State University, P. 0. Box 8005, Raleigh, NC 27695-8005, United States (wood anatomy)W. Wongkaew, Department of Botany, Faculty of Science, Kasetsart University,Chatuchak 10903, Bangkok, Thailand (EUColyptus coma!dulensis)

Acknowledgments

PROSEA Foundation, Jalan H. Juanda 22, P. 0. Box 332, Bogor 16122, Indonesia(use of text parts and illustrations of species overlapping between South-EastAsia and tropical Africa)S. van Otter100-Butler, Bowlespark 21, 6701 DR Wageningen, Netherlands(English language correction)N. Wulijarni-Soetjipto, Jl. Fahlawan 1/3, Bogor 16131, Indonesia (coordinationillustrators)

CONTRIBUTORS 11

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PROTA Board of Trustees and Personnel

Board of Trustees

M. J. Kropff(WU, Netherlands), chairZ. L. K. Magombo (NHBGM, Malawi), vice-chairV. Agyeman (FORTG, Ghana)H. Andriamialison (PBZT, Madagascar)H. GB. Carsalade (AGROPOLIS, France)D. Garrity (ICRAF, Kenya)M. Honadia (CNSF, Burkina Faso)S. D. Hopper(RBGKEW, United Kingdom)D-. F. Idiata (CENAREST, Gabon)L. S. Luboobi(MU, Uganda)E. Sukara (PROSEA, Indonesia)

Personnel

Regionol Office CentrolAfrico, Gobon

J. A. Bourobou Bourobou, Regional OfficerE. D. Eyone Edoh, Assistant Regional OfficerD. N. Omokolo, Contact Person CameroonM. K. D. Ben-Bala, Contact Person Central African Republic

Regionol Office EastAfrico, Ugondo

G. Nakabonge, Regional OfficerC. Bulafu, Assistant Regional OfficerA. Tsegaye, Contact Person EthiopiaJ. Elia, Contact Person Tanzania

Regional Office Inchon Oceon 1810nds, Modogoscor

ME. Rahelivololona, Regional OfficerA. Gunb-Fakim, Contact Person MauritiusS. Brillant, Contact Person Reunion

Regionol Office SouthernAfrico, Molowi

J. Kamwendo, Regional OfficerV. K. Kawanga, Contact Person ZambiaO. Oagile, Contact Person BotswanaS. Kativu, Contact Person Zimbabwe

Regionol Office (orig!ophone) WestAfrico, Ghono

JR. Cobbinah, General Editor

S. Britwum-ACquah, Regional OfficerBE. Ewudzie, Assistant Regional OfficerOA. Denton, Contact Person NigeriaA. B. Kanm, Contact Person Sierra Leone

Regionol Office ofroncophone) WestAfrico, Burhino Faso

A. Traor6, Regional OfficerC. Kouam6, Contact Person C6te d'TVoireF. Assogba-Komlan, Contact Person BeninH. Sanou, Contact Person MaliL. Mahamane, Contact Person Niger

Country Office Fronce

M. Chauvet, Technical Editor FrenchC. Dorthe, Country Officer

PROTABOARDOFTRUSTEESANDPERSONNEL 13

Country Office United Kingdom

O. Grace, Country Officer

Network Office A1rico, Kenyo

E. A. Omino, HeadDJ. Borus, Dissemination OfficerJ. Chege, Database OfficerB. Owuor, SecretaryD. Laur, Office Assistant

Network Office Europe, Netherlurids

J. S. SIemonsma, HeadR. H. M. J. Lemmens, General EditorL. P. A. Oyen, General EditorE. J. Bertrums, Databank ManagerC. H. BOSch, Editor/Dissemination OfficerM. Brink, EditorG. H. Schmelzer, Editor/Dissemination Officer

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14 TIMBERSl

Introduction

PROTA 7(I):'Timbers I'is the first of 2 volumes describing the wild and cultivatedplant species of tropical Africa used for their timber. Some of these are traded on theinternational timber market, but many are only used locally, for construction pur-poses or the production offurniture, implements and utensils. Bamboos of which thestems are used for construction are also included in this commodity group. Mostspecies have several other, secondary, uses. PROTA assigns one primary use and, ifrelevant, one or more secondary uses to all plant species used in Africa. For in-stance, the primary use of Chrysophyllum IOCourtior, urn De Wild. is as a timbertree, and thus it is treated in PROTA 7, but it has several secondary uses, e. g. thefruits are eaten and the bark is used in traditional medicine. The timber of Chryso-phyllum o16idum G. Don is also commonly used, butthe primary use of this speciesis as a fruit, and consequently it is described in PROTA a 'Fruits'.

Table I. Families treated in PROTA 7(I) and the division of species.

Family

ApocynoceoeAruucortoceoe

Arecueeoe

Auicennioceoe

Bombocuceoe

Buyseroceoe

01ustoceoe

CupressoceoeLouroceoe

Moluoceoe

Mengceoe

liftmosoceoe

114'070ceoe

MyrtoceoePupilionoceoePingceoe

Poorceoe

PodocorpoceoeRhizophoroceoeRutoceoe

SopotoceoeStingrouboceoeSterculioceoeTiliaceoe

Verbenoceoe

Genera

3

I

I

I

3

4

3

3

3

2

15

16

6

2

21

I

7

2

2

7

12

4

11

5

4

Majorspecies

2

o

o

2

2

3

3

3

3

2

17

11

7

5

18

3

3

2

I

4

10

I

7

o

4

Minor

species

2

I

I

o

2

5

o

I

9

o

9

28

2

4

35

3

5

o

5

8

18

3

16

6

4

Mentioned

species

Total

2

2

o

o

o

8

9

4

18

3

18

40

o

12

54

o

I

. 3

3

18

12

I

10

7

6

Total

species

6

3

I

2

4

16

12

8

30

5

44

79

9

21

107

6

9

5

9

30

40

5

33

13

14

139 1/3 167 231 511

In PROTA 7(I), some species are treated which, in addition to the primary use astimber, also have another primary use and consequently are described in 2 PROTAbooks. These species are Getbopentondro (L. ) Gaertn. (also in PROTA 16:'Fibres'),EUColypti4s coinoldulensis Dehnh. (also in PROTA 10:'Fuelplants'), Pterocorpz, sorigolensis DC. (also in PROTA 3:'Dyes and tannins') and Pterocorpus soyou, ctiTaub. (also in PROTA 3:'Dyes and tannins').In 'Timbers I' allprimary-use timber species of 25 importanttimber-containingfamilies are treated. 'Timbers 2' will comprise the timber species of the remainingfamilies. Comprehensive descriptions are given of 1/3 important timber species.These major timbers comprise mostly wild species, but also several cultivated orpartly domesticated species. The accounts are presented in a detailed format andillustrated with a line drawing and a distribution map. In addition, accounts of 167timbers of minor importance are given. Because information on these species is of-ten scanty, these accounts are in a simplified format and usually do notinclude adrawing or map. For another 231 species the information was too scarce to justify anindividual treatment and they have only been mentioned in the accounts of relatedspecies.

The 'primary use'timbers nottreated in PROTA 7(I) will be included in volume7(2). In this final volume, all'secondary use' timbers will be listed as 'Timbers withother primary use' and referred to other Handbook volumes.

Plant names

Family: Apartfrom the classicfamily name, the family name in accordance with theAngiosperm Phylogeny Group (APG) classification is also given where it differs fromthe classic name.

Synonyms: Onlythe most commonly used synonyms and thosethat may causecon-fusion are mentioned.

Vernacular names: Only names in official languages of regional importance in Africaare included: English, French, Portuguese and Swahili. It is beyond the scope ofPROTA to give an extensive account of the names of a species in alllanguages spo-ken in its area of distribution. Checking names would require extensive fieldwork byspecialists. Although regional forms of Arabic are spoken in several countries in Af-rica, the number of African plant species that have a name in written, classical Ara-bic is limited. Arabic names are therefore omitted. Names of plant products arementioned under the heading 'Uses'.

INTRODUCTION 15

Origin and geographic distribution

To avoid long lists of countries in the text, a distribution map is added for majorspecies. The map indicates in which countries a species has been recorded, eitherwild or planted. It should be realized that for many species these maps are incom-PIete because they are prepared on the basis of published information, the quantityand quality of which varies greatly from species to species. This is especially thecase for wild species which are not or incompletely covered by the regional Africanfloras, and for cultivated species which are only planted on a smallscale (e. g. inhome gardens). For some countries (e. g. Central African Republic, Chad, Sudan,Angola)there is comparatively little information in the literature. Sometimes they are

16 TIMBERSl

not covered by recent regional or national floras and although species may be presentthere, this cannot be demonstrated or confirmed. For some major species, a distribu-tion map has been omitted because there is too little information on distribution.

Properties

Wood appearance (i. e. colour, grain, texture), physical properties (density, dryingproperties, shrinkage rates, movement in service) and mechanical properties(strength, elasticity, hardness) of the wood are given on the basis of literature andas much as possible in quantitative terms and in a standardized order. This is fol-lowed by a description of the working properties and the durability of the wood.

Description

A inorphologicalcharacterization of the species is given. The description is in 'tele-grain' style and uses botanical terms. Providing a description for the general publicis difficult as more generally understood terms often lack the accuracy required in abotanical description. A line drawing is added for all major and some lesser-knownspecies to complement and visualize the description.

Anatomy

A wood-anatomical description has been prepared for each major timber species.The descriptions of the hardwood species are based on the IAWA (International As-SOCiation of Wood Anatomists)list for hardwood identification myheeler, Baas &Gasson, 1989), those of the softwood species on the IAWA list for softwood Identifi-cation (Richter, Grosser, Hemz & Gasson, 2004). All descriptions have been coin-piled in the PROTAWood Anatomy Workshop (16-25 May, 2007, Montpellier,France) as part of a training of African wood anatomists under expert guidance. Thecompilers of each description are mentioned at the end of the paragraph 'Anatomy'.For many descriptions the InsideWood database cottp://insidewood. lib. ncsu. edu) hasbeen used as a basis, but for all descriptions microscopic sections of wood from theslide collections of CIRAD muontpellier, France), the Royal Museum for Central at-rica (Tervuren, Belgium), the Royal Botanic Gardens (Kew, UK), and the NationalHerbarium of the Netherlands (Leiden, Netherlands) have been investigated duringthe wood anatomy workshop. The final descriptions have been included in the In-sideWood database.

The character codes given in parentheses O indicate characters which are variableor rarely occur in the materialstudied. In character code 23 a question mark (?)issometimes added, indicating the presence of smallpits but difficulties to see the pitoutline.

Management

Descriptions of husbandry methods including fertilizer application, irrigation, andpest and disease control measures are given under 'Management' and under 'Dis-eases and pests'. These reflect actual practices or generalized recommendations, opt-ing for a broad overview but without detailed recommendations adapted to the

I

widely varying local conditions encountered by farmers' Recommendations onchemical controlofpests and diseases are merely indicative and local regulationsshould be given precedence. PROTA will participate in the preparation of derivedmaterials for extension and education, for which the texts in this volume provide abasis, but to which specific local information will be added.

Genetic resources

The genetic diversity of many plant species in Africa is being eroded, sometimes atan alarming rate, as a consequence of habitat destruction and overexploitation. Thereplacement of landraces of cultivated species by modern cultivars marketed by seedcompanies is another cause of genetic erosion. Reviews are given of possible threatsfor plant species and of the diversity within species, and reference is made to theIUCN Red list of threatened species where relevant. Information on ex-situ germ-PIasm collections is mostly extracted from publications of Bioversity International(formerly the International Plant Genetic Resources Institute - IPGRl).

References

The main objective of the list of references given us to guide readers to additionalinformation; it is notintended to be complete or exhaustive. Authors and editorshave selected two categories of references;'major references are limited to 10 refer-ences (5 for minor species), the number of'other references'is limited to 20 (10 for inI-nor species). The references listed include those used in writing the account. Wherethe internet was used, the website and date are cited.

INTRODUCTION 17

,

- co -a 1< ai

I;a 56 co

-.

I-,

Alphabetical treatment of timbers

19

20 TIMBERSl

ACACIA CAFERA (Thunb. ) Wind

Protologue Sp. PI. 4(2): 1078 (1806).Family Mimosaceae (Leguminosae - Mimo-

soldeae)Chromosome number2n=26

Vernacular names Commonhookthoru (En)Origin and geographic distribution ACUcio

coffro is found mostly in northern and easternSouth Africa and Swaziland, but occurs also insouthern Botswana and southern MozambiqueIt is found in small numbers near Harare inZimbabwe, probably the result of an Introduc-tion long ago. It is planted in India

Uses The wood is used for fence poles andoccasionally for furniture. The Xhosa people ofSouth Africa used it for making traditionalsmoking pipes, and it also serves as firewoodThe bark is used for tanning; it produces palebrown leather. Baskets are made from the

twigs. A leaf decoction with milk is used by theZulu people of South Africa as an enema totreat abdominal complaints in children; theleaves are sometimes chewed for the same

purpose. A leaf decoction is also drunk to treatcolds and fever. A bark infusion is adminis-tered as a blood purifier. Leaves and pods areeaten by livestock, but may cause poisoning

Properties The heartwood is dark brownand distinctly demarcated from the creamysapwood. The texture is moderately fine tomoderately coarse. The wood is heavy andhard. The density is 980-1060 kg/ina at 12%moisture content. The wood seasons withoutserious defects

A number of proteracacinidins (proanthocya-Didiris) have been isolated from the heartwood,and several cyanogenic glycosides from theleaves

Botany Shrub or smalltree up to 14 in tall;hole often twisted, up to 60 cm in diameter;bark rough, sometimes fissured, reddish brownto blackish brown; crown spreading; branchletsglabrous to densely hairy, with pairs of hookedprickles up to 9 mm long just below the nodesLeaves alternate, hipinnately compound, with6-38 pairs of pinnae; stipules linear, 2.5-4 mmlong, caducous; petiole 0.5-4 cm long, withgland near top, rachis 2-23 cm long, sometimeswith prickles up to 3 min long and with glandsbetween top pairs of pinnae; leaflets in 16-64pairs per pinna, linear to linear-oblong, 2-12mm x 0.5-2.5 mm, oblique at base, withrounded to acute apex, glabrous to pubescentInflorescence an axillary spike 2-10 cm long,solitary or in fascicles; peduncle up to 4 cm

long. Flowers bisexual, regular, 5-merous,small, creamy white, sessile; calyx campanu-late, tube 0.5-1.5 mm long, lobes shorter; co-rolla campanulate, tube up to 2 min long, lobestriangular, up to I min long; stamens numer-ous, free, up to 6 min long; ovary superior, 0.5-2 mm long, sessile or stalked, style slender, upto 6 mm long. Fruit an oblong to linear pod4.5-19.5 cm x 0.5-2.5 cm, usually straight,glabrous to hairy, reddish brown scaly, longi-tudinally dehiscent, up to 10-seeded. Seedsflattened ovoid to oblong, 6-12 mm x 4-8 min,olive green to pale brown.ACocio is a large pantropical genus, comprisingmore than 1300 species; most of them are dis-tributed in Australia (more than 900), morethan 200 in America, and about 130 in AfricaACocto coffro belongs to subgenus ACUleiferum,which comprises all African ACociu specieswith non-spinescent stipules and hooked prick-Ies. ACocio coffro is variable, especially in itsindumentum, number of pinnae per leaf andleaflet size. It is sometimes confused with ACo-cio ortoxocontho DC. and Acercio hereroe, 1sts

Engl. , the first of which has scattered pricklesand the second smaller leaves and shorter peti-o1es

The seeds are probably dispersed by large her-bivores such as elephants and antelopes and bybaboons, allofwhich eatthe pods.

Ecology ACocio coffro occurs in open wood-land, wooded grassland and on dry rocky hills,often along watercourses, up to 1500 in alti-tude.

Management ACocio cuffro coppices well.Genetic resources and breeding Although

Acercio coffro is only found in the most south-ern parts of Botswana and Mozambique, it iswidespread in South Africa and occurs there ina variety of habitats. Therefore, it is unlikely tobe under threat of genetic erosion.

Prospects ACocio coffro will remain onlylocally important as timber tree, but it haspotential as a fast-growing and attractive or-namentalwhich is drought- and frost-resistant

Major references Brenan, 1970; CoatesPalgrave, 1983; Ross, 1967; Ross, 1975; Tim-herlake, Fagg & Barnes, 1999

Other references Bennie at a1. , 2002; Connat a1. , 1989; Neuwinger, 2000

Authors R. H. M. J. Lemmens

ACACIA 21

.

22 TIMBERSl

ACACIAGALPINllBurtt Davy

Protologue Bun. Misc. Inform. Kew 1922(Icy326 (1922)

Family Mimosaceae (Leguminosae . Mimo-soldeae)

Chromosome number2n=40Vernacular names Monkey thorn (En)Origin and geographic distribution ACocio

gulptnii occurs in Tanzania, Malawi, Zambia,Botswana, Zimbabwe, Mozambique and north-ern South Africa. Experimental plantationsexist in Madagascar.

Uses The wood is used for interior and exte-

nor joinery, fences, wagons, railway sleepersand as mining timber. It is suitable for floor-ing, shipbuilding, sporting goods and imple-merits. ACUcio go!pmit is occasionally plantedas ornamental or roadside tree.

Production and international trade Thewood is only used locally and nottraded on theinternational market.

Properties The heartwood is reddish brownto dark brown and distinctly demarcated fromthe creamy sapwood. The grain is often itregu-Iar, texture moderately coarse. The density ofthe wood is about 800 kg/ing at 11% moisturecontent. Shrinkage during drying is medium.At 11% moisture content, the modulus of rup-ture is 1/2 N/mm2, modulus of elasticity 13,140N/mm2, compression parallel to grain 61N/min2, shear 15.9 N/mm2 Janka side hard-ness 9070 N and Janka end hardness 10,100 N.Although the wood is tough and resilient, itworks well with sharp tools, but sawing re-quires considerable strength. A good finish canbe obtained with waxes and oil. The wood hasgood natural durability, being fairly resistant

to decay and termite attack, but the sapwood issusceptible to attack by Lyctus borers and bluestain fungiThe wood is almost free of tannins, but con-tains a range of flavonoids and meIacacidin. Anumber of proteracacinidins (proanthocyanid-iris) have been isolated from the heartwood

Description Small to fairly large tree up to30 in tall; hole usually straight, up to 60(-150)cm in diameter; bark rough, flaking or longitu-dinally furrowed, greyish to greyish brown,often with yellowish tinge; crown spreading,rounded; branchlets glabrous to shortly hairy,with pairs of hooked, blackish prickles up to Icm longjust below the nodes. Leaves alternate,hipinnately compound, with 6-14 pairs of pin-nae; stipules linear-oblong, c. 2 min long, cadu-cous; petiole c. 3 cm long, with small gland inupper half, rachis 6-11 cm long, with glandsbetween top pairs of pinnae; leaflets in (8-)13-35(-45) pairs per pinna, narrowly oblong tolinear-oblong, (2-)4-TIC15) mm x (0.591-3(-4) min, oblique at base, apex obtuse to acute,glabrous but often with some hairs at marginsInflorescence an axillary spike 4-11 cm long,often in fascicles on short lateral leaflessshoots; peduncle 0.5-1.5 cm long. Flowers bi-sexual, regular, 5-meTous, small, fragrant, ses-SIIe; calyx cup-shaped, 0.5-1.5 min long, red topurple; corolla c. 2 min long, with short lobes,red to purple; stamens numerous, free, up to 5min long, creamy yellow; ovary superior,stalked, style slender, up to 5 mm long. Fruitan oblong to linear pod 11-28 cm x 2.5-3.5 cm,straight, glabrous, reddish to purplish brown,longitudinalIy dehiscent, up to 12-seededSeeds flattened ovoid, 12-15 mm x 10-12 mm.

Other botanical information Acercio is a

large pantropical genus, comprising more than1300 species; most of them distributed in AUS-tralia (more than 9000, more than 200 in Amer-ICa, and about 130 in Africa. ACocio gulpiniibelongs to subgenus ACUleiferum, which in-cludes all African ACocio species with non-spinescent stipules and hooked prickles. ACociogobytnii is often confused with Acacia poly-ocontho Wind. , but the latter has a largergland near the base of the petiole, and a largernumber of pinnae per leaf. The wood of ACUciopolyocontho is used for similar purposes as thewood of ACocio galpinii, but its gum is moreJinportant.

The wood of ACocio burhei Benth. (black mon-key thorn) has also been used for furniture andwagons, but it is heavier and darker and sup-plies are limited. This species occurs in about

"

ACoci0 8010inii- rolld

,,,

o

.^

..

g

4

*

*

,,J

I.,

,JR

%

2

sels. Tracheids and fibres: 61: fibres with SIm-

PIe to minutely bordered pits; 66: non-septatefibres present; 70: fibres very thick-walledAxial parenchyma: 79: axial parenchyma va-siCentric; 80: axial parenchyma allform; 81:axial parenchyma lozenge-allform; 83: axialparenchyma confluent; (89: axial parenchymain marginal or in seemingly marginal bands);(90: fusiform parenchyma cells); 91: two cellsper parenchyma strand; 92: four (3-4) cells perparenchyma strand. Rays: 98: larger rayscommonly 4- to 10'seriate; 104: all ray cellsprocumbent; 1/5: 4-12 rays per mm. Mineralinclusions: 136: prismatic crystals present; 142:prismatic crystals in chambered axial paren-chyma cells(M. Thiam, P. Derienne & E. A. Wheeler)

Growth and development Young ACUciogulpinii trees grow fast. In Zimbabwe the av-erage heights of planted trees under conditionsoffrequent drought were 3 in after 3 years and5 in after 9 years, and average hole diameters10 cm after 3 years and 16 cm after 9 years. In2 locations in Madagascar, at 770 in altitudewith a mean temperature of 21'C and an an-nual rainfall of 1150 mm, and at 100 in alti-tude with a mean temperature of 27'C and anannual rainfall of 1600 min, 35-years-oldplanted Acacia gobyiniitrees had a mean holediameter of 26 cm and 32 cm, respectively(maximum 74 cm), and a mean height of 25 in(maximum 30 in). The tree is comparativelylong-lived. It is deciduous, losing its leaves inthe cool season. In the hot season it providesdappled shade allowing some sunlight to pene-trate. Trees may flower when leaness, but theyusually flower while leaves are developing inSeptember-January. Insects such as bees andwasps visit the flowers. The fruits take about 6months to ripen.ACocio gulpinii nodulates well with indigenousRhizobium and Brodyrhizobium bacteria.

Ecology ACUcto galpinii typically occurs inriverme woodland, but it can also be foundscattered on termite mounts and in open wood-land, at 350-1500 in altitude. It is most coin-mon on loamy or clayey soils. Older trees arefrost-tolerant

Propagation and planting Seeds germi-nate easily, but soaking the seeds in hot waterfor one night prior to sowing is advantageousSeeds can be sown in seedling trays filled withriver sand and covered with vermiculite. Seed-

lings generally show good survival, even underregular drought conditions. In experimentalplantings in Zimbabwe seedlings showed a

%

I$i^$4 ;'*;a ',:,$5 1:1;i\ t~ ,.

-*v .-:.. ~

..,

"

ACUcio gobyinii - I, tree herbit, . 2, legb, twig, 3,flowering twig, . 4, fruit.Redrown und udopted by AChmod Sotiri Nur-human

."

,"

the same region as ACocio gulpinii, and mayreach 27 in tall; it has fewer pinnae and leaf-lets per leaf. The chocolate-brown heartwood ofACUcio goetzei Harms is used for construction,heavy joinery and furniture; it is very hard,heavy and durable. ACocio goetzei occurs fur-ther north than ACUci0 801pinii, to Ethiopia,and has fewer pinnae pairs and thicker podvalves

Anatomy Wood-anatointcal description (IAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13:simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;25: intervessel pits small (4-7 pin); 26: in-tervessel pits medium (7-10 pin); 29: vesturedpits; 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; (41: mean tangentialdiameter of vessel Iumina 50-100 prn); 42:mean tangential diameter of vessellumina100-200 pin; 46: s 5 vessels per square milli-metre; (47: 5-20 vessels per square inithmetre);58: gums and other deposlts in heartwood ves-

ACACIA 23

3

.

24 TIMBERSl

survival rate of 86%

Management Under natural conditions, re-generation of ACocio gulpintiis usually good.In suitable habitats, it even acts as a pioneer.YoungAcociogo!pmittrees coppice well.

Yield Being a larger sized ACUcto, ACociogobyinti gives a comparatively high yield ofworkable wood. Individual trees may yield upto 2.5 ing ot'hole wood.

Handling after harvest Large planks canbe sawn from the bole and sometimes sawlogscan also be obtained from the larger branchesof the crown

Genetic resources AlthoughAcociogo!pintiis fairly widespread, it is not very common inmost regions. However, it is not under threat ofgenetic erosion.

Prospects ACUci0 8010iniiis an interestingtimber tree of drier regions. It nodulates welland grows fast, and so it would be a potentialcandidate for testing for afforestation in dryregions, and it is an excellent shade tree forlawns in parks and large gardens

Major references Bolza & Keating, 1972;Brenan, 1970; Coates Palgrave, 1983; Mlambo,Nyathi & Mli10, 2004; Mutshinya10, 2003a;Ross, 1975; Timberlake, Fagg & Barnes, 1999

Other references Bennie at a1. , 2002; deWinter, de Winter & kilnck, 1966; InsideWood,undated; Malan & Roux, 1975; Palmer & Pit-man, 1972-1974; Sutter & Rakotonoely, 1989;Takahashi, 1978.

Sources of illustration Coates Palgrave,1983; de Winter, de Winter & Kilnck, 1966;Palmer & Pitman, 1972-1974.


sidered valuable and is used for construction,local furniture, joinery, flooring, arts andhandicrafts, and for the production of shinglesFormerly it was used for the construction ofsmall boats. The tree is planted as an orna-mental along roads and is a source of nectar forhoney bees

Production and international trade The

commercial production of ACUcio heterophyllotimber is small, with in 1990 about 1200 ing oflogs giving 400 ina of sawn wood (at an averageprice of Us$ 1200/in3), and 480,000 shingles.

Properties The heartwood is pinkish yellowto orange-brown and distinctly demarcatedfrom the paler sapwood. The density is about600 kg/ina at 12% moisture content. At 12%moisture content, the modulus of rupture is154 N/mm2, modulus of elasticity 11,180N/min2, compression parallel to grain 47N/min2, cleavage 26 N/mm and Chalais-Meudon side hardness 2.2. The wood is easy towork. The nailing and gluing properties aregood, and the wood takes varnish well. It takesan excellent polish.

Botany Shrub or smallto medium-sized treeup to 20(-25) in tall; hole often short andcrooked, up to 150(-200) cm in diameter; crownspreading. Leaves in young plants alternateand hipinnately compound with I-5 pairs ofpinnae, each bearing 7-14 pairs of ovate-oblongleaflets 6-10(-20) min long, but soon replacedby elliptical and usually slightly curved phyl-10des 6-16 cm x 0.5-1.5(-2) cm, leathery, withnumerous longitudinal veins. Inflorescence ahead, in small clusters on short axillary shoots,30-40-flowered; peduncle 5-10 min long. Flow-ers bisexual, regular, 5-merous, small, paleyellow, almost sessile; calyx cup-shaped, I-1.5min long, with minute lobes; corolla 2-2.5 mmlong, lobed to halfway; stamens numerous,free, 3.5-4 mm long; ovary superior, ellipsoid,sessile, silky hairy, style c. 10 min long, stigmatubular. Fruit a narrowly oblong, flattened pod6-11 cm x 0.5-I cm, glabrous, brown, 5-10-seeded. Seeds oblong-ellipsoid, compressed, 6-7min long.ACUcio is a large pantropical genus, comprisingmore than 1300 species; most of them distrib-uted in Australia (more than 900), more than200 in America, and about 130 in Africa. ACU-cio heterophy!IQ belongs to subgenus Hetero-phyl!urn, which mainly comprises Australianspecies with phyllodes and without prickles.ACocio melonoxylon R. Br. colackwood) fromAustralia is planted in Reunion and MauritiusIt resembles ACocio heterophylla, but differs in

ACACIAHETEROPHYLLA (Lain. ) Wind.

Protologue Sp. PI. 4(2): 1054 (1806).Family Mimosaceae (Leguminosae - Mimo-

soldeae)Chromosome number2n= 52

Synonyms Rocospermo heterophyllum or, am. )Fedley (2003)

Vernacular names Mountain tamatind (En)Taroarin des hauts, tamarin de in Reunion,ch6ne de Bourbon (Fr).

Origin and geographic distribution Acaciaheterophyllu is endemic to Reunion. It has beenreported from Mauritius, but this is probablyerroneous. It has been planted in centralMadagascar, where it may be naturalized in afew localities

Uses The wood of ACUcio heterophyllo is con-

its phyllodes with reticulate veins, its headsarranged in lax racemes and narrower, twistedpods. Cytogenetic investigations suggest a re-cent autotetraploid origin of ACUcio hetero-phyl!o from ACocio melonoxylon. Another closerelative of ACUcio heterophyllo seems to beACUcio hog A. Gray from Hawaii, of which a fewspecimens have been planted in ReunionGrowth rates of ACUcio heterophyllo seedlingsare high, in good conditions about I in per yearduring 5 years after germination. It is esti-mated that trees need 100 years to reach a bolediameter of 70 cm. In trials in MadagascarACocio heterophyllo succeeded only at 900 inaltitude with an annual rainfall of 1700 mm;

10-year-old trees had a diameter at breastheight of 18 cm and were 18 in tall, but theyhad a bad shape. Rooting is superficial, whichexplains the often severe damage to the treesby cyclones. Flowering is often abundant, andthe flowers are visited by bees.

Ecology Acercio heterophyllo occurs in hu-inId forest at (800-)1000-1800(-2500) in alti-tude, and is locally abundant. The annual rain-fall should be at least 1500 min and divlded

evenly overthe yearManagement Natural germinatlon occurs

after disturbance of the forest, e. g. after fire.Under such circumstances regeneration maybeabundant. Seeds may remain viable for over 30years. Clear felling of the natural forest hasbeen practised to establish pure, even-agedstands of Acercio heterophyl!o as a result ofnatural regeneration. Regular weeding is nec-essary, 2 times per year for the first 3 years, toavoid smothering of the seedlings by weedyplants such as Solonum mountionum SCOPThinning is practised when the trees are 6 and10 years old, to 1250 stemsftia and 800stems/ha, respectively. The eventual densLty tobe reached after 50 years is 200 stemsftia. In1990 the area managed in this way in Reunionwas 1600 ha. Nowadays plantations also existPods are picked from the tree when they arebecoming brown and are dried for two weeks.Before sowing, the seeds have to be pre-treatedby immersion in boiling water for 15 inlnutes.Germination takes place I-3 weeks after sow-ing. Seedlings stay in the nursery for 6-8months before being planted out. Direct sowingin the field is also possible. Recently, in-vitroculture techniques have been developed.The yield of timber per tree in natural forest isoften comparatively small due to the poorshape and short length of the hole; wood de-fects such as knots are common. Proper SIIvi-

cultural practices may improve hole shape andlength, and thus timber yield

Genetic resources and breeding Thetimber of ACocio heterophy!!o has been ex-PIOited for many decades, but important standsstill remain in Reunion, which cover approxi-mately 7500 ha, mainly in natural forest. How-ever, monitoring of the populations of this nar-row endemicremains necessary

Prospects The demand for the timber, espe-cially for use in the local furniture industry, Ismuch greater than the supply. It has been sug-gested that a total area of 3000 ha of well-managed ACocio heterophyllo stands could bedeveloped in Reunion, with an estimated yieldof 4000 in31year of sawn timber. The high inI-tialinvestments needed seem worthwhile inview of the high-quality, valuable timber. Theuse of this indigenous timber tree in Reunionfor the establishment of production forest ispreferable to the planting of exotic species withregard to biodiverslty conservatlon.

Major references Borderes, 1991; MIChon,1998; Pojhjjj, 1990.

Other references Chauvet, 1968; Coulaud,Brown & Sinak-Yakovlev, 1995; du Puy at al. ,2002; Coulaud, Brown & Sinak-Yakovlev, 1992;Reynaud, 1991; Sarrailh, Baret & Rivi6re,2008; Takahashi, 1978


ACACIA 25

ACACLAMELANOXYLONR. Br.

Protologue W. TAIton, Hortus kew. (ed. 2) 5:462 (1813)

Family Mimosaceae (Leguminosae - Mimo-soldeae)

Chromosome number2n=26

Synonyms Rocospermo melonoxylon (R. Br. )Pealey (1986).

Vernacular names Blackwood, Australianblackwood, Tasmanian blackwood (En). Acaciaa bois noir, acacia a fleurs blanches (Fr). Acaciaaustralia, acacia preta (Po)

Origin and geographic distribution ACUciomelono, cylon is indigenous in eastern AUStra-Iia, from Queensland to Tasmania. It has beenplanted in many tropical and subtropical re-

Plantation SIIviculture of ACUciogions.

melonoxylon has a long history in New Zea-land, Brazil and South Africa. It was lntro-duced into South Africa in 1909, and has be-come well established in some natural forestsin the south of the country. It was also intro-duced into Tanzania before the First World

26 TIMBERSl

War. In tropical Africa many small-scale plan-tations exist from Eritrea and Ethiopia to Zim-babwe, Madagascar and Mauritius. In Rwandaabout 2500 ha were planted in the 1980s. ACo-cio melonoxy!on is naturalized in mountainforest fringes in Zimbabwe

Uses Its attractive appearance, fairly eventexture, hardness and good machining proper-ties make ACocio melonoxy!on wood suitablefor high-quality furniture, cabinet making andfancy veneer. In South Africa it became popu-Iar as a substitute for stinkwood (Ocoteo bul-loto (Burch. ) Baill. ). The wood is also used forlight construction, tool handles, musical in-struments, bentwork, turnery and fence posts.It serves as firewood and to produce charcoal.In South Africa ACUcio melonoxy!on is alsoconsidered useful as a nurse tree for the reha.hintation of disturbed natural forest. It is usedfor afforestation in Rwanda and Burundi. ACU-eta melonoxylon is planted for erosion controland in windbreaks, and also as an ornamentaltree. The foliage is browsed by livestock

Production and international trade Theannual production in Australia of ACUciomelonoxylon timber is about 10,000 ms. mainlyfrom Tasmania. It is all used within AustraliaACUcio melonoxylon is a valuable tree in thesouthern part of South Africa, and it has beenestimated that about 1500 ms/year can be bar-vested until 2010 from the remaining oldstands.

Properties The heartwood is golden brownto dark brown or almost black, often with areddish tinge and dark steaks, distinctly de-marcated from the 3-10 cm wide, white tostraw-coloured sapwood. The grain is usuallystraight, sometimes interlocked or wavy, givinga fiddle-back figure; the texture is fine to inod-erately fine. The wood surface is highly Ius-trous.

The wood is medium-weight. The density is515-710 kg/ina at 12% moisture content. Theshrinkage rates are moderately high, 2.0%radial and 6.2% tangentialfrom green to 12%moisture content, and 3.4% radial and 9.0%tangential from green to oven dry. The dryingtlme of the wood is variable; quarter-sawnwood takes twice as long as flat-sawn heart-wood. Boards 25 mm thick take 6-7 weeks todry from 80% to 13% moisture content. It isrecommended that all material, particularly 50mm or thicker, be air dried to 30% moisturecontent before kiln drying. Spring and twistmay be a problem, but can be largely attrib-uted to poor hole shape; distortion can be re-

duced by steam treatment. Even kiln drying at70"C (dry bulb) did not cause excessive surfacechecking. Once dry, the wood is moderatelystable

At 12% moisture content, the modulus of rup-ture is 96-110 N/min2, modulus of elasticity14,130-16,380 N/min2, compression parallel tograin 42-58 N/mm2, shear 12.7-15.3 N/mm2cleavage 76 N/mm radial and 101 N/mm tan-gential, Janka side hardness 4630-5610 N andJanka endhardness 6450 N

The wood is easy to work with hand and ina-chine tools. The polishing properties are excel-lent, and the wood glues and stains well. Thesteam-bending properties are good. The nailingproperties are poor: the wood splits easily andpre-boring is recommendedThe heartwood is moderately durable to dura-ble. It is readily attacked by termites. Thesapwood is moderately susceptible to Lyetusattack. The heartwood is very difficult to treatwith preservatlves, but the sapwood is treatedfairly easily. The pulping characteristics aregoodThe wood is an Jinportant cause of allergic con-tact dermatitis in Australia. QUIDones such asacamelin and hydroxyflavans such as meIaca-cidin are responsible for the sensitizing activ-ity. Occupational asthma caused by blackwoodhas also been reported in wood workersTests conducted in Ethiopia showed high pal-atability and nutritive value of the foliage forsheep.The heartwood contains catechin derivativesthat are potent inhibitors of protein kiriase

Botany Shrub or small to large tree up to35(-40) in tall; hole cylindrical, up to 150 cm indiameter; bark rough, brownish grey to verydark grey, longitudinalIy furrowed and scaly;crown ovoid. Leaves in young plants alternateand hipinnately compound, but soon replacedby narrowly elliptical to lanceolate and usuallycurved phyllodes 4-16 cm x 0.5-2.5 cm, Ieath-ery, with reticulate veins. Inflorescence a headc. 6 min in diameter, arranged in racemes, 30-55nowered; peduncle 5-13 mm long. Flowersbisexual, regular, 5-merous, small, creamy tovery pale yellow, almost sessile; calyx cup-shaped, with smalllobes; corolla c. 2 mm long,lobed to halfway; stamens numerous, free, c. 4mm long; ovary superior, ellipsoid, sessile,style slender, stigma tubular. Fruit a coiledand often twisted, flattened pod 6-15 cm x 48min, glabrous, brown, 6-10-seeded. Seedsbroadly ellipsoid, compressed, 3-5 min long,glossy black

ACUcio is a large pantropical genus, comprisingmore than 1300 species; most of them distrib-uted in Australia (more than 900), more than200 in America, and about 130 in Africa. ACU-eta melono"y!on belongs to subgenus Hetero-phyllum, which mainly comprises Australianspecies with phyllodes and without pricklesIn SouthAfrica mean diameter growth was I-2cm/year until trees were 22 years old. It wasestimated that the mean annual increment ofgross hole volume is about 16 in3ftia and on thebest sites 25 msftia; rotation objectives on goodsites are 40 years for sawlogs. In Rwanda dur-ing the first 12 years of growth, the increase inheight was 1.5-2.4 in/year and in diameter1.6-1.9 cm/year. In Tanzania 20-year-oldplanted trees were 25-32 in tall, with a holediameter of almost 50 cm at breast heightA major problem in plantation establishment isthat young trees often have poor apical doini-nance when grown in fulllight, resulting inlarge numbers of branches and trees of poorhole form. Trees may already flower after 2years. They are usually prolific seed producers.They are long-lived, with ages of over 200 yearsreported from Australia. Nodule-producingRhi20bium bacteria have been isolated fromthe roots.

Ecology In Tasmania ACUciu melonoxy!onoccurs in significant volumes in swamp forest,riverme rain forest and as understorey coinpo-nent of wet eucalypt forest. It shows a wideecological adaptability in mainland Australia.The mean annual temperature in Its range Is6-19'C, mean annual rainfall 750-2300 minIn tropical Africa it is usually planted in coin-paratively cool and wet upland regions, e. g. inTanzania at 1500-2500 in altitude. It growsbest in deep, moist and fertile soils, but growsalso on a wide range of sandy loams and allu-vial soils and in wet, nearly swampy placesSeedlings are shade-intoIerant and ACUciomelonoxylon ls a pioneer species. The seed ger-inmates after disturbances of the forest. InSouth Africa trees with comparatively muchheartwood were found in well-drained areaswith high rainfall and consistently high soilmoisture. ACociu melonoxy!on tolerates frostmoderately well, but planted seedlings are sen-sitive to frost

Management Pre-treatment of seeds jin-proves germination. Mechanical scarification isoptimal, but hot water treatment (3 minutes at90'C) also improves germination, as well asstratification at 4"C for 4 weeks. Pre-treatedseeds can have a germination rate of over 90%

The optimum temperature for germinatlon Is25'C. Inoculation with Rhizobium is recoin-mended in sterilized or new soil. Seeds can bestored for years in airtight containers at roomtemperature or in cool rooms; under naturalcircumstances seeds in the ground can main-tain viahinty up to 50 years. One kg contains55,000-85,000 seeds. A nursery of 2.5 in2 coin-prising 1000 seedlings Is required to plant 0.5-2 ha. Methods of in-vitro propagation havebeen developed in South Africa. Callus fromjuvenile explants of mature trees formed shootson a Murashige and Skoog's medium whensupplemented with growth hormones. Propaga-tion by root cuttings has been applied success-fully. Regeneration by root suckers after fellingis possibleApplication of phosphorus fertilizer at plantingis generally associated with improved earlygrowth; this may be allied to lncreased num-bers of root nodules. In South Africa a singledose of 150 g of superphosphate per tree Is ap-plied at time of planting. In less than 6 monthsseedlings can reach a height of 20 cm, makingplanting out possible. Close initial spacing Isneeded to develop proper hole form, e. g. 2.5 in x2.5 in. Recommended nurse trees are Pinusrodioto D. Don and EUCo!yptus spp. , but resultsare varying. In Rwanda the initial denslty ofplantations was about 2000 treesA1a, with athinning of 50% after 6 years. Pruning is nec-essary to produce good hole form. Thinning to afinal stand of 100-250 stems/ha is practised incommercial plantationsIn South Africa a system was developed inwhich ACUcio melonoxylon was planted in purestands in forest gaps. ACUcio melono, cylon hasbeen considered one of the most important In-vasive plant species in southern parts of SouthAfrica. It does not aggressiveIy invade closedevergreen forest, but outside the forest it is aserious invader of open and disturbed sites.Seed-feeding weevils of the genus Melonteriuscan be used as biological control and have beenimported in South Africa from Australia forthis purpose. For many years a policy of eradi-cation was followed because ACocio melonoxy-10n was perceived as a threat to the indigenousforest, but recent investigations support theopinion that it is gradually replaced by nativespecies in indigenous forest. It seems safe torestrict the control programme to areas whereexotics are inherently unacceptable, e. g. naturereserves. The species is, however, useful bothin relieving pressure on Indigenous timbers,and as a nurse tree for the rehabilitation of

ACACIA 27

28 TIMBERSl

disturbed natural forest. In the 1950s in Zim-

babwe, seeds were scattered deliberately inhigh-rainfall forest reserves at high altitudesto introduce ACocio melonoxylon as a coinpo-nent of the indigenous mountain forest. It stilloccurs there, especially in moist gullies, but notgregariously and is not considered a seriousinvader.

Acacia melonoxylon is susceptible to attack bythe fungus Armingrid meneo. Palatability ofthe young plants to herbivores including live-stock can be a major constraint to successfulplantation establishment. Insect damage totimber in depots is common, but generally oflow intensity and thus not influencing theprice. However, the stored timber is sprayedmonthly with insecticides such as cyperme-thrin. Locally in South Africa, helicopters havebeen used to extract ACUcio melonoxylon tim-her from ecologicalIy sensitive or inaccessibleparts of forest, based on single-tree selection.The high value of the timber compensates forthe high costs

Genetic resources and breeding A studyof alloenzyme variation in Australia indicatedhigh genetic diversity in ACocio melonoxylon.The rate of phyllode development (graduallyreplacing juvenile leaves) appeared to be ge-netically controlled and was negatively corre-lated with height growth in one-year-old seed-lings. The limited research done has shownthat genetic selection may Improve a number ofeconomicalIy important characteristics ofplanted trees. Variation between provenanceshas been reported for wood density, tree size,hole shape, drought and frost tolerance, androot suckering ability. Variation between indi-vidual trees within provenances has been re-ported for percentage heartwood, heartwoodcolour, wood density, bole shape and frost toI-erance. A limited number of seed stocks ofprovenances from throughout the naturalrange of Acacia melonoxylon is maintained atthe Australian Tree Seed Centre of CSIRO,Canberra

Prospects In South Africa ACocio melonoxy-10n is considered a useful, fast-growing timberspecies, which needs to be managed properly toensure that it does not become invasive. It re-quires large forest gaps for optimum timberproduction, while its spread can be controlledby keeping canopy gaps to a minimum. Thelarge genetic variation within ACociomelonoxylon indicates the importance of care-fully selecting genetic material for plantingand the potential value of a tree improvement

programme, in which vegetative propagation isa promising option. However, in tropical Africathe possibilities for economicalIy interestingtimber plantations of ACUcio melonoxylon arelimited due to its ecological requirements, andInferior to those of many indigenous species.

Major references Darrow, 1995; Gelden-huys, 1996; Pinkard & Beadle, 2002; Searle,2000; Seydack, 2003.

Other references Beln at a1. , 1996; Bekele-Tesemma, Birnie & Tengnas, 1993; CAB Inter-national, 2005; Chudnoff, 1980; Mbuya at al. ,1994; Polhi11, 1990; Takahashi, 1978; Timber-lake, Fagg & Barnes, 1999; van Wyk &Gencke, 2000; World Agroforestry Centre, un-dated.


ACACIANiGREscENS Onv

Protologue F1. trop. Mr. a 340 (1871).Family Mimosaceae (Leguminosae - Mimo-


Vernacular names Knobthorn (En)Origin and geographic distribution ACUciu

itigrescens occurs from Tanzania southward tonorth-eastern Namibia, Botswana and north-eastern South Africa

Uses The wood is used for parquet flooring,carving, turnery, fence posts, railway sleepersand mine props. It is occasionally made intofurniture, although it is usually considered tooheavy for this purpose. It is also used as fire-wood and for charcoal production. The foliageis browsed by livestock. Traditional healers ofthe Shona people in Zimbabwe make an oint-merit from burnt roots to treat convulsions. In

Tanzania a root decoction is used as an aphro-disiac. The bark is used for tanning.

Properties The heartwood is dark brown,with light and darker streaks, and distinctlydemarcated from the narrow, whitish yellowsapwood. The grain is often irregular, texturemoderately coarse, even. The wood is veryheavy, with a density of 1000-1200 kg/ms at12% moisture content. The wood seasons

slowly and care is needed to avoid surfacechecking. It is very hard. At 12% moisture con-tent, the modulus of rupture is 126 N/min2modulus of elasticity 14,810 N/min2 coinpres-SIon parallel to grain 73 N/min2, shear 17.5N/min2, Janka side hardness 19,080 N andJanka end hardness 19,080 N. It is difficult tosaw, even when green, and dulls tool edges. A

good finish can be obtained with waxes and oilThe wood has very good natural durability,being extremely resistant to fungal, borer andtermite attack. The heartwood is extremelyreslstantto preservatlve treatment.Promelacacinidins and a variety of tetrahy-droxyflavonoids have been isolated from theheartwood.

Botany Medium-sized deciduous tree up to20(-30) in tall; hole usually straight, up to 75(-90) cm in diameter, usually beset with prickleson large knobs, but these often absent in oldtrees; bark rough, fissured, dark brown toblackish; slash pale to deep pink or red; crownconical in young trees and rounded in maturetrees; branchlets usually glabrous, with pairsof hooked, blackish prickles up to 7 min longjust below the nodes. Leaves alternate, hipin-nately compound, with 2-4 pairs of pinnae;stipules small, caducous; petiole with or with-out gland, rachis sometimes with gland be-tween top I-2 pairs of pinnae; leaflets in I-2pairs per pinna, obliquely obovate-orbicular toobovate-elliptical, (7-)10-35(-50) mm x 7-30(-50) min, apex rounded to emarginate, usuallyglabrous. Inflorescence an axillary spike I-Toe12) cm long, often in fascicles at the base ofyoung shoots; peduncle 0.5-2.5 cm long. Flow-ers bisexual, regular, 5-merous, sinaH, creamywhite but reddish brown in bud, fragrant, ses-sile; calyx cup-shaped, 1.5-2 mm long; corolla2-2.5 min long, with short lobes; stamens rin-meTous, free, 3.5-6 min long; ovary superior,very shortly stalked, style slender. Fruit anoblong pod 6-18 cm x 1.5-2.5 cm, straight,glabrous, dark brown, longitudinaUy dehiscent,3-7-seeded. Seeds flattened orbicular to lens-shaped, 12-13 mm in diameterACocio is a large pantropical genus, comprisingmore than 1300 species; most of them distrib-uted in Australia (more than 9000, more than200 in America, and about 130 in Africa. ACo-

belongs to subgenusit Igrescens

ACUleiferum, which comprises all African ACU-eta species wlth nori-spinescent stipules andhooked prickles. Acacia itigrescens is character-ized by prickles on large knobs on the trunkand the large and few leaflets.Growth is slow, with maximum annual incre-merit of 60 cm in height, usually much less.Acercio itigrescens is completely deciduous andbare for several months. It frequently flowerswhen leaness. The flowers are an importantfood resource to giraffe in the late dry season,and it has been suggested that giraffe could bea pollen vector. Nodulation and nitrogen fixa-

tion have been confirmed.Ecology ACUcio itigrescens occurs in wood-

land and bushland, commonly near rivers, upto 1200(-1600) in altitude. It usually grows onshallow soils on rocky binsides and on alluvialsoils in the valleys. It is often common andIOCaUy dominant on loamy soils. The tree isresistant to fire

Management ACUcio itigrescens has proveneasy to produce in the nursery. The seeds areusually sown into seedbeds, but sowing directlyinto pots is also possible because of a high ger-mination rate. Seedlings suitable for trans-planting have been produced in less than 9months.

Genetic resources and breeding LocallyAcacia itigrescens has been subjected to heavycutting, but in general it is stillfairly common.

Prospects ACocio wigrescens w111 remain asource of durable timber for local use, e. g. forfence poles and flooring. It does not seem to bea promising plantation tree because of its slowgrowth

Major references Brenan, 1970; CoatesPalgrave, 1983; Palmer & Pitman, 1972-1974;Ross, 1968; Timberlake, Fagg & Barnes, 1999.

Other references Bleys, Mazibuko & Allen,1982; Bolza & Keating, 1972; Brenan, 1959;Gelfand at a1. , 1985; Greenway, 1941; Howellat a1. , 2002; Neuwinger, 2000; Takahashi,1978; Williamson, 1955.


ACACIA 29

C, O

ACACIAROBUSTABurch.

Protologue Tray. S. Africa 2: 442 (1824)Family Mimosaceae (Leguminosae - Mimo-

soideae)Chromosome number2n= 26.52Vernacular names Splendid thorn, splendid

acacia, ankle thorn (En). Mungu manzi, ega-inOSema (Po). Mgunga (Sw)

Origin and geographic distribution ACociorobusto occurs from Ethiopia and Somaliasouth to Namibia and northern and easternSouth Africa. It has been introduced elsewhere,

e. g. in South AsiaUses The wood is occasionally used for furni-

ture, shelves and yokes, although its use isrimited because of considerable warping. It isalso used as firewood and among the Minkendapeople of Kenya it is even the preferred type offirewood. The foliage and pods are browsed bylivestock. In traditional medicine, root powderis applied to sweUings, a root decoction is used

30 TIMBERSl

to treat dysmenorrhoea, female sterility andschistosomiasis, a stem bark decoction to treatgonorrhoea, abdominal pains and skin ail-merits, and leaves to treat snakebites. Somemedicinal applications of unknown plant partshave been recorded: to treat malaria and bu-

bonic plague. Acacia robusto is occasionallygrown as a bonsaiplant.

Properties The heartwood is pinkish brownto reddish brown and distinctly demarcatedfrom the wide whitish sapwood. The texture ismoderately coarse to coarse and even. Thewood Is moderately heavy, with a density ofabout 850 kg/ina at 12% moisture contentWarping is often considerable during drying.The wood is brittle and moderately durable,being moderately susceptible to borer and ter-mite attack; the sapwood is susceptible to sapstain. The pulping properties of the wood havebeen rated as good. The energy value of thewood is 18,100 kJ/kg.The crude protein content of the seeds is12.5%. The gum from the bark contains 18% ofprotein bound to an arabinogalactan. In screen-ing tests root bark of ACocio robusto showedstrong antibacterial activity against Stophylo-coccus aureus and moderate activity againstESCherichio coli, but these activities were notconfirmed in a later test.

Botany Shrub or small to medium-sized treeup to 20(-25) in tall; root system moderatelydeep, with strongly spreading lateral roots;bole up to 70 cm in diameter; bark smooth tofissured, grey-brown to dark brown or black;crown spreading; young branches smooth, withpaired, straight stipular spines up to 6(-11) cmlong. Leaves alternate, bininnately compound,with 2-6(-10) pairs of pinnae; rachis 2.5-7 cmlong, pubescent or glabrous; leaflets in 9-27pairs per pinna, obliquely oblong, (2-)3.5-13(-16) min x I-5(-7) mm, glabrous or ciliate onthe margins. Inflorescence an axillary globosehead 9-12 mm in diameter, usually in clusters;peduncle 2-2.5 cm long, with a pair of smallbracts in basal half. Flowers bisexual, regular,usually 5-merous, small, creamy white, fra-grant, sessile; calyx cup-shaped, with shortlobes; corolla shortly lobed, glabrous; stamensnumerous, free, up to 5 mm long; ovary supe-nor, I-celled, style slender. Fruit a linear pod7-19 cm x 0.5-3 cm, straight or curved, moreor less woody, glabrous, brown, longitudinalIyveined, dehiscent, up to 15-seeded. Seeds quad-rangular to ellipsoid, compressed, 8-15 mm x5-9 mm, smoothACocio Is a large pantropical genus, comprising

more than 1300 species; most of them distrib-uted in Australia (more than 900), more than200 in America, and about 130 in Africa. ACo-cm robustu belongs to subgenus ACUcio, whichcomprises all African ACUciu species withstraight spinescent stipules. ACUcio robusto isvariable, and is subdivided into 3 subspeciessubsp. cloutgero (E. Mey. ) Brenan (synonymAcercio cloutgero E. Mey. ), which may developinto a medium-sized tree and is characterized

by its pubescent leaf rachis and curved, coin-paratively thin pods; subsp. robusto, whichremains a small tree (up to 8 in tall) and ischaracterized by its glabrous leaf rachis andstraight, comparatively wide pods; and subsp.usQinbore, 1sts (Taub. ) Brenan (synonym: ACU-cio usumborensis Taub. ), which is character-ized by its glabrous leafrachis and very narrowpods. Intermediates between the subspeciesare rather common.

ACocio robusto grows comparatively fast. Treesare usually deciduous for a short period. Inno-rescences appear together with or after thenew leaves. Dispersal of the seeds is probablyby browsing animals. ACUcio robusto is nodu-lated by nitrogen-fixing Rhizobium bacteria. InAustralia it is considered invasive

Ecology ACUcio robusto occurs in woodlandand wooded grassland, often near rivers, wherelarge specimens can be found, up to 1800 inaltitude. The tree is drought and frost resis-tant

Management The germination rate of un-treated seeds can be very low, often only about3%; it is higher when the seeds have been in-gested by herbivores. Mechanical scarificationmay Improve germination to over 90%, andtreatment with boiling water or sulphuric acidmay enhance germination to over 80%. Bruchidspecies may attack the seeds, especially whenthese are on the ground. Therefore, seeds forpropagation should be collected from the can-opy and the period of storage of untreatedseeds should be minimized. Trees resprout wellafter coppicing.

Genetic resources and breeding Acerciorobusto is widespread and locally common andnot in danger of genetic erosion

Prospects The wood of Acacia robusto isconsidered of rather poor quality and of limiteduse only. Some types are spectacular whenflowering and may be good garden trees, evenin frost-prone regions.

Major references Brenan, 1970; CoatesPalgrave, 1983; Palmer & Pitman, 1972-1974;Timberlake, Fagg & Barnes, 1999.

~I

Other references Beentje, 1994; Brenan,1959; Churms & Stephen, 1984; Khan, 2001;Khan at a1. , 1980; Kokwar0, 1993; Luoga, Wit-kowski & Balkwi11, 2004; Neuwinger, 2000;Pakia & Cooke, 2003a; Robbertse, 1974.


ACACIA RowMAE 011v

Protologue F1. trop. Mr. 2: 353 (1871).Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names Mgunga(Sw)Origin and geographic distribution ACocio

70uumoe occurs in coastal regions of southernSomalia, Kenya and Tanzania, possibly alsonorthern Mozambique, and in southern andwesternMadagascar

Uses In Tanzania the wood is used for house

building and pestles, as firewood and for theproduction of charcoal. In Madagascar it isused for making furniture. Twigs are given asfodder to livestock in Madagascar.

Properties The heartwood is reddish brownand nicely marked, moderately heavy andhard. It is resistant to termite attack.

Botany Small to medium-sized tree up to20(-30) in tall; hole up to 130 cm in diameter;bark rough or smooth, grey-green to dark greyor dark brown, often with thick, woody prick-Ies; crown open, flat; branchlets shortly hairy,with pairs of straight or only slightly curved,blackish prickles 4-8 mm long just below thenodes. Leaves alternate, hipinnatelypound, with (3-)4-9 pairs of pinnae; stipulessmall, caducous; petiole (I-)2-4 cm long, withI-2 glands, rachis 1.5-7 cm long, with glandsbetween top pairs of pinnae; leaflets in (5-)8-31 pairs per pinna, obliquely oblong, 4-9(-12)min x 1.5-4 mm, shortly hairy especially be-neath, bluish green. Inflorescence an axillaryspike 6-10 cm long; peduncle I-3.5 cm long.Flowers bisexual, regular, 5-merous, small,creamy white but with reddish calyx, fragrant,almost sessile; calyx cup-shaped, 1.5-2 minlong, hairy; corolla 2-3 min long, with shortlobes; stamens numerous, free, 4-6 min long;ovary superior, shortly stalked, style slender,up to 5 mm long. Fruit an oblong pod 5-15 cmx 1.5-3 cm, flat, straight, woody, nearly gla-forous, dark brown, indehiscent, c. 9-seededSeeds oblong to disk-shaped, 8-13 mm x 7-9

found in Australia (more than 900), more than200 in America, and about 130 in Africa. ACo-cio rouumoe belongs to subgenus ACUleiferum,which accommodates analrican Acacia specieswith non-spinescent stipules and hooked prick-Ies. ACocio rouumoe resembles ACocio burheiBenth. , which has more strongly hooked prick-Ies, less numerous leaflets and narrower, de-hiscent podsACUci0 70uumoe flowers when new leaves de-velop. The indehiscent pods are possibly dis-persed by water

Ecology ACoci0 70uumue occurs In riverIneforest and swamp forest near the coast, often atthe inner margin of mangroves, in Madagascaralso in deciduous woodland and scrubland oncalcareous soils, usually at lower altitudes butin East Africa occasionally up to 700 in alti-tude.

Management The trees coppice readily andcan be managed for the production of branchesused as fodder

Genetic resources and breeding AlthoughACUcio rouumoe is not very widespread, thereare no indications that it is threatened by ge-netlc eroslon

Prospects Too little is known about Acerciorouumoe to evaluate its prospects as a timbertree of wider use. With its rather specific habi-tat requirements, it seems unlikely that it w111gain linportance

Major references Brenan, 1959; du Puy ata1. , 2002; Ratsinarson at a1. , 2000

Other references Beentje, 1994; Brenan,1970; Kernp, 1951


ACACIA 31

coin-

mm.

ACUcio Is a large pantropical genus, coinprislrigmore than 1300 species; most of them are

ACACIAXANTHOPHLOEABenth.

Protologue Trans. Linn. SOC. London 30(3)511 (1875).



Vernacular names Fever tree, sulphur bark,African thorn acacia (En). Arbre a fibvre (Fr).Arvore da Iebre, cainba (Po). Mgunga, inukonge(Sw)

Origin and geographic distribution ACocioxonthophloeo occurs from Somalia south tonorthern South Africa and Swaziland. It isoccasionally planted as an ornamental treeoutside its natural distribution area, e. g. in theUnited States and Taiwan

Uses ACUciu runthophloeo produces a gen-

.

32 TIMBERSl

.

Acercio tonthophloeo - wild

eral-purpose timber, which is used in construc-tion and for carpentry, boat building, boxwood,furniture, mortars, domestic utensils, troughsand fence poles. The wood is used as firewood,although the gum leaves a black tar-like de-POSit when burnt. It makes good quality char-coal. ACocio tonthophloeo is a popular gardentree and has been grown in hedges in Tanza-nia. In Kenya a bark decoction is used in tradi-tional medicine to treat indigestion, and inTanzania to treat sickle cell anaemia. In SouthAfrica Zulu people use powdered bark as anemetic to treat malaria, and also against eyecomplaints. The gum, which occurs in largequantities on the trunk, is reportedIy edibleand is eaten by monkeys. Leafy branches aregiven as fodder to livestock. Trees produce goodbee forage.

Production and international trade ACociorunthophloeo bark is traded on local marketsin South Africa, with an estimated trade vol-ume of 7500 kg in 1988. It is also importedfrom Mozambique

Properties The heartwood is pale brownwith a reddish tinge and distinctly demarcatedfrom the wide, paler sapwood. It is fairlyheavy, with a density of about 900 kg/in3 at12% moisture content. It should be dried with

care because it is prone to splitting and crack-ing. It finishes smoothly. The wood is suscepti-ble to borer attack

The bark contains tannin in considerable quan-tity, but it does not produce good leather. Barkextracts showed weak in-vitro antimalarialactivity and considerable antimicrobial activ-ity. Leaf extracts showed antibacterial activityagaJnst Stophylococcus oureus and ESChertchio

coli. The gum is water soluble and containsgalactose, arabinose, rhamnose, glucuronic acidand 4-0-methyl-orD-glucuronic acid

Description Medium-sized tree up to 25 intall; hole straight, up to 60 cm in diameter; barksmooth and powdery, lemon yellow to greenishyellow; crown open, with spreading branches;young branches first purple then yellowish,with paired, straight stipular spines up to 7(-8.5) cm long. Leaves alternate, hipinnatelycompound, with 3-6(-8) pairs of pinnae; rachis(2.5-)3-7 cm long, almost glabrous; leaflets in8-19 pairs per pinna, obliquely oblong, 2.5-6.5min x I-2 min, glabrous. Inflorescence an axil-Iary globose head c. 12 mm in diameter, usu-ally in clusters on older twigs; peduncle 1.5-2.5cm long, with a pair of bracts c. 3 mm long inbasal half. Flowers bisexual, regular, usually 5-merous, small, creamy white to pinkish or pur-PIish or yellow, fragrant, sessile; calyx cup-shaped, I-1.5 min long; corolla shortly lobed,glabrous; stamens numerous, free, up to 5 mmlong; ovary superior, I-celled, style slenderFruit a tinear-oblong pod 4-13.5 cm x 0.5-1.5cm, straight or slightly curved, pale brown,constricted between the seeds, indehiscent but

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breaking up into segments when fanen, (3-)5-10-seeded. Seeds orbicular to ellipsoid, coin-pressed, 4.5-5.5 min x 3.5-4 min, smooth, brown.

Other botanical information ACocio is a

large pantropical genus, comprising more than1300 species; most of them distributed in AUS-tralia (more than 900), more than 200 in Amer-ica, and about 130 in Africa. ACocio xontho-phioeo belongs to subgenus ACUcio, which coin-prises all African ACocio species with straightspinescent stipules. ACUcio xonthophloeo isvery characteristic with its smooth, yellow barkand indehiscent pods.

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; (23: shape of alternate pits po-Iygona1); 25: intervessel pits small (4-7 prn);26: intervessel pits medium (7-10 prn); 29:vestured pits; 30: vessel-ray pits wLth distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;47: 5-20 vessels per square minimetre. Tra-cheids and fibres: 61: fibres with simple to inI-nutely bordered pits; 66: non-septate fibrespresent; 70: fibres very thick-walled. Axial pa-renchyma: 78: axial parenchyma scanty para-tracheal; 79: axial parenchyma vasicentric; (80axial parenchyma allform); (81: axial paren-chyma lozenge-alitorm); 83: axial parenchymaconfluent; 90: fusiform parenchyma cells; 91:two cells per parenchyma strand. Rays: 98:larger rays commonly 4- to 10-senate; 104: allray cells procumbent; 1/5: 4-12 rays per minMineral inclusions: 136: prismatic crystalspresent; 142: prismatic crystals ln chamberedaxial parenchyma ceUs; (156: crystals inenlarged cells)Obj. Thiam, P. Derienne & E. A. Wheeler)

Growth and development The growth rateof seedlings is fast, to a maximum of 7 in tallin3 years, A growth rate of 1.5 in/year and 2 cmin diameter are common in young trees. Thetree is deciduous. The flowers are pollinated byinsects such as bees and butterflies. In south-ern Africa flowering occurs in September-November, fruiting in January-April. ACocioxonthophloeo suffers stripping of bark, brows-ing and breakage by elephants, but exhibitshigh resilience to disturbance. The pods are afavourite food of vervet monkeys, which mayplay a role in seed dispersal. The light podsegments are probably also dispersed by wind

and water.

Ecology ACUcio xortthophloeo occurs in swam-py localities and riverme forest, often on floodplains or in depressions, up to 2100 in altitudeIt often grows on alluvial black clay soils. Thispreferred habitat, which offers ideal breedingpossibilities for malaria mosquitoes, is the rea-son that ACocio runthoph!oeo was associatedwith malaria; hence the name 'fever tree'. Itoften grows gregariously and is often evendominant. The tree can tolerate moderate frostIt is nodulated by nitrogen-fixing Rhizobiumbacteria.

Propagation and planting Seed produc-tion is often poor as a result of predation byanimals. There are 24,000-30,000 seeds per kg.Germination is generally fair, reachlrig about70% after 2 weeks. Seed should be soaked inwater for 24 hours, in hot water overnight, orbe mechanicalIy scanfied. Seeds can be storedfor a long time in a dry place, but they are sus-ceptible to insect damage. When seedlings havereached the 2-1eaf stage 6-8 weeks after sow-ing, they should be transplanted from seedlingtrays into nursery bags. Care should be takennot to damage the long taproot. ACocio xontho-phloeo can be propagated by cuttings

Management ACocio runthophloeo often oc-CUTS in apparently even-aged stands of largetrees with few young plants, probably as a re-SUIt of prolific regeneration after flooding inthe past. Natural stands of ACUcio runtho-phloeo in South Africa comprise about 85sterns of allsize classes per ha

Harvesting The bark is removed with aknife and bark collectors usually focus on Iar-ger-sized trees. The tolerance of ACocio Ton-thophloeo to damage is high, and usually treesrecover from bark removal as well as from ele-

phant damage. However, excessive destructiveharvesting is locally common

Genetic resources In many riverIne localI-ties Acacia xonthophloeo occurs gregariouslyHowever, in many regions riverme woodlandshave declined as a result of clearing for agricul-ture, changes in climatic conditions and soilsannity, and because of tree damage by grow-ing elephants populations and overgrazing bylivestock. Locally, large ACUcio ranthophloeotrees have been felled for timber. In South of-rica there is considerable pressure on ACUciorunthophloeo populations because of excessivebark harvesting in a non-sustainable way

Prospects Acercio runthoph!oeo is a multi-purpose tree that is important for people as asource of timber, firewood, medicine and for-

ACACIA 33

34 TIMBERSl

age. It is a characteristic element of rivermehabitats in eastern and southern Africa, apopular tree amongst birds for nest buildingand an important food source for animals suchas vervet monkey and elephant, and as such itplays an important ecological role. Monitoringof natural populations is needed, and cultiva-tion of this fast-growing species should be en-couraged. In regions where bark is harvestedfor medicinal purposes on a large scale, meth-ods of sustainable harvesting should be devel-oped and promoted. After harvesting, barkshould be given sufficient time to grow backWith its open crown ACUcio xonthophloeo is apromising plantation tree in agroforestry sys-terns, and its popularity as an ornamental treeis also likely to increase.

Major references Botha, Witkowski& Shack-leton, 2002; Brenan, 1970; Coates Palgrave,1983; Hankey & Stern, 2002; unuya at al. ,1994; Palmer & Pitman, 1972-1974; Timberlake,Fagg & Barnes, 1999; van Wyk, 1972-1974

Other references inderson, Bridgeman &De Pinto, 1984; Beentje, 1994; Brenan, 1959;Chhabra & Uis0, 1991; Grace at a1. , 2002b;InsideWood, undated; Katerere & Eloff, 2004;Mugedo & Waterman, 1992; Neuwinger, 2000;Odee at a1. , 2002; 0tieno at a1. , 2005; Frozesky,Meyer & Louw, 2001; Thulin, 1993.

Sources of illustration Timberlake, Fagg& Barnes, 1999


oblong to broadly elliptical, up to 3 cm x 1.5cm, rounded at base and apex, almost glabrousInflorescence an axillary raceme (5-)10-20 cmlong, many-flowered. Flowers bisexual, regular,5-merous, yellow; pedice1 2-4 min long, withjoint near base; calyx shortly obconica1, 1.5-2mm long, glabrous, with short lobes; petalsfree, elliptical, 3-4 mm long, reflexed; stamens10, free, arithers with a stalked gland at apex;ovary superior, shortly stalked, narrowly ob-long, glabrous, style c. 3.5 min long. Fruit anarrowly oblong to linear and sickle-shapedpod (10-)20-30 cm x 1.5 cm, flattened, stalked,with thickened margins, glabrous, yellowish,silky, dehiscent with 2 valves, up to 15-seeded.Seeds flattened ellipsoid, 8-11 min x 6-8 mm,black in the lower part and red-orange in theupper partAdenonthero comprises about 12 species, whichare allindigenous to tropical Asia and northernAustralia, except Adenonthero montoroo. TwospecLes are occasionally planted as an orna-mental and roadside tree in tropical Africa:Adenonthero microspermo Tensm. & Binn. andAdenonthero PODonino L. The latter speciesoccurs occasionally naturalized in Madagascar;it differs in its leaflets that are pubescent be-low and in its completely red mature seeds.

Ecology Adenonthero montoroo occurs inevergreen forest up to 1100 in altitude. It isfound on latentic soils.

Genetic resources and breeding In gen-eral Ade, 10nthero montoroo is uncommon, butit is locally abundant, e. g. around the Bay offutongil

Prospects Very little is known about thisrecently described species. In view of its scat-tered occurrence and restricted distribution,harvesting the timber from wild stands shouldbe discouraged. Planting experiments mayclarify the possibilities as a plantation timber.The wood properties should be investigatedand might be good, as is the case with severalAsiaticAde, 10nthero species

Major references du Puy at a1. , 2002.Other references Viniers, 1995Authors R. H. M. J. Lemmens

ADENANTHERAMANTAROAVilliers

Protologue Bull. Mus. natl. Hist. nat. , sect.B, Adansonia 16: 227 (1995).

Family Mimosaceae (Leguminosae - Mimo-soideae)

Origin and geographic distribution Ade-norithero montoroo is endemic to Madagascar,where it occurs in the northern and easternparts

Uses The wood is used locally for construc-tion and canoes

Botany Medium-sized to fairly large tree upto 30 in tall; bole straight, up to 100 cm in di-ameter; bark scaly, reddish brown; youngbranches sparsely pubescent. Leaves alternate,hipinnately compound with (3-)4-6 pairs ofpinnae, these alternate to opposite; stipulestriangular, c. 2 mm long, caducous; petiole(2.5-)4-9 cm long, rachis (4-)6-22 cm long,grooved and slightly pubescent above; leaflets11-15 per pinna, alternate, shortly stalked,

AESCHYNOMENEEiAPHROXYLON (Gulll. &Perr. ) Taub.

Protologue Engl. & Prantl, Nat. Pnanzen-fom. 111, 3: 319 (1894)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

I


Synonyms Herminieru e!uphroxylon Gulll& Perr. (1832)

Vernacular names Ambatch, ambach, pithtree, balsa wood tree (En)

Origin and geographic distribution Aes-chynomene elophroxylo, " occurs widely in tropi-cal Africa, from Senegal east to Ethiopia andKenya and south to Zambia, Angola and Mo-zambique; also in Madagascar, but thereprobably introduced. It has been introduced inEgypt, Indonesia and SriLanka

Uses The lightweight wood is used for ca-noes, rafts and shields, but also for buildingpoles and furniture. The stems are used forfishing-net floats and sandals. The wood isattached to fishing spears to ensure they reap-pear at the water surface in case of a miss, ormay be attached to spears to serve as a buoy

Properties The pale coloured and spongywood is very light, with a density of 160-190kg/ina at 15% moisture content

Botany Large shrub or smalltree up to 9(-12) in tall; stems swollen, pithy, glandularhairy and with woody prickles up to 1.5 cmlong, often with dense mats of adventitiousroots; outer bark green, smooth. Leaves alter-nate, panpinnately compound with 9-20 pairsof leaflets; stipules broadly ovate, 10-13 mmlong, auricled on one side; petiole and rachistogether 4-16 cm long, densely bristly hairyand with short prickles; leaflets oblong, 8-26min x 4-10 mm, obliquely rounded at base,truncate or slightly notched at apex, glabrousabove, shortly hairy below. Inflorescence anaxillary raceme up to 4.5 cm long, I-4-flowered; bracts ovate, up to 7 mm long, brac-teoles broadly ovate, up to 16 mm long. Flowersbisexual, papilionaceous; pedice1 12-16 minlong; calyx glandular hairy, 2-11pped, the upperlobe 2-2.5 cm long and entire to slightlynotched, the lower lobe 1.5-2 cm long and shal-lowly 3-toothed; corolla with rounded, notched,orange yellow standard up to 4.5 cm in diame-ter and yellow wings and keel; stamens 10,fused into a sheath split into 2 groups of 5;ovary superior, I-celled, style slender, up-turned. Fruit a spiralIy coiled pod 10-14 cmlong when unrolled, glandular hairy, 6-17-jointed. Seeds kidney-shaped, up to 8 min indiameter, dark purplish brownAeschynomene comprises about 150 species,tropical America and Africa being richest inspecies, with a few species in tropical Asia.Continental Africa has about 50 specles,Madagascar 13. The stems and wood of some

other Aeschynomene species Is used for similarpurposes as those of Aeschynomene elophroxy-10n.

The stems of Aeschynomene crossicoulisHarms, a shrub with procumbent stems occur-ring from Senegal east to DR Congo, are usedas floats for fishing nets and lines, and as fueland fodder, whereas the leaves are added to amixture of other plants to treat rheumatism.The stems of Aeschynomene cristoto Vatke, ashrub widespread in tropical Africa includingMadagascar, are also used as floats for fishingnets and in raft and canoe construction, and

parts of the plant are applied to skin coin-plaintsAeschynomene PIundii Taub. closely resemblesAeschynomene elophroxylon, but is a smallershrub without prickles. It occurs at watersidesfrom Mali east to Kenya and south to Zambia.Its stems are used as floats for fishing netsAeschynomene @10phroxy!on ls adapted to fluc-tuating water levels. At the base of the holeunbranched roots up to 10 cm long are found aswell as larger roots of unlimited length growthThe plants root superficially Into the soil, andwhole thickets are frequently displaced in thewater by wind or currents. Numerous hemi-spherical protuberances are situated on thestem bark containing nitrogen-fixing bacteriaof the genus Brodyrhizobium. Aeschynomeneelophroxylon usually flowers at the beginningof the rainy season

Ecology Aeschynomene elophroxylon occursat lakesides and in pools and swamps, up to1350 in altitude. The lower part of the stem lsoften submerged, andAeschynomene elophroxy-10n is well suited to seasonalIy flooded localI-ties where the soilremains wet. The stems andbranches may form dense floating masses thatclog watercourses.

Genetic resources and breeding Aes-chynomene e!uphroxy!on is widespread andlocally common and not endangered by geneticerosion, although locally It has disappeareddue to changed climatic conditions, e. g. in InOStparts of Lake Chad

Prospects The wood of Aeschynomeneelophro, cylon and other Aeschynomene specieswill remain of local importance because of itsgreat floating ability. It could be a good substi-tute for the wood of balsa (Ochromo pyromi-dole (Cav. ex Lain. ) Urb. ) for special applica-tions such as model making. Aeschynomeneelophro, cylon deserves more attention as a nT-trogen fixing tree that is potentially useful forenhancing soil fertility in temporarily flooded

ABSCHYNOMENE 35

36 TIMBERSl

agricultural land, e. g. as a green manure inrice cultivation

Major references Arbonnier, 2004; Burkill,1995; du Puy at a1. , 2002; Jenik & Kubikova,1969

Other references A1azard, 1991; A1azard &Duboux, 1988; Beentje, 1994; Eggeling & Dale,1951; Ita, 1994; Raponda-Walker & Sillans,1961.


AFROCARPusFALCATus (Thunb. ) GNPage

Protologue Notes Roy. Bot. Gard. Edin-burgh 45: 383 (1988)

Family PodocarpaceaeChromosome number2n=24

Synonyms Podocorpus 101cotus (Thunb. )R. Br. ex Mirb. (1825), Nogeio Idledtus (Thunb. )Kuntze (1891), Podocarpus grocilior Pilg.(1903), Decussocurpus Idledtus (Thunb. ) deLaub. (1969), Decussocorpus grocilior (Pilg. ) deLaub. (1969), Afrocorpus grocilior (Pilg. )GNPage (1988)

Vernacular names Yenowwood, African fernpine, weeping yew, Outeniqua yellowwood (En)Bois jaune, pin foug6re africain (Fr)

Origin and geographic distribution Afro-corpus falcotus occurs in mountain forest fromEthiopia through Kenya, Tanzania and Mo-zambique to eastern and southern South at-rica; also in Swaziland and Lesotho. It isplanted as ornamental tree in South Africa,and occasionally outside its natural area ofdistribution, e. g. in Australia, the UnitedStates and India

Uses The wood, often traded as 'podo' or

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'yellowwood', is highly valued for ship building,e. g. for masts and planks, but it is also used forpoles, panelling, furniture, boxes, veneer andplywood. It is suitable for construction, floor-Ing, Joinery, Interior trim, mine props, vehiclebodies, railway sleepers, toys, novelties, agri-cultural implements, musical instruments, foodcontainers, vats, turnery, hardboard and parti-cle board. It is also used as firewood.

The ripe seed is edible, but resinous. Edible oilhas been extracted from the seeds in EthiopiaThe bark and seeds are used in traditional

medicine. Bark decoctions or infusions are used

as anodyne, also to treat stomach-ache. A barkdecoction is also applied to itching rash. Pul-verized seeds are applied to treat tuberCUIoidmeningitis and sunburn. In Ethiopia the seedoilis used in the treatment of gonorrhoea. Thebark has been used for tanning, although itonly contains 3-6% tannin. Afrocorpus foieotusis planted as ornamental and roadside tree;sometimes it is also used as container plantand Christmas tree. It is very useful for soilprotection against water erosion. It is alsoplanted as windbreak


international trade in Afrocorpus falcotus tim-her is very limited. The volumes sold by auc-tion in South Africa in the period 1996-1998varied from 16 mat0 33 ms. In 2006 the price of2.5 cm thick boards was 2280 Us$/ina.

Properties The heartwood is pale yellow topale yellowish brown, and not distinctly de-marcated from the sapwood. The grain isstraight, occasionally spiralIy, texture fine andeven. Reddish streaks of compression wood anddarker lines resulting from year rings may bepresent. Resin is absent, and the wood has nodistinctive odour

The wood is moderately lightweight, with adensity of 430-560(-620) kg/ms at 12% ino1s-ture content. It usually air dries without prob-Iems, but surface checking, splitting and dis-tortion may occur; close stacking an. d heavyweighting recommended. Kiln dryingshould be done at lower temperatures. Therates of shrinkage from green to oven dry areabout 3.6% radial and 5.4% tangential. Oncedry, the wood is stable in serviceAt 12% moisture content, the modulus of rup-ture is 50-68(-82) N/min2, modulus of elastic-ity 7600-8900 N/mm2, compression parallel tograin 35-42 N/min2, compression perpendicu-Iar to grain 6 N/min2, shear 7-8 N/min2 Jankaside hardness 2840-3740 N and Janka endhardness 4000-4930 N.

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The wood is easy to saw and work, both by ina-chine and hand tools, with little dulling effectson cutting edges. It can be planed to a smoothfinish. The wood has a tendency to split uponnailing, and pre-boring is recommended. Thegluing, painting, varnishing and staining prop-erties are moderately good. Steam bendinggives moderate to satisfactory results. Turninggives good results. The peeling properties aregood; good-quality veneer can be produced fromthe wood. The wood is not durable, being sus-ceptible to blue stain, powder-post beetle, pm-hole borer, longhorn beetle, termite and marineborer attacks. It is permeable to preservatives.The wood is resistant to acids

Podolide, a norditerpene dilactone with In-vitroantileukemic properties, has been isolated fromleaves of Afrocorpus falcotus. The tubulin-binding diterpene taxol has been isolated insmallamounts from young stems and leaves.

Adulterations and substitutes The wood

of Podocurpus lotifolius (Thunb. ) Mirb. closelyresembles that of Afrocorpus folcotus and isused for similar purposes; it is also known asyellowwood

Description Evergreen, dioecious, medium-sized to large tree up to 45G60) in tall; holebranchless for up to 25 in, straight and eviln-

drical, up to 200(-300) cm in diameter; barksurface greyish brown to reddish brown, rathersmooth, flaking in irregular pieces; twigsridged by decurrent leaf bases. Leaves ar-ranged spiralIy, simple and entire; stipulesabsent; petiole indistinct, short, twisted by c90' blade narrowly linear-lanceolate to linear-elliptical, (I-)2-4(-4.5) cm x (1.2-4(-6) mm,curieate at base, acute to acuminate at apex,leathery, glabrous, glaucous green to yellowishgreen, wLth a single main vein and single resincanal. Male cone axillary, solitary or morerarely in groups of 2-4, nearly sessile, 5-15min x c. 3 mm, brownish; scales many, ar-ranged spiralIy, each bearing 2 pollen sacsFemale cone terminal on short, leafy or scalybranchlet, solitary, with a single fertile scale.Seed drupe-like, globose to obovoid, 12-18 minlong, glaucous to greyish green, seed coat hard,woody, warty, enclosed in a somewhat fleshy,resinousintegument

Other botanical information Afrocorpuscomprises I-4 species, depending on the spe-cies concept, and is restricted to mainlandtropical Africa. It has been separated from Po-docorpus in 1988, mainly based on the absenceof a fleshy receptacle at the base of the seedand on chromosome number, but there is stilldisagreement about this. However, moleculardata support the recognition of Afrocorpt, s asdistinctfrom Podocorpt, s.On the basis of more slender leaves Afrocorpusgruci!tor (Pilg. ) C. N. Page from East Africa issometimes kept separate from Afrocorpus 101-cottts sensu stricto, which is then restricted tosouthern Africa.

Afrocorpus usQinborensis (Pilg. ) GNPage(synonyms: Afrocorpus dowei(Stapf) C. N. Page,Podocorpus dowei Stapf, Podocorpus usQin-borensis Pilg. ) occurs in mountain forest fromeastern DR Congo to Kenya and Tanzania. hisclose to Afrocorpus foieotus, differing in itsleaves more abruptly tapering at apex andslightly larger seeds with thicker woody part ofseed coat. It is used for SImlIar purposes asAfrocorpus falcotus; its wood is similar butmore lightweight, with a density of about 420kg/ina at 12% moisture content. The nomencla-ture is confusing and in need of clarificationPodocorpus usermborensis has been consldereda synonym of Afrocorpus monnii (Hook. f. )C. N. Page (synonyms: Decussocorpus monniz(Hook. f. ) de Laub. , Nogeio monnii (Hook. f. )Kuntze, Podocarpus monnit Hook. f. ), whichhas been described as an endemic from SaoTome and is planted in southern Nigeria and

AFROCARPUS 37

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38 TIMBERSl

western Cameroon. Some authors considerAfrocorpus as comprising only a single, van-able species.

Anatomy Wood-anatomical description orAWAsoftwood codes):Growth rings: (40: growth ring boundaries dis-tinct); (41: growth ring boundaries indistinct orabsent); 43: transition from earlywood to late-wood gradual. Tracheids: 44: tracheid pittingin radial walls (predominantly) uniseriate (ear-Iywood only); 54: latewood tracheids thin-walled (double wanthickness less than radiallumen diameter); (56: torus present (pits inearlywood tracheids only)). Axial parenchyma72: axial parenchyma present; 73: axial paren-chyma diffuse (evenly scattered throughouttheentire growth increment); (74: axial paren-chyma tangentially zoriate); 76: transverse endwalls smooth. Ray composition: 80: ray tra-cheids absent or very rare; 82: cell walls of raytracheids dentate. Cross-field pitting: (91:cross-field pits pinoid); 94: cross-field pitstaxodioid; 98: I-3 pits per cross-field (early-wood only). Ray size: 103: average ray heightmedium (5-15 cells).(P. Baas & I. Hemz)

Growth and development In South Africatrees reached a height of up to 8.2 in and holediameter of up to 14 cm 11 years after plant-ing. On fertile soils in Rwanda the annualgrowth can be over I in in height and ICm inbole diameter, at least until 15 to 20 years old.The tree is very sensitive to competition, andthinning is necessary for good diametergrowth.The pollen is dispersed by wind, but most of itdoes not get far from the male tree. Fruitingstarts when the tree is 10 years old. Seed pro-duction varies from year to year. Usually, treesbear seeds irregularly, mostly at intervals of2-4 years. Seeds take about one year to de-velop. They remain on the tree for a consider-able time. Fruit bats are the main dispersers;they feed on the fleshy part of the seed coat,discarding the woody part with the embryoColobus monkeys feed on the seeds, but alsorodents, bushpig and large birds such as horn-bills and turacos. However, it has been re-ported that seed that has passed the digestivetract of these animals usually does not germi-nate anymore. Seedlings seem to establish onlyafter mast seed years, and usually close to theparent tree. Investigation of the roots revealedthe presence of arbuscular mycorrhizae. Treesmay reach a very high age, up to 600 years

Ecology In East Africa Afroco, pus Idledtus

the mountains atforestOccurs In In

1500-2400(-3000) in altitude, often associatedwith Juniperus procero Hochst. ex Endl. It ischaracteristic for undifferentiated Afromon-

tane forest, but it can also be found in rainfor-est. Locally it occurs in nearly pure stands. Insouthern Africa Afrocorpus foieotus is uricom-mon, usually occurring as scattered trees or insmall clumps. It can be found there in moistcoastal forest, as well as in patches of moun-tain forest and wooded ravines. It prefersplaces with annual rainfall ofan

(800-)1200-1800(-2200) mm and mean annualtemperatures of 13-20'C. It is susceptible todrought; it tolerates light frost, but young seed-lings are susceptible. Afrocorpus falcotus ischaracterized as a non-pioneer, shade-tolerantspecies. It performs best on well-drained, deep,humus-rich and light-textured soils with pH of5-7.

Propagation and planting The 1000-seedweight is 0.9-2 kg. The germination capacity ofseeds varies from region to region, with a ger-ruination rate of 40-90%. Seeds collected from

the ground are often infected with the fungusPenicillium cloutforme, which reduces the ger-mination rate. The seed coat delays germina-tion for about one year and removal of thefleshy part promotes germination significantlySeeds germinate best at 25'C. The germinationrate of seeds stored for 12 months at room

temperature dropped to below 35%. Storage atI'C is possible and results in a germinationrate of up to 60% after 2 years, Fair seed vi-ability was also maintained at 4'C for 4 yearsSeeds should be dried to a moisture content

below 15% before storageSeeds can be sown into nursery bags or seedtrays, using a 1:1 mixture of well-rotted coin-post and washed sand. They should be coveredwith a thin layer of soil. At the time of trans-planting, care should be taken not to damagethe taproot.Methods of vegetative propagation have beeninvestigated. Leafy branch cuttings 6-11 cmlong and 2-3 mm in diameter derived fromstock plants of 3 months to 2 years old showedup to 80% rooting when indolebutyric acid wasapplied at doses of 20-80 pg per cutting. Semi-hardwood cuttings 10-12 cm long taken from4-5-year-old saplings treated with 2000 ppmindolebutyric acid showed a rooting rate of45%

Studies in mountain forest in Ethiopia showedthat natural regeneration of Afrocorpus foieo-tus was poor

I

Management Enhanced nutrient supplyincreases the growth rate of seedlings, espe-cially at high irradiance levels, but not onburned sites. Protection of plantations againstfire is necessary. In South Africa trees plantedin agroforestry systems showed better growth(average height 4.2 in and 90% survival after 7years) than in plantations (average height 3.4in and 58% survival)Afrocorpus foieutus is self-pruning, but inplantations wLth wide spacing pruning opera-tions are needed for proper hole developmentIn Rwanda thinning to 50% was carried outwhen the trees were 15 years old, and the re-sponse was very good: at age 26 the mean an-nual growth in diameter was 7 min/year in thethinned plot and only 2.5 min in the unthinnedplot

Diseases and pests Fusorium orysporumand Polyporus sp. are pathogenic to seeds andseedlings. An unidentified fungus causingblackish tumours on twigs and leaves causesvitality problems in relict populations of Afro-corpus foieotus in northern Ethiopia. Seeds arecommonly attacked by insects, which maycause considerable losses.

Harvesting In plantations on good sites,trees can be harvested for timber 40-50 yearsafter planting

Yield Trials in South Africa showed that the

growth rate and yield of Afrocorpus foieotusplantations (standing volume 47.7 mama 11years after planting) and also wood qualitycompared favourably to those of commercialpine plantations. Mean annual volume incre-merit over 20 years was calculated as 5.8matha-

Handling after harvest Logs are susceptl-ble to insect and fungal attacks and should beremoved from the forest and converted as soon

as possible after felling, or treated with pre-servatives

Genetic resources In Ethiopia Afrocorpusfoieotus has become threatened because of se-Iective logging for many decades, whereas thereproductive potential has rapidly declined. Inother countries populations also have dimin-ished seriously, e. g. In Tanzania, whereasmany of the larger Afrocorpus 101cotus treeshave disappeared from South Africa as wellAfrocorpus falcotus is included in the IUCNRed List as vulnerable. It is legally protected inSouth Africa. Ex-situ conservation and prove-nance trials have been carried out in SouthAfrica

In experiments in South Africa it was demon-

strated that seeds from provenances near theafforestation site showed better results after

germination than those from provenances fromfurther away

Prospects Afrocorpus 101cotus could makean excellent indigenous substitute for exoticpine tree plantations throughout its nativerange. Plantations have the capacity to pro-duce valuable timber with good export pros-pects in rotation cycles of reasonable time, andthey have important ecological value as wellStudies to determine the genetic variation,selection of provenances with superior woodcharacteristics and investigations on optimalSIIvicultural systems are needed. Immediateaction is needed for in-situ conservation of theremaining naturalstands

Major references Bekele-Tesemma, 2007;Bolza & Keating, 1972; Friis, 1992; Gelden-huys, 1993b; Napwijk, 2002; Leistner, 1966;Negash, 2003a; Palmer & Pitman, 1972-1974;Takahashi, 1978; World Agroforestry Centre,undated.

Other references Barker, Muller & Mill,2004; Coates Palgrave, 1983; de Laubenfels,1987; Geldenhuys & von dem Bussche, 1996;Dharani, 2002; Gure, Wahlstrom & Stenhd,2005; Hemz, 2004; Kabera, 1992; Kupchan ata1. , 1975; Leistner, Smith & Glen, 1995; unuyaat a1. , 1994; Negash, 2003b; Neuwinger, 2000;Page, 1988; Sharew, Grace & Legg, 1996;Sharew, Legg & Grace, 1997; Stahlhut at al. ,1999; Teketay & Granstr6m, 1997; van

Vuuren, Banks & Stohr, 1978; van Wyk &Gencke, 2000; Wubet at a1. , 2003

Sources of illustration Bekele-Tesemma,2007; Coates Palgrave, 1983; Leistner, 1966

Authors R. Aerts

AUNTSiLODENDRON 39

Aun4TSILODENDRONVILLOSUM (R. Vig. )Viniers

Protologue Bull. Mus. natl. Hist. nat. , sectB, Adansonia 16: 70 (1994)


Synonyms Diehrostochys Dinoso R. Vig.(1949)

Origin and geographic distribution A10nt-silodendron Dittosum is endemic to northern

MadagascarUses The stems are used for posts in house

construction

Botany Small tree; young branches gla-brous, with lenticels. Leaves alternate but clus-

40 TIMBERSl

tered at the apex of short lateral shoots, bipin-nately compound with 13-40 pairs of pinnae,these opposite; stipules densely crowded alongshort shoots, narrowly ovate, 7-12 mm long;petiole 8-13 mm long, rachis 3.5-8 cm long,grooved, woolly hairy and with glands betweenthe pinnae; leaflets 40-45 pairs per pinna, op-POSite, sessile, elliptical to oblong, up to 1.5 mmx 0.5 mm, truncate to rounded and asymmetri-cal at base and obtuse at apex, glabrous. Inno-rescence an axillary head, produced solitary orin clusters on short shoots; peduncle 1.5-2.5cm long, woolly hairy. Flowers bisexual, regu-Iar, 5-merous, sessile, white; calyx saucer-shaped, c. 1.5 mm long, woolly hairy in lowerpart, with short lobes; petals connate into atube 1.5-2 min long, lobes c. I mm long, acute;stamens 10, free, 4.5-6.5 min long, aritherswith a gland at apex; ovary superior, sessile,ellipsoid, woolly hairy, style c. 1.5 min long.Fruit a linear-oblong pod 6-8.5 cm x 0.5-I cm,with thickened and sharply ridged margins,glabrous to shortly hairy, red-brown, dehiscentwith 2 woody valves.A10ntsilodendron comprises 9 species, most ofthem restricted to southern Madagascar. It isclosely related to Diehrostochys and Gogne-bing. A10ntsilodendron oillosum flowers whennew leaves develop.

Ecology Aidntstlodendron Dinosum occursin dry deciduous woodland on sandy soils

Genetic resources and breeding A10ntsi-10dendron unlosum has a limited distribution

and may become easily endangered by habitatdestruction

Prospects Aidntsilodendron Dinosum willremain of minor importance because of itssmallsize and limited distribution

Major references du Puy at a1. , 2002.Other references Viniers, 1994Authors R. H. M. J. Lemmens

rough-bark flat-crown (En). Goane (Po). Mchane,inchani inbao, inchani inhawa, ingendagenda,inchapia turnbili(Sw)

Origin and geographic distribution A1bizioodinnthtfo!to is widespread, occurring fromSenegal east to Kenya, and south to Angola,eastern South Africa and Swaziland; also ineastern Madagascar.

Uses The wood of A1bizio odionthifolio isused for light construction (e. g. posts, rafters)and carving (e. g. images, spoons, masks, clubs).It is also suitable for light flooring, joinery,interior trim, furniture, cabinet work, boatbuilding, vehicle bodies, toys and novelties, toolhandles, baseball bats, boxes, crates, hard-board and particle board. The wood is used asfirewood, although it burns quickly; it is alsomade into charcoal

A1bizio adjonthifo!to is locally valued as ashade tree for crops, e. g. In cocoa and coffeeplantations in Sierra Leone, and is also plantedor retained for soilimprovement and conserva-tion (e. g. in Cameroon). The gum from the barkis sometimes used in local cosmetics. The

young leaves are eaten as a vegetable in DRCongo and Zimbabwe. A sauce is made fromthe seeds. The foliage of A1bizio adjonthifoliois browsed by cattle and wild ruminants, espe-clany the coppice-growthVarious plant parts are used in traditionalmedicine. Bark sap is applied to the eye totreat river blindness and conJunctivitis, andinternally against respiratory complaints, asan anodyne and to treat allergic reactions; it isalso applied to sores and to allay toothache. Abark infusion or decoction is administered to

treat scabies and other skin complaints, and totreat fever. Pounded bark is applied externalIy

ALBIZIAADIANTHIFOLIA (Schumach. )W. Wight

Protologue U. S. Dept. Agr. Bur. PI. Indus-try, Bull. 137: 12 (1909)


Synonyms A1bizio lostigioto (E. Mey. ) Onv(1871), A1bizio intermedio De Wild. & T. Durand(1901), A1bizio eoloensis De Wild. (1907), A1-biziogummifero auct. non (J. F. Ginel. ) C. Asm.


Vernacular names West African albizia,

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A1bizio odionthifolio - wild

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to boils and itching skin, and internally as avermifuge. A twig-bark decoction is adminis-tered as a purgative and anodyne. In tradi-tional South African medicine the bark of A1.bizio adjonthifo!to is used to improve memoryand to treat Alzheimer's disease. A root infu-SIon is apphed to treat eye complaints, andpowdered roots are adminlstered to women inlabour or with irregular menstruation. Theleaves are used internally against diarrhoeaand gonorrhoea, and externaUy to treatwounds and sore feet. A fruit extract is drunkto relieve stomach-ache. In southern Cameroon

the gum from the bark is used in hunting poi-son. In the Central African Republic bark andleaves serve as fish poison.

Production and international trade A1bizio

admitthifolio timber has no importance on theinternational market. However, small volumesmay be mixed and sold with timber of otherA1bizio spp. Parts of the plant are exportedfrom Madagascar as a medicine

Properties The heartwood is pale brown orgolden brown, sometimes with a greenishtinge, and distinctly demarcated from the c. 5cm wide whitish or pale yellow sapwood. Thegrain is straight or interlocked, texture moder-ately coarse to coarse.The wood is moderately light, with a denslty of520-580 kg/ina at 12% moisture content. Itdries slowly, but generally with little degradeThe rates of shrinkage are moderate: fromgreen to 12% moisture content 1.7% radial and4.1% tangential, and from green to oven dry2.3-2.4% radial and 6.5-7.0% tangential. At12% moisture content, the modulus of ruptureis 99-136 N/min2, modulus of elasticity 9300N/min2, compression parallelto grain 52.5-57.5N/mm2 cleavage 11.5-27.4 N/min and Chalais-Meudon side hardness 1.9-2.8

The wood generally saws and works easily withordinary hand and machine tools. The use of afiller is necessary to obtain a good finish. Thewood nails satisfactorily, and gluing and stain-ing properties are good. The wood dust maycause irritation to nose and throat. The heart-wood is only moderately durable, being suscep-tible to wood borer, marine borer and termiteattacks. The heartwood is resistant to Impreg-nation by preservatives, the sapwood moder-ately resistantto permeableA crude protein content of the foliage of 29 gper 100 g dry matter has been recorded. Theleaves of trees growing on acid soils in south-eastern Nigeria contained per 100 g dry mat-ter: N 3.72 g, P 0.13 g, K 088 g, Ca 0.48 g and

Mg 0.38 g. 01eanane-type triterpene saponlnswere isolated from ethanolic extracts of A1btzioodinnthifolio roots. Some of these compoundsshowed jinmunomodulatory and haemolyticactivities, particularly adjanthifoliosides A andB. They also demonstrated a cytotoxic effect onhuman Ieukaemia T-ceMs. The bark contains

large amounts (2 ing/g dry matter) of histamineand related jinidazole compounds.

Adulterations and substitutes The woodof A1bizio gummifero (J. F. Ginel. ) C. Asm. andA1bizio zygio (DC. ) J. F. Machr. is very similarto that of A1bizio odiunthifolio and used for thesame purposes

Description Smallto medium-sized decidu-ous tree up to BOG35) in tall; hole straight andcylindrical in closed forest but often crookedand/or twisted in more open localities, up to 95cm in diameter, without buttresses or withsmall, thick buttresses; bark yellowish brownto grey, smooth or rough, Inner bark granular,creamy to yellowish, wlth clear gum; crownflattened, with large, spreading branches;young branches densely yellowish or reddishpubescent. Leaves alternate, hipinnately coin-

ALEiziA 41

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A1bizio odionthifoli0 - I, 60se of 601e, . 2, ledj. 3,inflorescence, . 4, fruit.Redrown ond adopted by Achingd Satin Nur-human

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42 TIMBERSl

pound with (3-)5-10 pairs of pinnae; stipulesovate to lanceolate, up to 12 mm long, cadu-cous; petiole 1.5-7.5 cm long, near the base atupper side with a sessile gland, rachis 3-12 cmlong, yellowish or reddish pubescent; leaflets in5-17 pairs per pinna, sessile, obliquely rhombicto elliptical-obovate, up to 1.5(-2) cm x I cm,curieate to rounded at base, obtuse to acute atapex, pubescent below. Inflorescence an axil-Iary head on a 2-6 cm long peduncle. Flowersbisexual, regular, 5-merous, reddish or green-ish white, almost sessile, subtended by up to 8min long, caducous or persistent bracteoles;calyx narrowly obconica1, 2.5-5 min long, pu-bescent outside; corolla 6-12.5 min long, with5-9 min long tube, pubescent outside; stamensnumerous, 3-3.5 cm long, united into a tube formost of their length, red to pink or greenish;ovary superior, narrowly ellipsoid, 2-3 mmlong, gradually tapering into a 3-3.5 cm longstyle. Fruit an oblong, flat pod 9-19 cm x 2-3.5cm, with stipe c. 0.5 cm long, densely but finelypubescent, transversely veined, pale brownwhen ripe, opening with 2 papery valves, 7-10-seeded. Seeds flattened globose, 6.5-9.5 min x6.5-8.5 mm. Seedling with epigeal germina-tion; hypocoty1 2-6 cm long, epicotyl c. I cmlong; cotyledons oblong, c. I cm long, thick andfleshy, rounded, early caducous; first 2 leavesopposite, hipinnately compound with I pair ofpinnae

Other botanical information A1bizio coin-

prises about 120 species and occurs throughoutthe tropics. Approximately 35 species are foundin continental Africa and about 30 in Madagas-car. It is characterized by the head-like milo-rescence, with I-2 central flowers modified,functionally male and having a larger, nectar-producing staminaltube. Molecular analysesshowed that A1bizio is heterogeneous, and arevision of the genus is needed. A1bizio adjon-thifolio is frequently confused with A1bi2togummifero, which differs in its almost glabrousleaflets usually auricled at base, and glabrouspods. However, almost glabrous types of A1-bizio odionthifolio have also been recorded,and more research is needed to confirm theseparation of the two species, the more so sincehybrids have been recorded from Malawi andMozambique.

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pits

alternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 prn); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;43: mean tangential diameter of vesselluminaZ 200 pin; (46: s 5 vessels per square minime-tre); 47: 5-20 vessels per square minimetre; 58:gums and other deposits in heartwood vesselsTracheids and fibres: 61: fibres with simple tominutely bordered pits; 65: septate fibres pre-sent; 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: 79:axial parenchyma vasicentric; 80: axial paren-chyma allform; 81: axial parenchyma lozenge-allform; 83: axial parenchyma confluent; 91:two cells per parenchyma strand; 92: four (3-4)cells per parenchyma strand. Rays: (97: raywidth I-3 cells); (98: larger rays commonly 4-to 10-senate); 104: all ray cells procumbent;1/5: 4-12 rays per min. Mineral inclusions136: prismatic crystals present; 142: prismaticcrystals in chambered axial parenchyma cells;(143: prismatic crystals in fibres).(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development Trees can growrapidly, with a recorded maximum early heightgrowth of 2 in per year. However, growth ofplanted trees is often much less and even slowduring the first years' In planting experimentsin Ghana 3-years-old trees reached a height ofon average 1.5 in with a stem diameter of 1.5cm. The roots develop nitrogen-fixing nodulescontaining Bradyrhizobium bacteria. A1bizioodionthifolio trees live in association with ar-buscular mycorrhizae. They usually flower atthe end of the dry season, just after or with theemergence of new leaves. Insects such as beesare the main pollinators. The fruits dehisce onthe tree and the papery valves with seeds stillattached are spread by wind.

Ecology A1bizio udionthtfolio has a remarka-bly wide ecological adaptation, occurring inhigh-rainfall forests as well as seasonalIy dryforests and even in wooded savanna. It occursmost frequently in moister types of semidecid-uous forest, where it is characteristic for sec-ondary forest, forest edges, roadsides andabandoned farmland. It is common in manyregions. In East and southern Africa and inMadagascar, it is found in lowland rainforest,deciduous woodland and wooded grassland, upto 2000 in altitude. It behaves mostly as a PIO-neer species. A1bizto odionthifolio occurs on awide range of soil types, often on deep and

.

.I

sandy soilsPropagation and planting Seedlings are

classified as strong light demanders. The ger-mination rate of seeds in the light and darkmay be equal, but seedlings soon die in theshade. Regeneration is most abundantin forestdisturbed by logging. In burnt forest, seedlingsare less abundant. There are about 25,000seeds per kg. Seeds should be collected frompods stillattached to the tree to reduce damageby insects, and they should be dried jinmedi-ately after collection. They can be stored for upto 3 months if ash is added to reduce insectdamage. In cultivation seed may be used forpropagation, but wildlings are sometimes alsoused for planting. Experiments in Ghanashowed that A1bizio odiorrthifolio can be sric-cessfully propagated vegetativeIy by root cut-tings.

Management In forest management thereis generally no focus on A1bizio adjonthifoliobecause it is not a preferred timber tree, andsometimes even considered an aggressive colo-nizer. It is occasionally planted as an auxiliarytree in agroforestry systems. In planting ex-perlments in Ghana A1bizio odionthifolioshowed a survival rate of 89% three years afterplanting

Diseases and pests Seeds suffer a highincidence of insect attack, probably often bybruchid beetles as in other A1bizio spp.

Yield The yield of timber per tree is oftenlow for A1bizio odionthifoliu because the bole isoften low branching and crooked.

Handling after harvest Freshly harvestedlogs float in water and can be transported byriver. Treatment of the logs with preservativesis necessary when they are left in the forest forsome time. Logs are sometimes hollow

Genetic resources A1bizio odiunthtfolio iswidespread and locally common in secondaryforest. It is therefore not easily liable to genetic

with its leaf litter, reducing erosion with itslarge rooting system, and protecting crops fromtoo much sun. However, the results of screen-ing trials were quite variable, from disappoint-ing due to slow initial growth in Tanzania andZambia, to very good locally in Ghana. Theexistence of different ecotypes has been sug-gested, and this deserves more research atten-tion because it may offer possibilities for opti-inizing the use of A1bizio odionthifolio in agro-forestry systems and afforestation programmesunder different ecological conditions. With itsnat, wide-spreading crown, It has value as anornamental shade tree. A1bizio odiorrthifolio isan important and widely used medicinal plant,and more research is desirable on its activecompounds, some of which have already showninteresting pharmacological activities.

Major references Arbonnier, 2000; Bolza &Keating, 1972; BurkiU, 1995; du Puy at al. ,2002; Katende, Birnie & Tengnas, 1995; Keay,1989; Neuwinger, 1998; Neuwinger, 2000; Ta-kahashi, 1978; World Agroforestry Centre,undated

Other references Aubr6ville, 1959c;Beentje, 1994; Brenan, 1959; Brenan, 1970;Clarke, 2000; Coates Palgrave, 1983; Cobbinaat a1. , 1990; Danquah, 2000; Debray, Jacque-min & Razafindramba0, 1971; GIIbert & Bou-tique, 1952; Haddad, Laurens & Lacaille-Dubois, 2004; Haddad at a1. , 2003; Hawthorne,1990; Hawthorne, 1995; Honu, 1993; Inside-Wood, undated; Latham, 2004; Palmer & Pit-man, 1972-1974; Saville & Fox, 1967; Wilks &Issemb6, 2000; Zambia Forest Department,1979b.

Sources of illustration Hawthorne, 1990;White, 1962; Wilks & Issemb6, 2000


A1, BIZiA 43

erosion and protection measures are notneeded

Prospects A1bizio admitthifolio is a multi-purpose tree. In general it Is not a preferredtimber tree because of its often short andcrooked bole, but where it occurs in closed for-est its hole may be longer and more straight,and may be harvested indiscriminately fromother A1bizio spp. which are more common inclosed forest (e. g. A1bizio Ierrugineo (Gulll. &Perr. ) Benth. ). A1biziu odionthifolio is oftenrecommended as an auxiliary tree in agrofor-estry systems, improving the soil with its ni-trogen-fixing root nodules, providing mulch

ALBIZiAALTISSiMA Hook. f.

Protologue Hook. , Niger F1. : 332 (1849)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Chromosome number2n= 26

Pithecellobium altissimumSynonyms(Hook. f. ) 011v. (1871), Cothormion o1tissimum(Hook. f. ) Hutch. & Dandy (1928), Ar-throsomone0 o1tissimo (Hook. f. ) G. CG. GIIbert& Boutique (1952).

Origin and geographic distribution A1bizioo1tissimo occurs from Sierra Leone east to

Uganda, and south to Zambia andAngolaUses The wood is used locally for furnlture

.

44 TIMBERSl

and implements, e. g. tool handles. The bark isused in traditional medicine in Sierra Leoneand DR Congo; a decoction is used as an ano-dyne to treat toothache and stomach-ache, andagainst pulmonary affections, and externalIy totreat sores. The bark also serves as fish poison.Scraped inner bark beaten up in water is usedas soap to wash clothes. In DR Congo a leafdecoction is used in a vapour bath to treatcolds. Burned leaves are applied to snakebitesThe fruits are used for tanning and dyeing, andto prepare Ink and soap. The fruit pulp and theseeds are edible. In Nigeria fermented seedscalled 'OSo' are used as condiment in soups

Production and international trade A1bizioo1tissimo is traded with other A1bizio spp. as'kibizia'.

Properties The heartwood is pale brown toyellowish brown, often with darker stripes, anddistinctly demarcated from the whitish sap-wood. The grain is often wavy or interlocked,texture moderately fine. The wood is moder-ately heavy and hard. The shrinkage rates are4.6% radial and 7.4% tangential from green tooven dry. At 12% moisture content, themodulus of rupture is about 128 N/min2modulus of elasticity 8920 N/mm2 compressionparallel to grain 60 N/mm2 and shear 10N/min2. Reports on workability vary from easyto difficult; when finished the wood has an at-tractive polish. The wood is durableSaponins are present in different parts of theplant, especially the bark, which explains theuse as fish poison and soap and possibly alsosome medicinal applications. The bark showedantifungal activity against pathogens affectinghumans and plants. Tinidazole, a compoundwith antifungal and antibacterial activities,has been isolated from the seeds. analysis offermented seeds showed 25.3% protein, 16.9%lipid and 10.0% carbohydrate. Several bacteriaare responsible for the fermentation process

Botany Small to fairly large tree up to 35 intall; hole often short and bent, up to 80 cm indiameter; bark scaly, dull grey to brown; crownopen, spreading, flat-topped, often with pen-dent branches; young branches shortly hairyLeaves alternate, bipinnately compound, with4-8 pairs of pinnae; petiole and rachis shortlyhairy, with glands on the upper side betweenthe pinnae; leaflets opposite in 10-25 pairs perpinna but lowest leaflet solitary, slightlyobliquely oblong, 7-17 mm x 2.5-6 min, gla-brous. Inflorescence an axillary globose head,I-3 together; peduncle I-4.5 cm long. Flowersbisexual, regular, usually 5-merous, white,

sessile; one to several central flowers in eachflower head larger; calyx cup-shaped, 3-3.5min long, with short teeth; corolla funnel-shaped, 5-8.5 min long, glabrous; stamens nu-merous, united at base, c. 12 mm long; ovarysuperior, I-celled, style finform, about as longas stamens. Fruit a narrowly oblong pod 10-28cm x I-2 cm, curved or spiralIy twisted, coin-pressed, red-brown to blackish, constrictedbetween the seeds, up to 20-seeded, breakingup into I-seeded segments. Seeds oblong tolens-shaped, flattened, 6-9 min x 6.5-7 min,smooth, brown. Seedling with epigeal germina-tion.

A1bi2io comprises about 120 species and occursin all tropical regions. In continental Africaabout 35 species occur, in Madagascar about 30species. Some African species, including A1-bizi0 o1tissimo, with pods breaking up into I-seeded segments have been included in Co-Ihormio, ,. However, all transitions from inde-hiscent segmented pods to dehiscent podswhich are indistinctly segmented can be found,and thus these species have been transferred toA1bizio

Another species with pods breaking up into I-seeded segments is A1biaio rhombifolio Benth(synonym: Cothormion rhombifolium (Benth. )Keay), which occurs from Senegal to SierraLeone and differs from A1bizi0 o1tissimo in its

fewer and larger leaflets, stalked flowers andshorter pods. The wood of A1bizio rhombifolio,which is a smalltree up to 10 in tall, is used inSierra Leone to make bed-postsA1bizi0 o1tissimo nodulates with Brodyrhizo-binm bacteria. In experiments in C6te d'TVoire,inoculation had a positive effect on plantheight and stem diameter measured at 4 and11 months after inoculation. In C6te d'Ivoire

seeds usually germinate 6-30 days after sow-ing, and fruiting occurs from March to May.

Ecology A1bizi0 o1tissimo characteristicallyoccurs in riverme forest and freshwater swampforest, but can also be found in secondary for-est

Management The 1000-seed weightis about100 g.

Genetic resources and breeding A1bizioo1tissLmo is widespread and there are no indi-cations that it is threatened by genetic erosion.

Prospects A1bizi0 o1tissimo will probablyremain a minor timber tree because of its spe-cific ecological requirements and short, oftenbent bole. If industrial production of 'OSo' be-comes feasible, possibilities for domesticationof the species deserve investigation

Major references Brenan, 1970; Burkill,1995; Popoola, Jolaoso & of o1abi, 2004;Raponda-Walker & Sillans, 1961; Viniers,1989

Other references Aubr6ville, 1959c;Brenan, 1959; de in Mensbruge, 1966; Diabateat a1. , 2005; Eggeling & Dale, 1951; Gilbert &Boutique, 1952; Hayman & Gray, 1987;Kuster-Laine, 1985; Neuwinger, 2000; Takaha-shi, 1978


A1, BIZLAANTUNESIANA Harms

Protologue Bot. Jahrb. Syst. 30: 75 (1901).Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names Purple-leaved albizia,

purple-leaved false thorn (En). Muiando (Po)Origin and geographic distribution A1bizio

antu, lesiono Is widespread, from eastern DRCongo, Rwanda, Burundi and Tanzania southto Namibia, Botswana, Zimbabwe, Mozam-bique and South Africa

Uses The wood of A1bizio gritunesiono usused for heavy construction, loinery, furniture,boat building, plywood, drums and implementsin Zambia and Zimbabwe. It is one of thewoods preferred by the woodcarving industryin southern Zimbabwe. The wood is suitable forcabinet work, interior trim and railway sleep-ers. It also serves as firewood. The roots havenumerous uses in traditional medicine. Theyare applied externalIy to treat sore eyes, cuts,ulcers, pneumonia, painful and swollen legs,and internally as an Infusion or decoction totreat sore throat, tonsillitis, tuberculosis, gon-orrhoea and other sexuaUy transmitted dis-eases, abdominal pains, depressed fontanelle ininfants and infertility in women, as an aphro-disiac, and to prevent abortion. A bark infusionis taken to treat constipation and applied ex-ternally to cuts, whereas crushed leaves areused as an enema for their purgative actionand as a dressing to treat oedema of the legsThe bark has been used for tanning, and theflowers are a source of nectar for honey bees.

Properties The heartwood varies from palebrown to purplish brown, sometimes darkerstriped, and not distinctly demarcated from theup to 6.5 cm wide, whitish sapwood. The grainis irregularly interlocked, texture moderately

The wood is moderately heavy, with a densityof 640-785 kg/ms at 12% moisture content. It

dries slowly, but with little degrade. Shrinkagerates from green to 12% moisture content are1.3% radial and 2.0% tangential. Once dry, it isstable in service. The wood saws and worksweU, but blunts tools rather rapidly. Surfacesmay tear because of interlocked grain; a cut-ting angle of 10' is recommended for planingand low speeds are needed in moulding. Thewood may split upon nailing. It does not turnwell, but mortises, peels and slices satisfacto-rily. The sawdust causes irritation to nose andthroat. The wood is considered durable andresistant to termites. It is highly resistant toimpregnation by preservatives.Roots of A1bizio ontunesiono showed signifi-cant in-vitro activities against the tapewormHymeno!epis diminuto ('rattapeworm') and thetrematode Schistosomo monsoni, causal agentof schistosomiasis

Botany Small to medium-sized deciduoustree up to 18(-25) in tall; bole usually shortand branchless for up to 5 in butsometimes upto 12 in, straight or twisted, up to 75 cm Indiameter; bark surface grey to brown, smoothor rough and Teticulate, inner bark with red-

flattened, umbrella-dish exudate; crown

shaped; young twlgs glabrous or very shortlyhairy. Leaves alternate, hipinnately compoundwith I-3(-4) pairs of pinnae; stipules awl-shaped, caducous; petiole 4-8 cm long, groovedabove, glabrous, near the base at upper sidewith a sessile gland, rachis 8--14 cm long, gla-brous or sparsely pubescent; leaflets in (3-)4-8(-9) pairs per pinna, almost sessile, obliquelyThornbic-ovate to elliptical-oblong, up to 5(-7)cm x 2.5(-4) cm, rounded to slightly notched atapex, glabrous, distinctly glaucous below. In-florescence an axillary head on 2-11 cm longpeduncle. Flowers bisexual, regular, 5-meTous,greenish yellow; pedicelup to 2 mm long; calyx3-5.5 min long, with long tube toothed at apex,pubescent outside; corolla 5-11 min long, pu-bescent outside; stamens numerous, 1.5-3 cmlong, united at base, with white filaments;ovary superior, gradually tapering into an upto 3 cm long style. Fruit an oblong, flat pod 12-23 cm x 2.5-5 cm, almost glabrous, indistinctlytransversely veined, pale brown when ripe,opening with 2 papery valves, c. 8-seeded.Seeds lens-shaped, 7-9 min in diameterA1bizio ontunestuna trees usually flower inAugust-November and frults are rlpe in April-September. The pod valves with seeds stillattached are spread by windA1bizio comprises about 120 species and occursthroughout the tropics. Approximately 35 spe-

A1. BIZIA 45

coarse

I

46 TIMBERSl

cies are found in continental Africa and about30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is needed

Ecology A1bizio gritunesiono occurs in de-ciduous woodland and wooded grassland at(250-)900-1700 in altitude.

Management Propagation can be done byseed. Seeds should be collected before the podsdehisce, which makes collection difficult. Wild.lings are sometimes collected for plantingPlanted trees can be managed by lopping andpollarding

Genetic resources and breeding Thereare no indications that A1bizio Gritunesiono isthreatened by genetic erosion. However, thecommon practice of harvesting the roots fortraditional medicine may severely reduce popu-Iations of the species locally

Prospects Although the wood of A1biziogritunesionu is of excellent quality, it is of littleeconomic Importance because of the often smallsize and poor shape of the hole. However, treesof good stature do exist and planting experi-merits are desirable to evaluate possibilities asa timber-plantation tree. It is surprising thatsuch a well-known medicinal tree has hardlybeen tested on its pharmacological activities

Major references Bolza & Keating, 1972;Brenan, 1959; Bryce, 1967; Chilufya &Tengnas, 1996; Gelfand at a1. , 1985.

Other references Braedt at a1. , 2000;Brenan, 1970; Coates Palerave, 1957; GIIbert& Boutique, 1952; Molgaard at a1. , 2001; Ndu-bani & H6jer, 1999; Neuwinger, 2000; Palmer& Pitman, 1972-1974; van Wyk & van Wyk,1997


inner bark fibrous, with sticky gum; youngbranches sparsely pubescent, glabrescentLeaves alternate, hipinnately compound with4-9 pairs of pinnae; stipules minute, caducous;petiole 1.5-2.5 cm long, in the basal half ofupper side with a sessile gland, rachis 3.5-5.5cm long, pubescent; leaflets in (2-)4-17 pairsper pinna, sessile, obliquely oblong to obovate-oblong, up to 8 mm x 4 mm, rounded at apex,pubescent on both sides. Inflorescence an axil-Iary head on 2-4 cm long peduncle. Flowersbisexual, regular, 5-merous, greenish white,almost sessile; calyx narrowly obconica1, 1.5-2mm long, glabrous; corolla 5-6 mm long, gla-brous; stamens numerous, (1.5-)2-2.5 cm long,united into a tube at base, white; ovary supe-nor, shortly stipitate, glabrous, gradually ta-permg into a 2-2.5 cm long style. Fruit an ob-long to broadly oblong pod 7.5-16 cm x 2.5-6cm, with short stipe, slightly swollen, withthick and woody fruit wall, glabrous, withoutveins, several-seeded. Seeds ellipsoid, 10-16min x 4.5-8 min, black.A1bizio orenico!o flowers from August to No-veinber. The flowers are pollinated by insectssuch as butterflies

A1bizio comprises about 120 species and occursthroughout the tropics. Approximately 35 spe-cies are found in continental Africa and about

30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is neededA1bizi0 60inensis R. Vig. , a shrub or small treeup to 15 in tall with a hole diameter up to 50cm, has thick, woody and indehiscent pods likeA1bizio orenicolo, but it has more numerousand smaller leaflets (up to 5 min x 1.5 min)The wood of A1bizio botnensts is used in north-western Madagascar for furniture. A1bizio ino-sthororum R. Vig. closely resembles A1bizioorenico!u, but differs in its smaller leaflets (upto 3 mm x 1.5 min) produced on short shoots.The wood of A1bizio mosthororum is used inhouse building and as fuelwood in south-western Madagascar.

EcologyAlbizio orenicolo occurs widespreadbut scattered in dry deciduous woodland andscrubland up to 700 in altitude, on sandy soilsand sandstone

Genetic resources and breeding Thereare no indications that A1bizio orenicolo isthreatened by genetic erosion

Prospects It is unlikely that the utilization

ALBIZIAARENICOLA R. Vig.

Protologue Notul. Syst. (Paris) 13(4): 335(1948).


Origin and geographic distribution A1bizioorenicolo is endemic to Madagascar, where it iswidespread in the western part of the island.

Uses The durable wood is used for poles inhouse construction.

Botany Small to medium-sized deciduoustree up to 15 in tall; bark grey, usually smooth,

of A1bizio orenico!o timber will increase beyondits currentlocalimportance

Major references Capuron, 1970; du Puy ata1. , 2002Other references Rabarimanariv0, 2000;Stiles, 1998.


ALBiziAAYLMERIHtitch. ex Broun & Massey

Protologue F1. Sudan: 175 (1929)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Origin and geographic distribution A1btzio

oylmeriis endemic to central Sudan.Uses The wood of A1bizio oylmeriis used as

timber and is suitable for construction, lightflooring, interior trim, toys and novelties,turnery, veneer, plywood, hardboard and parti-cle board.

Properties The heartwood is reddish brown,often darker striped, and distinctly demarcatedfrom the up to 5 cm wide, creamy to yellowlshsapwood. The grain is often spiral, texture fineto coarse and even

The wood is moderately heavy, with a densityof about 740 kg/ing at 12% moisture content. Itshould be dried slowly and carefully to avoidserious degrade. The rates of shrinkage aremoderate: from green to oven dry 3.8% radialand 5.9% tangential. Once dry, the wood isstable in servlce

At 12% moisture content, the modulus of rup-ture is 1/3 N/mm2 modulus of elasticity 11,600N/mm2, compression parallel to grain 68.5N/mm2 shear 13.7 N/mm2 radial and 11.8

N/min2 tangential, cleavage 7.3 N/min radialand 7.4 N/mm radial, Brinell side hardness 28N/mm2 radial and 30 N/mm2 tangential, andBrinellend hardness 57 N/mm2The wood saws and works well, but causesrapid blunting of saw teeth and tool edges. Itfinishes to a smooth or rough surface depend-ing on the wood texture. The nailing, screwLng,gluing and staining properties are allgood. Thewood dust may cause irritation to nose andthroat. The wood is moderately durable, beingresistant to termite attack, but susceptible tomarine borers. It is resistant to Impregnationby preservatives.

Botany Medium-sized tree up to 25 in tall;hole up to 80 cm in diameter, often slightlytwisted; bark surface brown, fissured. Leavesalternate, hipinnately compound; leaflets in 5-8 pairs per pinna, obliquely broadly oblong to

rhombic, 2.5-5 cm x I-3 cm, rounded to obtuseat apex, whitish and pubescent below. Inflores-cence an axillary head on c. 5 cm long pedun-cle. Flowers bisexual, regular, 5-merous, sessileor with short, thick pedicel, greyish pubescent;stamens numerous, far exserted, united at baseinto a tube about as long as corolla; ovary su-perlor, gradually tapering into a long, slenderstyle. Fruit an oblong, flat pod 17-20 cm x c. 3cm, glabrous.A1bizio comprises about 120 species and occursthroughout the tropics. Approximately 35 spe-cies are found in continental Africa and about30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow.ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is needed.

Ecology A1bizio oy!inert occurs in woodedsavanna

Management Logs should be processedquickly after felling because the wood is sus-ceptible to discoloration by blue-stain fungi

Genetic resources and breeding A1biziouy!inert has a small area of distribution andmay be threatened by genetic erosion, but istoo poorly known to establish its conservatlonstatus.

Prospects Apart from sporadic tests of thewood, very little is known about A1bizio oyl-inert. There is no recentinformation on lts usesand conservation needs.

Major references Bolza & Keating, 1972;Gottwald & Noack, 1966; Takahashi, 1978.

Other references hadrews, 1952; Bege-inarin, 1963-1969; FAO, 2001; GOTashi, 2001.


A1. BIZiA 47

ALEizlABERNIERiE. Fourn. exVilliers

Protologue Legum. Madagascar: 254 (2002)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Origin and geographic distribution A1bizio

bemien is endemic to Madagascar, where it iswidespread in the western part of the island

Uses The wood is used in house constructionand for planks. The bark is used for makingrope, and it is also suitable for tanning hides

Properties Seed extracts are toxic to mice,and showed haemolytic activity in red bloodceUs of sheep as well as antibacterial activity.

Botany Shrub or small to medium-sized,deciduous tree up to 25 in tall; hole up to 100

48 TIMBERSl

cm in diameter; bark whitish to pale grey,finely fissured; crown umbrella-shaped; youngbranches sparsely pubescent, glabrescent.Leaves alternate, bipinnately compound with4-15 pairs of pinnae; stipules minute, cadu-cous; petiole I-2 cm long, in the apical half ofupper side with a sessile gland, rachis 1.5-9 cmlong, pubescent; pinnae ending in a largeinucronate gland; leaflets in 10-35 pairs perpinna, sessile, slightly obliquely oblong, up to 5mm x I min, obtuse to acute at apex, glabrousexcept for hairs at margins. Inflorescence anaxillary head on 2-4.4 cm long peduncle. Flow-

bisexual, regular, 5-merous, greenishwhite, almost sessile; calyx cylindrical to ob-conical, 1.5-2 min long, pubescent; corolla 5-7.5 mm long, with tube 3-5.5 mm long, pubes-cent; stamens numerous, 2-2.5 cm long, unitedinto a tube at base, white in lower part, pink-purplish in upper part; ovary superior, nar-rowly ellipsoid, with 0.5-I min long stipe, gla-brous, gradually tapering into a c. 2.5 cm longstyle. Fruit an oblong, flat pod 8-21 cm x 3-5cm, with short stipe, slightly pubescent to al-most glabrous, transversely veined, brown,several-seeded. Seeds oblong-ellipsoid, 10-12min x 5.5-6 mm, blackA1bizio bemien usually flowers from October toDecember


30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is needed.A1bizio polyphyl!o E. Fourn. has small glandsat the tips of pinnae like A1bizio bemien, butmore pinnae per leaf and more and smallerleaflets (up to 3.5 min x I mm). In western andsouthern Madagascar its wood is used for con-struction and dugout canoes, as well as forfirewood, and the bark is used to make rope

Ecology A1bizio bemien occurs in deciduouswoodland and scrubland up to 100 in altitude,on all soil types. It is mainly found in moistlocations, such as clayey depressions and riverbanks.

Genetic resources and breeding A1biziohemieriis common over a large area and notthreatened by genetic erosion.

Prospects A1bizio bemien timber witlproba-bly remain of local importance for constructionpurposes. It is unlikely that it will become jin-

ers

portant on a commercialscale.Major references du Puy at a1. , 2002; Ra-

bansoa, 1999.Other references Boiteau, Bolteau & A1-

10rge-Bolteau, 1999; Capuron, 1970; Rabari-manariv0, 2000


ALBIZIABOmNllE. Fourn.

Protologue kiln. Sal. Nat. , Bot. , s6r. 4, 14:378 (1860).


Origin and geographic distribution A1biziobotuiniL Is endemic to Madagascar, where it iswidespread in the western and central parts ofthe island.

Uses The wood is used for dugout canoes. ItIs suitable for peeled veneer and plywoodcorestock. It is also used as firewood and forcharcoal production.

Production and international trade Thewood of A1bizio hojuiniihas been exported fromnorth-western Madagascar as rotary-cut ve-neer. It is traded in small quantities only andmixed with wood of other A1bizio spp

Properties The wood is whitish to slightlypinkish. The density is about 500 kg/ing at 12%moisture content. The wood dries rapidly. Theshrinkage rates are low, from green to oven dry2.7% radial and 5.2% tangential. The wood issoft and easy to work. The durability of thewood is low, it is liable to fungal attack andsusceptible to termltes and Lyetus borers.Seed extracts are toxic to mice, and showedhaemolytic activity in red blood cells of sheepas well as antibacterial actrvity against severalpathogenic bacteria. Pod extracts showed toxic-ity against crab brood and inhibit germinationof rice and bean seeds

Botany Small to medium-sized deciduoustree up to 20(-35) in tall; hole up to 40(-130)cm in diameter; bark pale grey to purplish,cracking into thin plates; young branches pu-bescent, glabrescent. Leaves alternate, hipin-nately compound with (I-)2-3(-4) pairs of pin-nae; stipules minute, caducous; petiole 1.5-7.5cm long, in the basal half of upper side with asessile gland, rachis (1.2-11.5 cm long, gla-brous to pubescent; leaflets in (2-)3-8(-9) pairsper pinna, with petiolules 1.5-3 mm long,slightly obliquely elliptical to elliptical-obovate,up to 4 cm x 2.5 cm, rounded to obtuse at apex,leathery, slightly pubescent to glabrous. Inflo-

'I

rescence an axillary head on 3-8 cm long pe-duncle. Flowers bisexual, regular, 5-merous,greenish white, sessile; calyx narrowly obconi-cal, c. 2 min long, pubescent; corolla 4-5 minlong, with 2.5-3 mm long tube, pubescent;stamens numerous, 9-12.5 min long, unltedinto a tube at base, white; ovary superior, errip-sold, shortly stipitate, glabrous, gradually ta-permg into a c. 11 min long style. Fruit an ob-long to oblong-elliptical flat pod 10-29 cm x 2-5 cm, with short stipe, thick-papery, glabrous,with fine transverse veins, several-seeded.Seeds narrowly oblong to narrowly obovoid,12-13 min x 5-6 mm, black.A1bizio botuintiflowers from July to JanuaryA1bizio comprises about 120 species and occursthroughout the tropics. In continental Africaapproximately 35 species are found and about30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is neededA1bizio gunsporso (Drake) R. Vig. is a smalltree up to 15 in tall, occurring mainly on sandysoils in western Madagascar. It resembles A1-bizio botuinii, but it is distinguished by thegolden-yellow pubescence on young leaves,flowers and pods. The durable wood of A1btziogunsporso is used, e. g. for coffins. A1bizio gre-

(Baill. ) R. Baron resembles A1biziobotuinii in having few pinnae with few, largeleaflets per leaf, but differs in its distinctlypedicellate flowers. It occurs in western Mada-gascar on a wide range of soiltypes. Its wood isused for dugout canoes. A1bizio odoroto R. Vig.is a medium-sized tree up to 20(-30) in tall,occurring in western Madagascar, scattered oncalcareous soils; its leaves resemble those ofA1bizio greueo, IQ, but can be distinguished bybeing completely glabrous and havLng dis-tinctly acuminate leaflets. Its wood is occasion-ally used for furniture. A1bizto tu!earensisR. Vig. is another species with few and coin-paratively large leaflets, but is characterlzedby the greyish white pubescence on youngshoots, leaves and flowers. It is a common treein southern Madagascar, up to 15(-20) in tall,with a hole diameter up to 70 cm. Its good-quality wood is used for construction, beams,planks and coffins

Ecology A1bizio botuinii occurs in deciduouswoodland up to 1800 in altitude, mostly onsandy soils.

Genetic resources and breeding A1bizio

botuiniiis common over a large area and notthreatened by genetic erosion.

Prospects Little is known about the woodproperties of A1bizio botuinii, but the wood isreportedIy suitable for veneer production. Thismay offer possibilities for commercial exploita-tion as a timber tree, but research is needed onecology, growth rates and regeneration toevaluate methods of sustainable production.

Major references Capuron, 1970; du Puy ata1. , 2002; Raherinirina, 1999.

Other references hadrianavalona, 2001;Gu6neau, Bedel & Thie1, 1970-1975; Parant,Chichignoud & Rakotova0, 1985; Rabarimana-riv0, 2000


A1. BIZIA 49

ueono

ALBIZiAFERRUGiNEA (Guill. & Perr. ) Benth.

Protologue London Journ. Bot. 3: 88 (1844)Family Mimosaceae (Leguminosae - Mimo-


Synonyms A1bizio origo!errsis WeIw. ex 011v.(1871).

Vernacular names West African albizia

(En). Mufufuta, kambala escura (Po)Origin and geographic distribution A1bizio

Ierrugineo is widespread, occurring from Sene-gal east to Uganda and south to northern All-gola.

Uses The wood of A1bizioferrugineo is knownunder the trade names tatandza (Gabon),awlemfosamina (Ghana) and musase (Liberia).It is used for construction, light and heavyflooring, staircases, furniture, cabinet making,joinery, turnery, implements (e. g. mortars),

I.

A1bizio Ierrt, gineo - wild

D

..

o

,^

..

I

50 TIMBERSl

carvings, glued laminated timber and veneerIt is also suitable for carpentry, mine props,vehicle bodies, toys and novelties, musical in-struments, railway sleepers, boxes and crates.In south-eastern DR Congo the wood is coin-monly used for charcoal production.A1bizio forrugineo is planted as an ornamentalshade tree and roadside tree. The leaf litter

improves the soil, and the tree is being testedfor shade and mulch in agroforestry. The foli-age is eaten by goats. The leaves are some-times used for washing clothes. The flowersproduce nectar collected by bees. Branchletshave been used as firesticks

The bark is used in traditional medicine; adecoction is administered to treat dysentery,bronchial affections and pain caused by fever,and it is applied externalIy to sores, pimplesand other skin complaints. It is also used as avermifuge. A bark maceration is drunk andapplied as a wash to treatjaundice. Leaf decoc-tions are used externalIy to treat headache,and as a wash or steam inhalation against fe-ver (including malaria) and toothache; they arealso applied as a wash to kill head lice. A leafmaceration is administered as an enema to

induce abortion. The bark is also used in vet-

erinary medicine. In Gamero0n the root barkand the gum from the stem bark are used inthe preparation of arrow poison. The leavesserve as fish poison in the Central African Re-public.

Production and international trade A1bizio

Ierrugineo timber has some importance on theinternational market, although it is considereda lesser-used timber. It is often mixed with

timber of other A1bizio spp. In Ghana the ex-port volume of sawn A1bizio timber in the firsthalf of 2004 was 240 ms. with a value of about

Us$ 250,000Properties The heartwood is yenowish brown

to reddish brown, with a golden or occasionallypurplish shine, and distinctly demarcated fromthe 3-7 cm wide pale yellow to pinkish brownsapwood. The grain is interlocked, texturecoarse. Radialsurfaces show a nice figureThe wood is moderately heavy, with a densityof 500-700(-820) kg/ina at 12% moisture con-tent. It dries slowly, but generally with littledegrade. The rates of shrinkage are moderatelylow, from green to oven dry 2.2-3.5% radialand 4.6-5.4% tangential. Once dry, the wood isstable in service

At 12% moisture content, the modulus of rup-ture is 65-1/4 N/mm2, modulus of elasticity9200-14,500 N/mm2, compression paraUel to

grain 41-65 N/mm2, shear 6-8.5 N/mm2, cleav-age 16-20.5 N/min, Janka side hardness 3870-5600 N and Janka end hardness 4800-6760 N.

The wood generally saws and works easily withordinary hand and machine tools, but the forcerequired depends on the density. A small cut-ting angle is needed to avoid tearing. The useof a filler is recommended to obtain a good fin-ish. The wood holds nails and screws well, butpre-boring is advised to prevent splitting. Thegluing, steam-bending and veneering proper-ties are all good. The wood dust may causeirritation to nose and throat.The heartwood is durable. It is resistant to

fungi and dry-wood borers, and usually also totermites, although wood from Liberia is report-edly liable to termite attack. The heartwood isresistant to impregnation with preservatives,the sapwood is permeableThe bark from roots and hole contains highamounts of saponins, and the root bark alsoalkaloids. The content of protein in the leaveswas 38%, and that of crude fibre 41%, both ofwhich are comparatively high. In another testthe crude protein content was only about 25%,while the leaves had a GIN ratio of 11.4, a 11g-rim/N ratio of 10.3 and a polyphenoVN ratio of0.82, indicating that the leaf litter is slow todecompose.

Description Large deciduoustree up to 45(-50) in tall; hole branchless for up to 22(-30) in,straight and cylindrical, up to 100(-130) cm indiameter, without buttresses or sometimeswith small, thick buttresses up to 1.5 in high;bark yellowish brown to dark grey, rough andscaly, inner bark fibrous, yellowish to orange-brown, with darker stripes and clear or honey-coloured gum; crown dome-shaped, heavilybranched, with fairly spreading branches;young branches densely rusty hairy. Leavesalternate, bipinnately compound with 3-7pairs of pinnae; stipules awl-shaped, caducous;petiole (2-)3-4.5(-6) cm long, at the middle ofupper side with a sessile gland, rachis (2-)4-15cm long, rusty hairy; leaflets in 6-14(-20) pairsper pinna, sessile, elliptical to oblong, slightlyoblique, I-2.5 cm x 0.5-I cm, rounded at apex,densely pubescent belolv. Inflorescence an axil-Iary head on (2-)5-10 cm long peduncle. Flow-ers bisexual, regular, 5-merous, white to yel-low, subtended by up to 7 min long bracteoles;pedicel up to 3 mm long; calyx 3-6 mm long,with long tube and short lobes, densely rustypubescent outside; corolla 9-13 mm long, withc. 6 mm long tube, rusty pubescent outside;stamens numerous, 3-5.5 cm long, united into

11

I, ,,2

I"

3

--.

I

densely hairy leaves and flowers, and in Itsstamen filaments being red above (whitish orgreenish in A1bizioferrugineo). The two speciesare much confused in the literature, and usedfor similar purposes, as timber, firewood, char-coal, ornamental, forage and medicinal plantA1bizio congrio is a smaller tree, rarely up to35 in tall, and occurs from C6te d'TVoire east-wards to Ethiopia and Kenya, and south toeastern Tanzania, Zambia and northern All-gola. In West Africa A1bizio conorio occursparticularly in the transition zone betweensavanna and dry forest, whereas A1bizio Ier-

characteristic for semi.Is morerug, neo

deciduous forest. In East Africa the formeroccurs in wooded grassland and riverIne forestat 850-1700 in altitude, the latter in lowlandrainforest at 800-1200 in altitude.A1bizio moldcophyllo (ARIch. ) Walp. may beconfused with A1bizio Ierrugineo, but it is asmalltree up to 15 in tall, with grey to whitishhairs on the calyx. It occurs from Senegal toUganda in wooded savanna and dry forest, andits wood is used to make pestles and as fire-wood. Its gum is used to mend broken pottery,its foliage is browsed by camels, and its rootsare used to treat conjunctivis and backacheA1bizio tongonyicensis Baker f. (paperbarkalbizia) also shows some resemblance to A1-bizio Ierrugineo, but it differs in its peeling,brownish red, papery bark revealing thecreamy young bark, its broader leaflets, itsflowers usually produced before the new leaves,and its thicker pod valves. It is a small to me-dium-sized tree up to 20 in tall of deciduouswoodland from Kenya and Tanzania south toMozambique and South Africa and from therewest to Angola. Its light and whitish wood Isoccasionally used, e. g. for carving. Root powderis rubbed into scarifications to treat swollen

legs and is used as a tonic. A root infusion isdrunk against impotence and a bark decoctionto treat cough. Pods and seeds of A1bizLo ton-gonyicensis are poisonous to livestock; youngpods are most toxic

Anatomy Wood-anatoimcal description OAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13:simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); 26: intervesselpits medium (7-10 prn); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 43: mean tan-

\

.--.

*

^

4

A1bi2io fryrugineo - I, powering twig, 2, leg/let,3, flower, . 4, fruit.Redrown und oddpted by AChmod Sotiri Nur-hornon

a tube in lower half; ovary superior, c. 3 minlong, gradually tapering into an up to 3 cmlong style. Fruit an oblong, flat pod 15-24 cm x3-5 cm, glabrous, transversely veined, yellow-ish brown when ripe, opening with 2 paperyvalves, c. 10-seeded. Seeds flattened globose toellipsoid, 7-10 mm x 4.5-8 mm. Seedling withepigeal germination; hypocoty1 4-6 cm long,epicoty1 I-2.5 cm long; cotyledons c. I cm long,thick and fleshy, rounded, early caducous; first2 leaves opposite, one pinnately compound andthe other hipinnately compound


prises about 120 species and occurs throughoutthe tropics. Approximately 35 species are foundin continental Africa and about 30 in Madagas-car. It is characterized by the head-like milo-

with I-2 central flowers modified,rescence,

functionally male and having a larger, nectar-producing staminal tube. Molecular analysesshowed that A1biziu is heterogeneous, and arevision of the genus is neededA1bizio conurio WeIw. ex 011v. closely resem-bles A1bizio Ierrugineo. It differs in its less

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A1, BIZIA 51

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gential diameter of vessellumina ^ 200 prn; 46:S 5 vessels per square minimetre; (47: 5-20vessels per square minimetre); 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 65: septate fibres present; 69:fibres thin- to thick-walled. Axial parenchyma:79: axial parenchyma vasicentric; 80: axialparenchyma anform; 81: axial parenchymalozenge-anform; 83: axial parenchyma conflu-ent; 91: two cells per parenchyma strand; 92four (3-4) cells per parenchyma strand. Rays:(97: ray width I-3 cells); (98: larger rays coin-monly 4- to 10'seriate); 104: all ray cells pro-cumbent; 1/5: 4-12 rays per mm. Mineral in-clusions: 136: prismatic crystals present; 142prismatic crystals in chambered axial paren-chyma cells(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development In exposedsites, seedlings may grow up to 3 in tall in 5years' In Ghana seedlings reached a height of2in within 2 years. In a selection test for agro-forestry species on acid, aluminium toxic soilsin southern Garnero0n, seedlings reached aheight of hardly 2 in in 20 months, which wasless than 20% of the height of the best perform-ing species (/rigo edulis Mart. and Pteroco, pussontolinoides L'Her. ex DC. ). The roots developnitrogen-fixing nodules. A1bizio Ierrugineo ishighly dependent on arbuscular mycorrhizaeThe leaves show sleep movements at nightLeaves fall after the rainy season, and newflushes are red. In Liberia trees flower in Feb-ruary and March, and the fruits ripen in De-cember to February. The fruits dehisce on thetree and the papery valves with seeds still at-tached are spread by wind

Ecology A1bizio Ierrugirteo occurs in low-land semi-deciduous and evergreen forest, inUganda up to 1200 in altitude. In West AfricaIt Is most common in semi-deciduous forest, butit can also be found in lower numbers in for-ested savanna on the one hand, and evergreenforest on the other hand

Propagation and planting While the ger-mination rate of seeds in the light and darkmay be equal, seedlings are strong light de-manders and soon die out in the shade. Regen-eration may be abundant at the start of therainy season. There are about 14,000-20,000seeds per kg. Germination takes 4-10 days.Mechanical scarification in addition to soakingin water for 24 hours enhances germination; upto 80% germlnation can be reached after pre-treatment. Usually direct seeding is practised

Wildlings are sometimes collected for plantingExperiments in Ghana showed that A1bizioIerrugineo can be successfully propagated vege-tatively by root cuttings, without the applica-tion of rooting hormone

Management Trees coppice well, but ratesof regrowth, wood and leaf production havebeen slow in experiments

Diseases and pests In C6te d'Ivoire a highincidence of attack by bruchid beetles on theseeds of A1bizioferrugineo has been recorded.

Handling after harvest Freshly harvestedlogs do not float in water and cannot be trans-ported by river. They are usually conveyed bytruck. Although logs are moderately durable,treatment is recommended ifthey are to be leftin the forest for some time, especially to protectthe sapwood against sapstain.

Genetic resources Although A1bizio Ierru-gineo is widespread in tropical Africa, it isclassified as vulnerable in the TUGN 2006 Red11st of threatened species, due to overexploita-tion. In Ghana it is considered still common,but under serious pressure from heavy exploi-tation

Prospects In West Africa A1bizio fryrugineois considered a potential substitute for someeconomic timber species that are threatened byexploitation. However, research is needed tostudy its silviculture and interaction withother crops and other tree species in agrofor-estry systems. In Nigeria A1bizio Ierrugineohas been recommended for planting in fuel-wood plantations.

Major references ATIBT, 1986; Bolza &Keating, 1972; Burki11, 1995; 01RAD ForestryDepartment, 2003; Farmer, 1972; Latham,2004; Neuwinger, 1998; SIepel, Poorter & Haw-thorne, 2004; Takahashi, 1978; Voorhoeve,1979.

Other references Addae-Mensah &Ayarkwa, 1998; Adjanohoun at a1. , 1989; Dan-quah, 2000; Ghana Forestry Commission, 2004;GIIbert & Boutique, 1952; Gillon at a1. , 1992;Habte & Musok0, 1994; Hawthorne, 1990;Hawthorne, 1995; InsideWood, undated; Kan-megne at a1. , 2000; Katende, Birnie & Tengn5. s,1995; Neuwinger, 2000; Normand & Paquis,1976; 0yun, 2006; Twumasi, 2001; Ulzen-Appiah & Flawatsror, 1993; van Wyk & vanWyk, 1997; Viniers, 1989; Wester & Hogberg,1989; World Agroforestry Centre, undated

Sources of illustration Viniers, 1989Authors K. Twum-Ampofo

ALBIZIAGLABERRiMA (Schumach. & Thonn. )Benth.

Protologue London Journ. Bot. a 88 (1844)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names Muanza (Po). Mkenge

maii, ingelenge (Sw).Origin and geographic distribution A1bizzo

globerrimo is widespread from GUTnea Bissaueast to Kenya and south to Angola, Zimbabweand Mozambique; also in Comoros and westernMadagascar.

Uses The wood of A1bizio globerrimo (tradename in Uganda: white nongo) is used for fur-niture in Uganda. In Tanzania it is used forconstruction, stools, beehives, tool handles andgrain mortars, and in Malawi for doors, bedsand implements (e. g. mortars). It is also suit-able for light and heavy flooring, Interior trim,joinery, mine props, shipbuilding, vehicle bod-Ies, railway sleepers, toys and novelties, boxesand crates, carving and plywood making. Thewood serves as firewood and for charcoal pro-duction. In Uganda A1bizio globerrimo Is con-sidered a good shade tree for coffee, tea, ba-nana and cocoa plantations. The foliage is usedas forage for livestock. The flowers are a beeforageIn southern Nigeria the bark is applied exter-nany to treat fever. In Tanzania a cold waterextract of the root bark is used to treat bilhar-zia, and in Benin a root decoction Ln a mixturewith other plant ingredients Is used as a bathto treat anaemia. In Gamero0n the dried pul-verized twig bark is applied to scarifications totreat blenorrhagia, a decoction of the twig barkis drunk against chest pain, and ash of burnedroots is applied to scarifications to treat livercomplaints

Properties The heartwood varies from dirtywhite to reddish brown, sometimes with darkerstripes, and is distinctly demarcated from thewhite sapwood, which is up to 7.5(-10) cmwide. The grain is straight, sometlmes Inter-locked, texture moderately coarseThe wood is moderately heavy, wlth a densityof about 660 kg/in3 at 8.5% moisture content. Itdries slowly, but with little degrade. At 8.5%moisture content, the modulus of rupture is 93N/mm2 modulus of elasticity 11,400 N/mm2,compression parallel to grain 55 N/mm2, shear13 N/mm2 Janka side hardness 5300 N andJanka endhardness 5910 N

The wood saws and works well, with littleblunting of saw teeth and tooledges. A reduced

cutting angle Is required to prevent tearing inplaning operations. A filler is needed to obtaina smooth finish. The nailing, screwing andgluing properties are satisfactory. The wooddoes not turn well. It is moderately durable,with some resistance to termite attack, butsusceptible to pinhole borers and marine bor-ers. The heartwood is resistant to impregna-tion by preservatives, the sapwood permeable.

Botany Medium-sized deciduous tree up to30 in tall; hole straight and cylindrical or ir-regular, up to 100 cm in diameter; bark surfacegrey, smooth or shallowly fissured, often withnumerous lenticels; crown flattened, umbrella-shaped; young twigs shortly halry. Leaves al-ternate, bipinnately compound with I-3(-4)pairs of pinnae; stipules awl-shaped, caducous;petio}e 2-4.5 cm long, grooved above, shortlyhairy, with a sessile gland near the middle ofupper side, rachis 2-4 cm long, sparsely pubes-cent; leaflets in 3-7(-8) pairs per pinna, withpetiolules 0.5-2 mm long, obliquely rhombic-ovate, up to 7(-9) cm x 3(-4) cm, obtuse toacute at apex, minutely pubescent to almostglabrous. Inflorescence an axillary head on 2-4cm long peduncle. Flowers bisexual, regular, 5-merous, white; pedice1 1.5-7 mm long, finelygreyish hairy; calyx 1.5-2.5 mm long, with longtube toothed at apex, pubescent outside; corolla3-5.5 mm long, with 2-3 mm long tube, pubes-cent outside; stamens numerous, 6-13 minlong, united at base, filaments white; ovarysuperior, 1.5-2 mm long, with c. 0.5 min longstipe, gradually tapering into an up to 12 minlong style. Fruit an oblong, flat pod 12-26 cm x3-4 cm, with stipe up to 8 mm long, pubescentor almost glabrous, indistinctly transverselyveined, glossy or dullbrown when ripe, openingwith 2 papery valves, c. 10'seeded. Seeds flat-tened globose to ellipsoid, 8-12 mm x 6-8 minSeedling with epigealgermLnationA1bizio globerrimo trees grow slowly. Theyform N-fixing root nodules. The pod valveswith seeds stillattached are spread by windA1biziu comprises about 120 species and occursthroughout the tropics. Approximately 35 spe-cies are found in continental Africa and about30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses have shown that A1bizio isheterogeneous, and a revision of the genus Isneeded. A1bizio globerrimo is often confusedwith A1bizio zygio (DC. ) J. F. Machr. , whichdiffers in its sessile leaflets. A1bizio globerrimo

ALBlziA 53

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is variable and 3 varieties have been distin-guished, of which var. globerrimo and var.globrescens (011v. ) Brenan (synonym: A1bizioglobrescens 011v. ) are widespread, the formerfrom Guinea Bissau to Sudan and Uganda, andthe latter from DR Congo and Kenya to Zim-babwe, Mozambique and Madagascar. Thelower side of var. globerrimo leaflets is inI-nutely pubescent, that of var. g!o6rescens leaf-lets glabrous. Var. inPIOopwensis Brenan isIntermediate and only known from Tanzania

Ecology In West and Central Africa A1biziogloberrimo occurs most commonly in semi-deciduous forest, but sometimes also in logged-over evergreen forest. In Garnero0n it is char-acteristic of secondary forest. In East Africa itcan be found in lowland rainforest, rivermeforest and evergreen bushland, in Madagascarin deciduous woodland

Management Seeds should be collected be-fore the pods dehisce, which makes collectingdifficult. They can be stored for up to 3 months,but are liable to weevil attacks; adding ashreduces insect damage. To obtain good germi-nation, seeds can be Immersed in boiling water,allowed to cool, and soaked in water for 24hours before sowing. The seedlings are classi-fled as non-pioneer light demanders. Wildlingsare sometimes collected in the forest for plant-ing. Planted trees can be managed by coppicingand pollarding. Logs may have brittle heartand tension wood seems to be rather frequentThe logs have to be treated with a preservativesoon after felling to prevent insects attackingthe sapwood.

Genetic resources and breeding A1biziogloberrimo is widespread and locally common,and notthreatened by genetic erosion.

Prospects Little is known about A1biziogloberrimo, probably partly because it is un-common in several countries within its distri-

bution area, and partly because it is confusedwith other A1bizio spp. As a multipurpose spe-cies it deserves more attention in agroforestryprogrammes.

Major references Bolza & Keating, 1972;Brenan, 1959; Burki11, 1995; Katende, BITnie &Tengnds, 1995; Takahashi, 1978

Other references Beentje, 1994; Brenan,1970; du Puy at a1. , 2002; Gilbert & Boutique,1952; Gros, 1994; Hawthorne, 1990; Haw-thorne, 1995; Lovett at a1. , 2006; Neuwinger,2000; Williamson, 1955


atBIZIAGUMMiFERA (J. F. Ginel. ) GASm.

Protologue Bull. Misc. Inform. Kew1930(5): 218 (1930).


Synonyms A1biato SOSsu myind. ) Chiov. (19/2).Vernacular names Peacock flower, smooth-

bark flat-crown (En). Farroba de Lala, inpepe(Po). Mkenge, inchapia turnbili, inchani inbao,mshai(Sw)

Origin and geographic distribution A1biziogummifero is widespread, occurring from east-ern Nigeria to western Ethiopia and Kenya,and south to Zimbabwe and Mozambique; alsoin central Madagascar

Uses The wood (trade names: red nongo,mepepe, omulera)is used for light construction,furniture, cabinet work and various imple-merits. It Is also suitable for mine props, lightflooring, joinery, interior trim, panelling, fram-ing, toys and novelties, sporting goods, boxes,crates, carvings, peeled and sliced veneer, ply-wood, hardboard and particle board. Logs aretraditionally used for the construction of ca-noes. The wood is sometimes used as firewood

and for making charcoal. The wood pulp issuitable for paper productionA1bizio gummifero is planted as an ornamentalshade tree. It is valued as a shade tree for

crops, e. g. in coffee plantations in Ethiopia, teaplantations in Malawi and vanilla plantationsin Madagascar, and is also planted or retainedfor soil conservation and improvement. Thegum from the bark is sometimes used in con-fectionery.Various plant parts are used in traditionalmedicine. In Kenya a bark infusion is taken to

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treat malaria, in Uganda to hasten parturitLonThe pounded bark is used in Kenya as a snuffto treat headache, and in Tanzania it is appliedexternalIy to treat scabies. In eastern DRCongo a bark maceration is used as a bodywash and drink to treat psoriasis. The rootsand leaves are purgative and are used inMadagascar to treat diarrhoea and eye trou-bles. In Malawi roots are soaked in water for10 minutes, and the liquid is drunk to relievethe pain caused by sprains. In Madagascar aleaf decoction is reputed to have antitussiveactivity and is administered to treat asthma;leaves are applied to sores and fractures. InKenya pounded roots are added to a bath totreat skin diseases, and an extract of crushedpods is drunk to treat stomach-ache. InUganda roots are used to treat sleeping sick-ness. The foliage is browsed by goats. The flow-ers produce nectar for bees, which often buildnests in the trunk. The leaves are said to

quicken the rlperilng of bananasProduction and international trade The

timber of A1bizto gummifero is mainly usedlocally and probably not much traded interna-tionally. Production and trade statistics are notavailable

Properties The heartwood is yellowish brownor reddish brown, often with a golden tinge,and distinctly demarcated from the 7-10 cmwide pale yellow or white sapwood. The grainis straight or interlocked, texture medium to

oftenQuarter. sawn surfaces are

properties are satisfactory, and steam bendingproperties moderate. The wood dust may causeirritation to nose and throat.

Reports on durability of the heartwood arecontradictory, but in general it is susceptible tofungal, wood borer and termite attack. Theheartwood is resistant to impregnation by pre-servatlves

In an experiment in Ethiopia, the leaves andtwigs of A1bizio gummifero contained per 100 gdry matter: N 3.8 g, P 0.2 g, K 1.5 g, 11gnin 26g, soluble polyphenols 8.5 g; they had a 01Nratio of 12. Mulching a maize crop wLth theleaves and twigs resulted in a smaller yieldincrease than green manure of the other spe-cies tested, probably due to the low P and Kcontent of the A1bizio gummifero green ina-

ALBiziA 55

coarse.

striped.The wood properties of A1bizio gummifero arevariable and depend on the origin of the wood,that from Madagascar being heaviest andstrongest. The wood is moderately light tomoderately heavy, with a density of 430-800kg/in3 at 12% moisture content. It dries slowly,but generally with little degrade. The shrlnk-age rates from green to oven dry are 2.9-3.5%radial and 8.2-8.7% tangential. Once dry, thewood is fairly stable in service. At 12% ino1s-ture content, the modulus of rupture was 75N/min2 modulus of elasticity 8900 N/min2,compression parallel to grain 40.5 N/mm2,shear 12 N/min2 cleavage 82 N/mm and Jankaside hardness 2840 N in a test of wood from

UgandaThe wood generally saws and works fairly eas-ily with ordinary hand and machine tools, butsawn and planed surfaces tend to pick up. Theuse of a filler is necessary to obtain a good fin-ish. The wood holds nails and screws well anddoes not split easily. The gluing and staining

nure

A dichloromethane extract of A1bizio gummi-fryo root bark showed considerable in-vitroantitrypanosomal activity, with an IC50 valueof 0.07 ILg/inI, which confirms its use as a tradi-tional treatment of sleeping sickness. Extractsalso showed in-vitro antimalarial activity

against Piersmodium folciporum, althoughmuch less than chloroquine used as referencedrug. The presence of triterpenoid saponins,sapogenin Iactones and macrocyclic sperminealkaloids (budmunchiamines) has been re-ported forthe stem bark. These lastcompoundswere active against gram-positive and grain-negative bacteria

Adulterations and substitutes The wood of

A1bizio zygio (DC. ) J. F. Machr. is very similarto that of A1bizio gummifero and is used forthesame purposes.

Description Medium-sized deciduous treeup to 30 in tan; hole straight and cylindrical,up to 75(-100) cm in diameter, without but-tresses or with small, thick buttresses; barkyellowish to grey, usually smooth, inner barkwith clear gum; crown flattened; youngbranches finely pubescent, but soon glabres-cent. Leaves alternate, bininnately compoundwith (3-)5-7(-8) pairs of pinnae; stipuleslanceolate, up to 7 mm long, caducous; petiole2.5-4.5 cm long, near the base of upper sidewith a sessile gland, rachis 4.5-11 cm long,pubescent; leaflets in 6-17 pairs per pinna,sessile, obliquely rhombic, up to 2 cm x I cm,often auricled at proximal side of base, obtuseto acute at apex, glabrous but with some hairson inidrib and margins. Inflorescence an axil-jary head on a 2.5-5 cm long peduncle. Flowersbisexual, regular, 5-meTous, reddish white,almost sessile, subtended by up to 6 mm long,

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56 TIMBERSl

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revision of the genus is needed. A1bizio gum-milero is related to A1bizio zygio, and hybridsbetween the two species have been recorded.A1biaiu gummifero is frequently confused withA1bizio odinnthifolio (Schumach. ) W. Wight,which differs in its pubescentleaflets and podsA1bizio grandibrocteoto Taub. is anotherclosely related species. Like that of A1btziogummi/erg, its wood is known under the tradename 'red nongo'. It differs from A1bizio gum-milero in having fewer leaflets per pinna (as inA1bizio zygio) and broad bracts and stipules.Hybrids between A1bizio gummifero and A1-bizio grondibrocteoto have been recorded. A1-bizio grondibrocteoto occurs in eastern DRCongo, Rwanda, Burundi, southern Sudan,south-western Ethiopia, western Kenya,Uganda and northern Tanzania. Its wood issimilar to that of A1bizio gummifero and usedfor the same purposes. It is also used as fire-wood and for charcoal production. In DR Congoa leafinfusion, together with other ingredients,is used in a vapour bath to treat fever, inUganda the pounded leafis taken mixed withother ingredients to treat diarrhoea, and inKenya a root infusion is drunk to treat tonsilli-tis. A methanolic extract of leaves of A1bizio

grandibrocteoto and saponins isolated fromthis extract have shown in-vitro antitumouractivity against KB and MCF7 celllines. LikeA1bizio gummifero, A1bizio grondibrocteoto isplanted as an ornamental and shade tree, andto improve the soilA1bizio Dirtdis E. Fourn. , a tree up to 20 in tallfrom Madagascar, may be confused with A1-bizio gummifero because of its obliquely Thorn-bic leaflets, but it differs in having stipels atthe base of pinnae and stamens united at basein a much shorter tube. In northern and east-ern Madagascar the wood of A1bizio Dirtdis isused for flooring and furniture. A1bizio inoho-100 Capuron is close to A1bizio Dirtdis, but hasoblong to elliptical leaflets (not rhombic). It is asmalltree up to Toe15) in tall, rather frequenton sandy soils in southern Madagascar. Itswood is used for construction. A1bizio momoeoViniers is a smalltree up to 15 in tall with ahole diameter up to 60 cm, widespread in west-ern and southern Madagascar. It resemblesA1bizio gummifero, but can be distinguished byits grooved leaf rachis, almost completely gla-brous leaflets and pubescent pods. Its wood isbrown and hard and used for construction and

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caducous bracteoles; calyx obconica1, 2-5 mmlong, minutely pubescent outside; corolla 7-12mm long, with 4.5-6 mm long tube, pubescentoutside; 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 styleFruit an oblong, flat pod 10-21 cm x 2-4 cm,with stipe c. I cm long, glabrous, transverselyveined, pale brown to reddish brown when ripe,opening with 2 papery valves, 9-12-seededSeeds flattened globose to broadly oblong, 8-12min x 7-loinm.


prises about 120 species and occurs throughoutthe tropics. Approximately 35 species are foundin continental Africa and about 30 in Madagas-car. It is characterized by the head-like inno-rescence, with I-2 central flowers modified,functionally male and having a larger, nectar-producing staminal tube. Molecular analysesshowed that A1bizio is heterogeneous, and a

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canoes

Anatomy Wood-anatointcal description OAWAhardwood codes):

Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13:simple perforation plates; 22: intervessel pitsalternate; 23?: shape of alternate pits polygo-na1; 25: intervessel pits small (4-7 pin); 26intervessel pits medium (7-10 pin); 29: ves-tured pits; 30: vessel-ray pits with distinct bor-ders; similar to Intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 F1m;43: mean tangential diameter of vessellumina;a 200 prn; 46: s 5 vessels per square minimetre;(47: 5-20 vessels per square minimetre); 58:gums and other deposits in heartwood vesselsTracheids and fibres: 61: fibres with simple tominutely bordered pits; 65: septate fibres pre-sent; 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: 76axial parenchyma diffuse; (79: axial paren-chyma vasicentric); 80: axial parenchyma all-form; 81: axial parenchyma lozenge-alitorm;(83: axial parenchyma confluent); (90: fusiformparenchyma cells); 91: two cells per paren-chyma strand; 92: four (3-4) cells per paren-chyma strand. Rays: 98: larger rays commonly4- to 10'seriate; 104: all ray cells procumbent;(1/4: S 4 rays per min); 115: 4-12 rays per mmMineral inclusions: 136: prismatic crystalspresent; 142: prismatic crystals in chamberedaxial parenchyma cells(P. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Trees are capa-ble of growing rapidly. The roots develop nitro-gen-fixing nodules containing Bradyrhizobiumbacteria. A1bizio gummifero trees live in asso-ciation with arbuscular mycorrhizae

Ecology A1bizio gummifero occurs in rain-forest and riverIne forest, sometimes also insavanna vegetation close to forest, usually athigher altitudes, up to 2500 in, but sometimesnear sea-level. It is locally common. In Zim-babwe it is reportedIy fire resistant and onlyslightly sensitive to frost

Propagation and planting Seedsfor plant-ing should be collected from the trees beforethe pods dehisce, to avoid insect damage. Thereare 10,000-15,000 seeds per kg. Fresh seedsmay have a germination rate of up to 80% in 3-10 days, and do not requlre pre-treatmentHowever, in Ethiopia seedcoat-induced dor-mancy has been recorded, hampering complete,rapid and uniform germination. Scarificationimproves the germination capacity and vigourof the seeds. Seeds can be stored for more than

one year in a sealed container in a cool place,

after adding ash to reduce insect damageStored seeds should be soaked in water before

planting. A fine and firm seedbed is requiredfor even germination and vigorous seedlinggrowth. Wildlings are sometimes collected forplanting.

Management The addition ofNPKfertilizeris recommended for seedlings. In planting ex-penments in Ethiopia, A1bizio gummiferoshowed a survival rate of 94%. Young plantedtrees can be managed by CDPpicing and loppingThey are often damaged by strong wind, fromwhich they should be protected.

Handling after harvest Freshly harvestedlogs float in water and can be transported byriver. Treatment of the logs with preservativesis necessary if they are to be left in the forestfor some time, to avoid damage by fungi orInsects.

Genetic resources A1bizio gummifero iswidespread and locally common, also in secon-dary forest. It is therefore not easily liable togenetic erosion

Prospects A1bizio gummifero is a multipur-pose species. It seems to have good prospects asa commercial timber tree in sustainably inari-aged forests and in afforestation projects. Fur-ther tests are needed to confirm Its promise asan auxiliary tree in agroforestry systems. Onceadequate vegetative propagation techniqueshave been developed, A1bizio gummifero de-serves to be promoted for planting

Major references Bekele-Tesemma, BITnie& Tengnas, 1993; Bolza & Keating, 1972;Burki11, 1995; du Puy at a1. , 2002; Katende,Birnie & Tengnas, 1995; Keay, 1989; Takaha-shi, 1978; Thuhn, 1989; Tigabu & Oden, 2001;World Agroforestry Centre, undated.

Other references Beentje, 1994; Brenan,1970; Chifundera, 2001; Coates Palgrave, 1983;DebeUa at a1. , 2000; Debray, Jacquemln &Razafindramba0, 1971; Decary, 1946; Eggeling& Dale, 1951; Freiburghaus at a1. , 1996; GII-hert & Boutique, 1952; Hamill at a1. , 2000;Hines & ECkman, 1993a; InsideWood, undated;Kokwar0, 1993; thief at a1. , 2005; Maundu ata1. , 2001; Of una at a1. , 1996; Rukunga &Waterinari, 1996; Sommerlatte & Sommerlatte,1990; Williamson, 1955; Wubet at a1. , 2003.

Sources of illustration Coates Palgrave,1957

AuthorsA. Maroyi

ALBlziA 57

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A1, BIZiAscHiMPERLANA Onv

Protologue F1. trop. Mr. a 359 (1871)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names Large-podded albizia,

forest long-podded albizia (En). Mkenge (Sw)Origin and geographic distribution A1bizto

schimperiono is widespread, from eastern DRCongo, southern Sudan and Ethiopia, south toZimbabwe and Mozambique

Uses The wood of A1btzio schimperiono is usedfor construction, joinery, tool handles, bee-hives, stools, grain mortars, spoons, needles,plywood and matchboxes. It also serves asfirewood and for charcoal production. A1btzioschLmperiono is planted in agroforestry sys-terns as a shade tree and for soil conservationand Improvement. It is browsed by cattle andbranches are lopped and fed to goats. Reportsof its forage value range from poor to fair. Theflowers serve as a source of nectar for honey-bees. In Kenya a root infusion added to por-ridge is used against headache and other painsThe stem bark is used in Tanzania to treatwarts. The bark is used as a cough remedy andas a soap substitute, while the ash of the barkis added to snuff tobacco to give pungency. InEthiopia ground seeds mixed with water areused as an Insecticide. Wool can be colouredyellow with the leaves

Production and international trade There

are no trade statistics for A1bizio schimperionotimber, but it has recently been included incommercial trade from the East UsambaraMountains in Tanzania

Properties The wood is fairly strong, workseasily and is not resistant to termites. Leavesand twigs harvested from mature trees at theend of the rainy season contained 25.6 g crudeprotein per 100 g dry matter, but the digestfoil-ity is rather low. Spermine alkaloids and sev-eraltriterpenes (Iupeol, Inpenone, o1eanoic acidand bederagenin) have been isolated from thebark of A1bizio schimperiono.

Botany Medium-sized deciduous tree up to30(-35) in tall; hole straight and cylindrical orlow-branched, up to 70 cm in diameter; barksurface grey or brown, smooth or rough; crownflattened or rounded, often umbrella-shaped;young twigs brown pubescent. Leaves alter-nate, hmmnately compound with (I-)2-7(-11)pairs of pinnae; stipules awl-shaped, caducous;petiole near the base of upper side with a ses-SIIe gland, rachis densely to sparsely pubes-cent; leaflets in 6-21(-23) pairs per pinna, al-

most sessile, obliquely rhombic or oblong, up to2(-3) cm x I(-1.5) cm, rounded to acute atapex, pubescent below. Inflorescence an axil-Iary head on 2.5-5 cm long peduncle. Flowersbisexual, regular, 5-merous, white or pale yel-low; pedice1 I-6 min long, brown pubescent;calyx 1.5-2.5 mm long, with long tube toothedat apex, brown or sometimes grey pubescentoutside; corolla 3-7.5 mm long, brown or some-times grey pubescent outside; stamens numer-ous, 7-12 min long, united at base, withcreamy white filaments; ovary superior, gradu-ally tapering into slender style. Fruit an ob-long, flat pod 15-34 cm x 2-6 cm, with 1.5-2cm long stipe, shortly pubescent, transverselyveined, dullbrown when ripe, tardily dehiscentor indehiscent, c. 8-seeded. Seeds flattenedglobose to ellipsoid, 9-11 mm x 6.5-8 mm.Seedling with epigealgermination.A1biaio comprises about 120 species and occursthroughout the tropics. Approximately 35 spe-cies are found in continental Africa and about

30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. Mo-Iecular analyses showed that A1bizio is hetero-geneous, and a revision of the genus is neededA1bizio schimperiono is variable, especially inleaf characteristics and indumentum colour

Three varieties have been distinguished: varschimperiono occurring throughout the rangeof the species, var. omaniensis (Baker f. )Brenan characterized by few leaflets per pinnawith the terminal pair largest and restricted toTanzania, and var. tephrocolyx Brenan charac-terized by the grey colour of the hairs on calyxand corolla and occurring in southern SudanandUgandaA1bizio schimperiono trees grow slowly. Theroots develop nitrogen-fixing nodules contain-ing Bradyrhizobium bacteria. A1bizio schimpe-norio trees live in association with arbuscular

mycorrhlzae. Flowering occurs at the end of thedry season; pods take about 5 months to ina-ture. The pods tend not to split open and there-fore rot on the groundA1bizio omuro (Roxb. ) Boivin (bitter albizia)resembles A1bizio schimperiono, but differs insmaller leaflets and shorter pealcels. It is ashrub or smalltree up to 15 in tall, occurringfrom Eritrea and Ethiopia south to northernSouth Africa, and in India and Sri Lanka. Itswood is used for poles and Implements, and asfirewood and for charcoal production. Tests inTanzania showed that A1bizio omuro could be

:

promoted for commercial utilization for rumi-ture and construction. It is also planted for soilimprovement and conservation, its foliage isused as fodder, and its roots, leaves and podsare used in traditional medicine. A root infu-SIon is drunk to treat pneumonia, tuberculosis,infertility of women and as an aphrodisiac,whereas roots are applied externalIy to wartsand against uterus complaints. The bark isused to treat jaundice and mouth inflamma-tions, the leaves for the treatment of diarrhoea,oedema and wounds, and the pods as an emeticand for the treatment of cough and malariaThe roots are also used as a soap substituteA1bizio horueyi E. Fourn. also differs from A1-bizio schimperiono by its smaller leaflets andshorter pedicels, and from A1bizio omuro by itsleaflets, which are acute and asymmetrical atapex in A1bizio horueyi and obtuse and sym-metrical at apex in A1biz, o omuro. It is a smalltree up to 15 in tall and occurs from Kenyasouth to northern South Africa. Its wood isused for building poles and as firewood. InTanzania the roots are used to treat epilepsy,cystitis, vomiting, Infertility and to preventmiscarriage. The foliage is browsed by ITve-stock

A1bizio forbesti Benth. , occurring from Tanza-nia south to South Africa, also resembles A1-bizio schimperiono, but is characterized by Itssessile flowers with larger calyx and narrowerseeds. Its wood is used in southern Africa for

posts of native housesEcology A1bizio schimperiono occurs In up-

land forest and evergreen bushland at 900-2600 in altitude. It may be dominant in theupper canopy in southern Sudan and EthiopiaIn Tanzania it grows in regions with a meanannual temperature of 14-20'C and an annualrainfall of at least 1000 min, often in open sec-ondary forest and moritane riparian forest.

Management A1bizio schimperiono is capa-ble of regeneration in more or less closed forest,with seedlings surviving in the shade. Theseeds are often damaged by insects. Methodshave been developed to separate sound anddamaged seeds using near infrared spectros-copy. The 1000-seed weight is 90-125 g. Me-chanicalscarification as wellas treatment withsulphuric acid or hot water (boiling or 75'C)are effective ways of overcoming seed coat in-duced dormancy. Seedlings may be planted outafter 4 months in the nursery. Wildlings aresometimes collected from the forest for plant-ing. Planting in pure stands Is not recoin-mended because of the greater risk of pest

problems. Planted trees can be managed bycoppicing

Genetic resources and breeding A1bizioschtmpertono is widespread and locally coin-mon, and not threatened by genetic erosion.However, locally populations are under great

because of deforestation and poorpressure

regeneration, e. g. in Ethiopia.Prospects Although A1biziu schtmperiono is

a slow-growing tree, it is locally preferred intraditional agroforestry systems, e. g. in Ethio-pia and in the West Usambara Mountains inTanzania. It is locally a highly valued multi-purpose tree. Little Is known about many as-pects of this tree, and research is needed toevaluate its potential role as a timber andagroforestry tree on a wider scale

Major references Bekele-Tesemma, Birnie& Tengnas, 1993; Brenan, 1959; Sommerlatte& Sommerlatte, 1990; Thulin, 1989; Tigabu &Oden, 2003.

Other references Demel Teketay, 1996;Graham at a1. , 2000; Hines & ECkman, 1993b;Kaitho at a1. , 1998; Kokwar0, 1993; Lovett ata1. , 2006; Mahmoud at a1. , 1995; Msanga &Maghembe, 1986; Neuwinger, 2000; Roe at al. ,2002


ALBiziA 59

ALBIZIAVERSICOLOR WeIw. ex 011v.

Protologue F1. trop. Mr. 2: 359 (1871)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names Poison-pod albizia, large-

leaved false thorn (En). Mchani ridovu, inkenge,innduruasi, induruasi(Sw).

Origin and geographic distribution A1bi2iouersicolor is widespread from DR Congo east toKenya, and south to Namibia and SouthAfrica

Uses The wood is locally used for smallboats,tool handles, mortars and other kitchen Im-PIements, containers, casks and musical in-struments. It is suitable for light construction,light flooring, joinery, furniture, cabinet work,decorative work, veneer, plywood, drainingboards, hardboard and particle board. It is alsoused as firewood (although it may spark badly)and for charcoal production. The bark has beenused for tanning and the flowers serve as asource of nectar for honey bees. The inner barkis used for making rope. Roots boiled with wa-ter can be used as a substitute for soap. A1biziouersicolor is planted as an ornamental shadetree.

.

60 TIMBERSl

Root and bark decoctions are used as an

arithelmintic and purgatrve, and to treat swol-Ien glands and venereal diseases. Dried andpowdered roots are taken or sniffed to treatheadache and sinusitis, and a root macerationis taken against gonorrhoea. A bark decoctionis used to treat anaemia, and it is applied ex-ternally to treat ophthalmia and skin rash. Abark maceration is taken against cough, andbark powder is sniffed for the same purpose.

Properties The heartwood is pale to purplishbrown, often darker striped, sometimes almostblack; It is distinctly demarcated from thewhite sapwood, which is up to 5 cm wide. Thegrain Is wavy or interlocked, texture coarse.The wood is moderately heavy, with a densityof 560-770 kg/ina at 12% moisture content. Itdries slowly with little degrade, but surfacechecking occurs in excessively cross-grainedpieces. The rates of shrinkage are low: fromgreen to 12% moisture content 1.1% radial and1.8% tangential, and from green to oven dry1.8% radial and 3.0% tangential. Once dry, it isstable In service

At 12% moisture content, the modulus of rup-ture is 52-57 N/mm2, modulus of elasticity6500-7700 N/min2, compression parallel tograin 32-41 N/min2, shear 10-12 N/min2cleavage 54 N/mm radial and 69 N/mm tangen-tialand Janka side hardness 4630 N

The wood saws and works well, but the sur-faces of quarter-sawn boards may pick up. Theuse of a filler is needed to obtain a smooth fin-

ish. The wood does not hold nails well, and pre-boring is required. The jointing and gluingproperties are good, but steam bending proper-ties are usually poor. The wood dust may causeserious irritation to nose and throat.

The wood is moderately durable, but liable toattacks by pinhole borers and marine borersReports on its resistance to termites vary fromsusceptible to resistant. The heartwood is re-SIStant to Impregnation by preservatives, thesapwood is permeablePods and seeds are poisonous to livestock;young pods are most toxic, but livestock ismore often poisoned as a result of eating fallenpods. Cattle, sheep and goats may develop hy-persensitivity, intermittent convulsions andhigh temperature, and may die from heart fail-ure, but most animals recover without treat-merit. The disease is called albiziosis, and iscaused by the presence of methylpyridoxine.Treatment of poisoned sheep with pyridoxinehydrochloride resulted in recovery. Kaempfero1glycosides and several triterpenes (Iupeol, Iu-

penone, betulinic acid and acacic acid Iactone)have been isolated from the bark of A1biziuuersicolor.

Botany Small to medium-sized deciduoustree up to 20 in tall; bole usually short andbranchless for up to 5 in but sometimes up to12 in, straight and cylindrical, up to 60(-150)cm in diameter; bark surface greyish brown,rough and fissured; crown flattened, open, withspreading branches; young twigs rusty brownhairy. Leaves alternate, hipinnately compoundwith I-4(-5) pairs of pinnae; stipules awl-shaped, caducous; petiole 5-7 cm long, groovedabove, near the base of upper side with a ses-SIIe gland, rachis 8-20 cm long, rusty brownhairy; leaflets in 3-6 pairs per pinna, almostsessile, obliquely broadly obovate to almostorbicular, up to 6.5 cm x 5 cm, rounded toslightly notched but inucronate at apex, Ieath-ery, densely yellowish or reddish brown hairybelow. Inflorescence an axillary head on 3-6cm long peduncle. Flowers bisexual, regular, 5-merous, white to greenish yellow; pedicel up to2 min long; calyx 4.5-8 mm long, with longtube toothed at apex, rusty brown pubescentoutside; corolla 8-12 min long, rusty brownpubescent outside; stamens numerous, 2.5-5.5cm long, united at base, filaments reddish;ovary superior, gradually tapering into a longand slender style. Fruit an oblong, flat pod 10-30 cm x 3-6.5 cm, almost glabrous, indistinctlytransversely veined, yellowish brown to red-dish brown when ripe, opening with 2 paperyvalves, c. 6-seeded. Seeds flattened ellipsoid, 9-13 min long.In southern Africa A1bizio Dersicolor trees usu-

ally flower in August-December and fruits areripe in December-March. The pod valves withseeds still attached are spread by wind. Thegrowth rate of trees is moderate. The rootsdevelop nodules containing nitrogen-fixingbacteria.


30 in Madagascar. It is characterized by thehead-like inflorescence, with I-2 central flow-ers modified, functionally male and having alarger, nectar-producing staminal tube. MD-Iecular analyses showed that A1bizio is hetero-geneous, and a revision is needed.

Ecology A1bizio uersicolor occurs in decidu-ous woodland and wooded grassland up to 1700in altitude. It prefers well-drained soils with ahigh water table.

Management Seeds should be collected be-

fore the pods dehisce, which makes collectingdifficult. Seeds on the ground are often infestedby insects. One kg contains 6000-8000 seedsSeed germination is usually good, up to 90%,and is completed in 30 days. Fresh seeds do notrequire any treatment, but stored seeds shouldbe soaked in water for 6 hours before sowingThey can be stored for long periods when keptunder dry and insect-free conditions. Seeds arepreferably sown in seedling trays filled with amixture of river sand and compost (4:1)Propagation by cuttings and root suckers hasbeen successful. Planted trees can be managedby lopping and pollarding. Harvested logs areoften of poor quality, being short and irregularor with heart rotin larger sizes

Genetic resources and breeding Thereare no indications that A1bizio uersicolor Isthreatened by genetic erosion, although it isIOCaUy uncommon within its wide area of dis-tribution, e. g. in Uganda

Prospects A1biziu uersicolor has good pros-pects as an auxiliary tree in agroforestry sys-terns, improving the soil with its nitrogen-fixing root nodules, providing mulch with itsleaf litter, reducing erosion with its large root-ing system, and protecting crops from too muchsun. This, together with other local uses, espe-clany as timber tree and medicinal plant,makes it a multipurpose tree worthy of cultiva-tion on a wider scale. However, its toxic fruitComponents warrant some Caution In areassubject to heavy grazing. It is also recoin-mended as an ornamental tree for large gar-dens and parks

Major references Bolza & Keating, 1972;Chilufya & Tengnas, 1996; Katende, Birnie &Tengnas, 1995; van Wyk, van Heerden & vanOudtshoorn, 2002; World Agroforestry Centre,undated

Other references Bryce, 1967; Gelfand ata1. , 1985; Kokwar0, 1993; Neuwinger, 1998;Neuwinger, 2000; 0molo, Chhabra & Nyagah,1997; Palmer & Pitman, 1972-1974; Scott,1950; Soldan, van Inzen & Edelsten, 1996;Zambia Forest Department, 1979b


Pau de raio, sancalama (Po). Nongo, inkenge(Sw)

Origin and geographic distribution A1biziozygio is widespread in tropical Africa, occur-ring from Senegal in the west to Kenya in theeast and northern Angola and Tanzania in thesouth.

Uses The wood is known under the tradenames nongo (general), red nongo (Uganda)and okuro (Ghana). It is used for indoor con-struction, light flooring, furniture, canoes, Im-PIements (e. g. pestles, hoe-handles), carving,veneer and plywood. It is also suitable for join-ery, interior trim, ship and boat building, vehi-cle bodies, toys and novelties, turnery, boxes,crates, matches, hardboard and particle boardThe wood iseommonly used asfirewood and forcharcoal production. The wood pulp is suitablefor paper productionA1bizio zygio is planted as an ornamentalshade tree, roadside tree and fire break. It isvalued as a shade tree, e. g. in cocoa and coffeeplantations, and as a support for yarn. It isconsidered useful for soilimprovement. Youngleaves are eaten cooked as a vegetable or addedto soups and sauces. The foliage is ranked aslow-quality fodder, but suitable for use in goatfeeds and sometimes also given to sheep andcattle. The flowers produce nectar which iscollected by bees. The gum from the bark isused in the food industry (e. g. as a stabilizer inthe preparation of ice cream), cosmetic Indus-try (e. g. as a thickener) and pharmaceuticalindustry (e. g. as a drug coating)In traditional medicine, bark sap is instilled inthe eyes to treat ophthalmia. A bark decoctionis administered to treat bronchial diseases,fever (including malaria) and female sterility,

A1, BIZiA 61

ALBIZIAZYGiA (DC. ) J. F. Machr

Protologue Contr. Gray Herb. 59: 3 (1919)Family Mimosaceae (Leguminosae - Mimo-


Vernacular names West African albizia (En).

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A1bizio zygio- wild

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62 TIMBERSl

and as a purgative, stomachic, antidote, vermi-ruge and aphrodisiac. Pounded or rasped barkis applied externalIy to treat yaws, sores,wounds and toothache. Ground roots are added

to food to treat cough and as an expectorant.Leaf decoctions are used to treat fever and di-arrhoea

Production and international tradeAlbizio

zygio tlinher has some Importance on the in-ternational market, although it is considered alesser-used species. It is often mixed with tim-ber of other A1bizio spp. In Ghana the exportvolume of sawn A1bizio timber in the first halfof 2004 was 240 in3 with a value of about Us$250,000

Properties The heartwood is yellowishbrown or pinkish brown to dark brown, some-times with a red tinge, and distinctly demar-cated from the wide pale yellow to grey sap-wood. The grain is straight or interlocked, tex-ture fine to coarse. Planed surfaces of the woodare lustrous

The wood is moderately heavy, with a densityof 500-720 kg/ina at 12% moisture content. Itdries slowly, but generally with little degradeThe rates of shrinkage are moderate, fromgreen to oven dry 3.0-3.4% radial and 5.7-7.1%tangential. Once dry, the wood is stable in ser-

colonic bacterial enzymes leading to release ofthe drug. The gum is also promising as bindingagent in tablet formulations, especially whenhigh mechanical strength and slower releaserates are desired.

The foliage has a high crude protein content(32%), but with a low in-vitro dry matter di-gestibility value (41.5%); it has been ranked asa low-value fodder


of A1bizio gummifero (J. F. Ginel. ) GASm. isvery similar to that of A1bizio zygio and is usedfor similar purposes

Description Medium-sized deciduous treeup to 30 in tall; hole branchless for up to 15 in,straight and cylindrical, up to 80(-120) cm indiameter, without buttresses or with small,thick buttresses; bark pinkish or greenish grey,smooth or shallowly fissured, inner bark fi-brous, yellowish to orange-brown, with darkerstripes and brownish gum; crown dome-shaped,with spreading branches; young branches pu-bescent, but usually soon glabrescent. Leavesalternate, hipinnately compound with (I-)2-3(-4) pairs of pinnae; stipules awl-shaped, ca-ducous; petiole 4-10 cm long, near the base of

vice

At 12% moisture content, the modulus of rup-ture is 69-1/8 N/min2, modulus of elasticity8400-12,000 N/min2, compression parallel tograin 42-65 N/mm2, shear 11.5-16.5 N/min2,cleavage 18-108 N/mm, Janka side hardness2790-5910 N and Janka end hardness 4500 N

The wood generally saws and works fairly eas-11y with ordinary hand and machine tools, butsawn surfaces tend to be woolly. A filler is rec-ommended to obtain a good finish. The woodholds nails and screws well, but pre-boring isadvised to prevent splitting. The gluing, stain-ing and veneering properties are satisfactory,and steam bending properties moderate. Thewood dust may cause irritation to nose andthroat.

The heartwood is fairly durable, but it is sus-ceptible to termite attack and powder post bee-tles. The heartwood is resistant to impregna-tion with preservatives, but the sapwood ispermeable.Experiments showed that A1bizio zygio gumhas good potential for use as compression coat-ing for drugs targeting the colon, being capableof protecting the core tablet in the physiologicalenvironment of the stomach and smallintes-

tine, but susceptible to degradation by the

,

2

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3

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4

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A1bizio zygio - I, flowering twig, 2, led/let, . 3,flower, 4, fruit.Redrown Grid odopted by Ishoh Syomsudin

I

upper side with a sessile gland, rachis 2-13 cmlong, pubescent but glabrescent; leaflets in 2-5pairs per pinna, sessile, obliquely rhombic toelliptical, increasing in size from base to apexof pinna, up to 8(-11.5) cm x 4(-6) cm, obtuseto acute at apex, glabrous. Inflorescence anaxillary head on 2.5-4 cm long peduncle. Flow-ers bisexual, regular, 5-merous, reddish white,sessile, subtended by up to 3 min long bracte-o1es; calyx 2-4 mm long, with long tube toothedat apex, pubescent outside; corolla 6-9(-10.5)min long, with c. 5 mm long tube, pubescent

long,outside; stamens numerous, 2-3united into a reddish tube for most of their

length; ovary superior, c. 1.5 mm long, gradu-ally tapering into an up to 2 cm long styleFruit an oblong, nat pod 10-20 cm x 2-4 cm,with stipe up to 1.5 cm long, glabrous, trans-versely veined, glossy brown when ripe, open-ing with 2 papery valves, 9-12-seeded. Seedsflattened globose to ellipsoid, 7.5-12.5 mm x6.5-9 min. Seedling with epigeal germination;hypocoty1 4-4.5 cm long, epicoty1 I-3 cm long;cotyledons rounded to broadly elliptical, thickand fleshy, early caducous; first 2 leaves OPPo-site, with one pair of pinnae


prises about 120 species and occurs throughoutthe tropics. Approximately 35 species are foundin continental Africa and about 30 in Madagas-car. It is characterized by the head-like inno-rescence, with I-2 central flowers modified,functionally male and having a larger, nectar-producing staminaltube. Molecular analysesshowed that A1bizio is heterogeneous, and arevision is needed. A1bizio zygio is related toA1bizio gummifero, and hybrids between thetwo species have been recorded

Anatomy Wood-anatomical description OAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13:simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;(25: intervessel pits small (4-7 prn)); 26: in-tervessel pits medium (7-10 pin); 29: vesturedpits; 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; 42: mean tangentialdiameter of vessellumina 100-200 pin; 43:mean tangential diameter of vessellumina ;a200 prn; 46: s 5 vessels per square minimetre;(47: 5-20 vessels per square minimetre); 58:gums and other deposits in heartwood vesselsTracheids and fibres: 61: fibres with simple to

minutely bordered pits; 65: septate fibres pre-sent; 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: 76:axial parenchyma diffuse; 79: axial paren-chyma vasicentric; 80: axial parenchyma all-form; 81: axial parenchyma lozenge-anform;83: axial parenchyma confluent; 91: two cellsper parenchyma strand; 92: four (3-4) cells perparenchyma strand; (93: eight (5-8) cells perparenchyma strand). Rays: 98: larger rayscommonly 4- to 10-senate; 104: all ray cellsprocumbent; 1/5: 4-12 rays per mm. Mlneralinclusions: 136: prismatic crystals present; 142prismatic crystals in chambered axial paren-chyma cellsre. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Seedlings de-velop quickly, and can reach 3 in tall after 2years. Trees are capable of growing rapidly andreached a bole diameter of 80 cm in secondaryforest of 40 years old. The roots develop nitro-gen-fixing nodules containing Brodyrhizobiumbacteria. A1bizio zygio trees live in associationwith arbuscular mycorrhizae. In Ghana treesflower from February to May, and ripe fruitscan be found from November to April. Thefruits dehisce on the tree and the papery valveswith seeds stillattached are spread by windIn a plantation in northern C6te d'Ivoire, 5.5-year-old A1bizio zygio trees were on average3.7 in tall, with a stem diameter of 7 cm. Treeson old termite mounds were over 6 in tall witha stem diameter over 10 cm. In plantations inthe evergreen forest zone in southern C6ted'TVoire A1bizio zygio trees had a mean annualdiameter growth of 1.6 cm, against 2.1 cm inthe semi-deciduous forest zone

Ecology A1bizLo zygio occurs in lowlandsemi-deciduous and evergreen forest, in EastAfrica up to 1400 in altitude. In West Africa itis most common in secondary forest in thesemi-deciduous forest zone, but it can also befound in lower numbers in both forested sa-vanna and evergreen forest. It is locally coin-mon and shows no preference for wet or drysites. It is tolerant to acid soils and shows some

droughttolerance.Propagation and planting Seedlings are

classified as non-pioneer light demanders. Re-generation depends on the presence of me-dium-sized to large canopy gaps in the forestConsequently, seedlings and saplings of A1biziozygio are more abundant in forest with loggingdamage than in undisturbed forest.Seeds for planting should be collected from thetrees before the pods dehisce. The 1000-seed

cm

ALEiziA 63

64 TIMBERSl

weight is 50-60 g. In C6te d'Ivoire the germi-nation rate of seeds soaked in water for 24

hours was 8-30% in 35 days; it was much bet-ter after treatment with sulphuric acid or me-chanical scarification. In Ghana pre-treatmentof the seeds with 98% sulphuric acid for 5 min-utes followed by thorough rinsing in water jin-proved germination by up to 85%, and me-chanical scarification followed by soaking for 6hours in water gave a germination rate of up to95% in 8 days. In Uganda it is recommendedthat seeds be soaked in water overnight beforeplanting. Seeds can be stored for some time ina sealed container in a cool place, after addingash to reduce insect damage. Wildlings aresometimes collected in the forest for planting.A1bizio zygio can be successfully propagatedvegetativeIy by root cuttings; the application ofindole-3-butyric acid improved rooting rates

Management It is necessary to protect youngtrees from cattle and wild herbivores, and toremove weeds during the first 2-3 years ofgrowth. Planted trees can be managed by cop-picing and pollarding

Diseases and pests In Cote d'TVoire a highincidence of attack by bruchid beetles on theseeds of A1bizio zygio has been recorded. Sen-ous attacks by borer beetles were recorded inthe savanna zone, where many planted treeswere killed.

Handling after harvest Freshly harvestedlogs sometimes float in water and can then betransported by river. They are also conveyed bytimber trucks. Although logs are moderatelydurable, treatment is recommended ifthey areleft in the forest for some time, especially toprotectthe sapwood

Genetic resources A1bizio zygio is wide-spread and locally common, particularly insecondary forest. This means that it is not eas-11y liable to genetic erosion. However, harvest-ing the bark for medicine and forthe gum maylocally reduce population sizes of the tree.

Prospects A1bizio zygio is a multipurposespecies. It seems to have good prospects as acommercial timber tree in sustainably man-aged forests, and also as a source of gum usefulin the food, cosmetic and pharmaceutical in-dustry. It has good prospects as an auxiliarytree in agroforestry systems, improving the soilwith its nitrogen-fixing root nodules, providingmulch with its leaflitter, reducing erosion withits large rooting system, and protecting cropsfrom too much sun. It deserves to be included

in agroforestry testing programmesMajor references anim Kwapong & Tekle-

balmanot, 1995; Bolza & Keating, 1972;Burki11, 1995; Katende, Birnie & Tengnas,1995; Neuwinger, 2000; Odeku, 2005; Odeku &Fell, 2005; Takahashi, 1978; Viniers, 1989;WorldAgroforestry Centre, undated.

Other references Anim Kwapong & Tekle-haimanot, 2001; Arbonnier, 2004; Arthur, Met-tle & Owusu Sekyere, 1998; Arthur at a1. , 1999;Asante, 1994; Beentje, 1994; Danquah, 2000;Dupuy, 1998; Ghana Forestry Commission,2004; Gilbert & Boutique, 1952; Gillon at al. ,1992; Hawthorne, 1990; Hawthorne, 1995; In-sideWood, undated; Irvine, 1961; Kokwaro,1993; Kpikpi, 1992; Normand & Paquis, 1976;Okafor, 1997; Sommerlatte & Sommerlatte,1990; Wester & Hogberg, 1989

Sources of illustration Viniers, 1989.Authors MM. Apetorgbor

ALSTONLABOONEIDe Wild.

Protologue Repert. Spec. Nov. Regni Veg.13: 382 (1914).

Family ApocynaceaeChromosome number2n=42

SynonymsAlstonio congensis auct. non EnglVernacular names instonia, cheesewood,

stool wood, pattern wood (En). Binien, ekouk(Fr).

Origin and geographic distribution AISto-itto boonei occurs from Senegal and Gambia towestern Ethiopia and Uganda

Uses The wood of AIStonio boonei, caned al-stonia in international trade, is used for lightconstruction, light carpentry, open boats,moulding, furniture, interior jolnery, imple-merits, boxes, crates, matches, pencils, sculp-

.

AIStonio boonei- wild

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:.

tures, and for veneer and plywood. It is locallypopular for the production of household imple-merits because of its good working propertiesand stability. In Ghana it is used for the fa-mous Asante stools, and in Nigeria for soundboxes of musical instruments of the Yoruba

people. The wood is also used as firewoodThe bark of the bole is important in traditionalmedicine. In local markets in West and CentralAfrica it is often amongst the most commonplant materials sold as crude drugs. A barkdecoction is widely used to treat malaria, ty-phoid fever, gonorrhoea, yaws, asthma anddysentery, and Is also applied to sores, ulcers,snakebites, rheumatic pain and toothache, andas a galactagogue. A maceration of the bark istaken to treatjaundice, cough and sore throat,and is applied externalIy to treat skin coin-plaints. The bark is also used as an arithelmin-tic. The latex is applied to snakebites, skincomplaints and swellings caused by filaria in-fections, and in concoction to treat fever. Theleaves are applied topicalIy to reduce oedemasand to treat sores. The latex has been used as

birdlime and as an inferior alternative for rub-her. AIStonio booneiis a useful shade tree for

coffee, tea and banana plantations.Production and international trade A1-

though AIStonio timber of Asian origin (mainlyfrom Indonesia and Malaysia) is important onthe international timber market, it is only oc-casionally exported from African countries, andthen probably mostly in mixed consignments oflight-weight timber. Although Ghana exported64,500 ina and 19,500 ina in 2000 and 2001respectively, and Gamero0n 210 ina in 2001, itis of greater importance locally. The bark ofAIStonio booneiis commonly sold on local mar-kets for medicinal purposes. In Yaound6 (Cam-eroon) the price of the bark varies from I-3FCFAper grain, depending on prescription.

Properties The heartwood is creamy whiteand indistinctly demarcated from the up to 20cm wide sapwood. The wood darkens upon ex-POSure to light. The grain is straight, occasion-any wavy, texture moderately coarse. Growthrings indistinct or absent. The wood has a dis-agreeable smellwhen greenThe wood is lightweight. The density is 360-420 kg/ina at 12% moisture content. Theshrinkage rates are moderately low, from greento oven dry 3.8-4.0% radial and 5.2-5.4% tan-gential. The wood dries rapidly, with slight riskof distortion and checking. Once dry, it is sta-ble.

At 12% moisture content, the modulus of rup-

ture is 48-73 N/mm2, modulus of elasticity5790-10,500 N/min2, compression parallel tograin 27-37 N/mm2, shear 6-7 N/min2, cleav-age 10 N/min and Janka side hardness 1820 NThe wood is easy to saw, although the presenceof latex may cause clogging of sawblades. Thewood is slightly sticky when freshly sawn dueto the latex. The peeling characteristics aregood. Nailing and screwing usually do notcause problems, although the nail- and screw-holding capacity may be poor, and glueing,painting and varnishing give good results. Thesurfaces produced are generally smooth ifsharp cutting edges are used. The steam-bending properties are poor.The wood is not durable. It is easily attackedby fungi and very prone to blue stain. It is sus-ceptible to dry-wood borers, powder-post bee-tles and termites. Consequently, it should betreated as soon as possible after felling. It isonly suited for indoor use, and cannot be usedin contact with the ground or exposed to theweather. However, it is easy to treat with pre-servatives, absorbing over 480 kg/ina usingopen tank or pressure systemsIn tests with mice and rats anti-inflammatory,antipyretic and analgesic properties of AIStoniobooneistem bark have been demonstrated, butit is not efficient in treating malaria. Severalalkaloids have been isolated from the bark.

One of these is the indole alkaloid echitamine,which showed several pharmacological activi-ties (e. g. hypotensive activity and relaxing ac-tivity of smooth muscles) in animal test inod-els. However, this compound showed only verylimited antiplasmodialeffect. The triterpenoidsIupeol and eramyrin, as well as esters of thesecompounds have been isolated from stem androot bark; these showed anti-inflammatoryproperties. Alcoholic extracts of the bark (andto a lesser degree of the leafy showed a broad-spectrum activity against both gram-negativeand grain-POSitlve bacteria, as well as fungiThe latex may cause blindness


of the closely related AIStonio congensis Englis used indiscriminately from that of AIStonioboonei. The wood of both species has similar

to that of Triplochiton seieroxylonuses

K. Schum. and Antioris tonedrid Leseh. In

drugstores AIStoniu boonei has been used toadulterate HOJOrrheno lionbundo (G. Don)T. Durand & Schinz and its latex has been

mixed with that of Funtumio elostico (Preuss)Stapfand other latex-producing species.

Description Large deciduous tree up to 45

ALSTONiA 65

66 TIMBERSl

5

.

\

4

thers up to 1.5 min long; ovary superior, ovoid,pubescent, 2-celled, style slender, 3-8 mm long,ending in a pistil head composed of a basalring, a cylindrical central part and an elongatecleft stigmoid apical part. Fruit composed of 2tinear foilicles 22-57 cm long and 2-4 mm indiameter, densely pubescent, many-seededSeeds oblong, flattened, 4.5-6 min x 1.5-2 mm,brown, with long hairs up to 2 cm long on theends. Seedling with epigeal germination; hypo-coty1 2.5-3.5 cm long, epicoty1 I-2 cm long;cotyledons leafy, persistent for 2-3 months,shortly stalked; first leaves opposite

Other botanical information AIStonio coin-

prises about 40 species and has a pantropicaldistribution. Only two species are indigenousin Africa, both belonging to section AIStonioAIStonio congensis Engl. differs from AIStonioboonei in its usually sessile leaves, glabroussepals and glabrous follicles, and also in sur-face sculpturing of the lower leaf epidermis(stomata concealed below wax chains in AISto-itto boonei, exposed in AIStonio congensis). Inmuch of the older literature concerning WestAfrica, the name AIStonio congensis has beenerroneously used for AIStonio booneiAIStonio ingcrophyllo Wall. ex G. Don is occa-SIonally planted as an ornamental in tropicalAfrica, e. g. in Sierra Leone, C6te d'Ivoire, All-gola and Zimbabwe. It originates from tropicalAsia, where is it a fairly important timber treeand medicinal plant. Its wood is heavier,stronger and harder than that of AIStortiobooneiandAlstonio congensis.

Anatomy Wood-anatointcal description (IAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13:simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); 25: intervessel pits small (4-7 prn); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;46: s 5 vessels per square minimetre; 47: 5-20vessels per square milllmetre. Tracheids andfibres: (61: fibres with simple to minutely bor-dered pits); (62: fibres with distinctly borderedpits); 66: non-septate fibres present; 68: fibresvery thin-walled; (69: fibres thin- to thick-walled). Axial parenchyma: 78: axial paren-chyma scanty paratrachea1; 86: axial paren-chyma in narrow bands or lines up to threeceUs wide; 87: axial parenchyma reticulate; 92

3

,2

AIStonio boonei - I, buse of 601e, . 2, port of670nch with ledues, . 3, inflorescence, 4, flower, 5,fruit, . 6, seed.Redrawn und adopted by Achingd Satin Nur-hornon

in tall, with latex; hole branchless for up to 25in, up to 100(-140) cm in diameter, fluted atthe base or with steep buttresses up to 4(-8) inhigh; bark smooth or rough, usually with largelenticels, greyish or pale brown, Tnner barkcreamy or pale yellow, wLth copious latex;crown layered, with branches in whorls. Leavesin whorls of 4-8, simple and entire; stipulesabsent; petiole up to 2.5 cm long, flattened andgrooved above, with narrowly triangular col-leters at base; blade obovate, 6-20(-25) cm x2.5-7.5 cm, curieate to obtuse at base, roundedto shortly acuminate at apex, leathery, gla-brous, many-veined with lateral veins Joiningnear the margin. Inflorescence a terminal,compound, umbel-like cyme up to 23 cm long;peduncle 2-6(-13) cm long, stout; bracts up to2.5 mm long. Flowers bisexual, regular, 5-meTous, fragrant; pedice1 2-5(-7) min long;sepals up to 2(-2.5) mm long, fused at base,pubescent; corolla cream-coloured or pale yel-low, pubescent, with tube 5-14 mm long, nar-rowed around the middle, lobes obliquely ovate,2.5-6 min long; stamens inserted in upper partof corolla tube, filaments up to I mm long, an-

a

,

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I II j. :,'*,, $4*,

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four (3-4) cells per parenchyma strand; 93:eight (5-8) cells per parenchyma strand. Rays:97: ray width I-3 cells; 106: body ray cells pro-cumbent with one row of upright and/or squaremarginal cells; 107: body ray cells procumbentwith mostly 2-4 rows of upright and/or squaremarginal cells; 1/5: 4-12 rays per min. Secre-tory elements and cambial variants: 132: Iati-cifers or tanniferous tubes. Mineral inclusions

(136: prismatic crystals present); (137: pris-matic crystals in upright and/or square raycells); (141: prismatic crystals inchambered axial parenchyma cells); (154: morethan one crystal of about the same size per cellor chamber)(E. Ebanyenle, A. A. Oteng-Amoako & P. Baas)

Growth and development Saplings mayreach 2 in tall in the first year, and 10-year-oldtrees of 15 in tall and 25 cm in bole diameterhave been recorded in Ghana. The tree is de-

ciduous at the end of the rainy season. In WestAfrica trees flower from November to Januaryand fruit from January to February. The flow-ers are pollinated by Insects, especially byhoney bees, but also by butterflies, wasps andbeetles. The seeds are dispersed by windAIStonio booneiforms endomycorrhizae

Ecology AIStonio boonet occurs in prlmaryas well as secondary, moist evergreen to drysemi-deciduous forest up to 1200 in altitude. Itis in general a light demander, but toleratesshade when young. Seedlings are most commonin medium-sized to large forest gaps and aban-doned plantations, and are not found in deepshade. AIStonio booneiis less abundant in re-

gions with high rainfall. It is sometimes foundin swampy locations and along streams, butusually occurs in drier habitats than AIStoniocongensis. A comparatively high density ofAIStonio boonei trees indicates former distur-

bance of the forest cover by humans. AIStontoboonei regenerates well in forest gaps andabandoned cultivation areas, but in closed for-est and older secondary forest only larger treesare found.

Propagation and planting Germination ofseed takes 18-25 days. Immersion in water for24 hours or manual scarification enhance ger-mination from about 30% to 85%, whereastreatments with sulphuric acid and boilingwater destroy the seeds. There are about33,000 seeds per kg.

Management The trees coppicereadily. Theyare often damaged by strong winds and breakat the levelofthe branch whorls

Harvesting Logs frequently have brittle

heart

Handling after harvest The logs float eas-ily in water, and can be transported by riverThey are very susceptible to insect and fungalattack and should be dried quickly or treatedwith preservatives after felling. They are notwell suited for rotary peeling because they areoften deeply fluted. The return after sawing ispoor, being reduced by the latex canals in thewood and by the nodes of whorled branches.

Genetic resources AIStonio boonetis wide-

spread and in many regions common, also insecondary forest, and does not seem to be eas-11y liable to genetic erosion. However, cuttingAIStonio boonei trees for timber and collection

of the bark for medicinal purposes may causeserious threats to local populations. In the Djabiosphere reserve in Gamero0n, for instance,AIStonio booneiis endangered because its barkis much collected to treat malaria, and this isalso the case locally in Ghana. The problem isaggravated by the fact that AIStonio booneibark recovers poorly after damage by exploita-tion. In Senegal AIStonio booneiis completelyprotected

Prospects AIStonio boonei has potential toserve as a substitute of Triplochiton seieroxy!onK. Schum. , which is an important low-weighttimber species in West Africa. However, exportprospects are doubtful because of the poor du-rability and limited uses of the wood. It may beuseful as an auxiliary plant in agriculture be-cause it is easy to propagate, grows moderatelyfast and develops a taproot which does not in-terfere with the superficial root system of thecrop

The effectiveness of AIStonio boonei and other

AIStonio species in the treatment of malaria Iscontroversial, and more research is stillneededto confirm the claimed medicinal activities

Major references Burki11, 1985; Chudnoff,1980; CTRAD Forestry Department, 2003; Sidi-yasa, 1998; SIepel, Poorter & Hawthorne, 2004;Voorhoeve, 1979; World Agroforestry Centre,undated.

Other references Abu, 1992; Burki11, 2000;de Jong, 1979; InsideWood, undated; Jayeola,1998; Mapongmetsem at a1. , 1998-1999; 0je-wole, 1984; 01ajide at a1. , 2000; Railc at al. ,2000; Rudjiman at a1. , 1993; Sam, 1993; Taka-hashi, 1978; Te0, 2001; Vivien & Faure, 1985;Wilks & Issemb6, 2000

Sources of illustration de Jong, 1979;Wilks & Issemb6, 2000.

Authors F. Falla

non-

ALSTONIA 67

11

68 TIMBERSl

ALSTONiACONGENSIS Engl.

Protologue Bot. Jahrb. Syst. a 64 (1887)Family ApocynaceaeSynonyms AIStonio gilletii De Wild. (1907).Vernacular names instonia, cheesewood,

stool wood, pattern wood (En). Emien (Fr)Songati(Po)

Origin and geographic distribution AISto-itto congensts occurs from south-western Nige-ria to the Central African Republic, easternand southern DR Congo, and northernAiigola.

Uses The wood is used for light construction,light carpentry, open boats, moulding, furni-ture, interior joinery, implements, boxes,crates, matches, pencils, sculptures (e. g.masks), and for veneer and plywood (interior).It is locally popular for the production ofhousehold implements because of its goodworking properties and stabilityA bark decoction is used to treat malaria, gon-orrhoea, diarrhoea and other intestinal prob-Iems, rheumatic pain, and as a galactagogue,and the bark is also applied as an antidote forarrow poison and as an arithelmintic. The latexis used to treat leucorrhoea, ulcers, scabies,yaws and headache. Lightly roasted leaves aresmoked in a pipe as a remedy for cough. Thetree is occasionally used as an ornamental orshade tree

Production and international trade A1.

though AIStonio timber of Asian origin (fromdifferent species; mainly from Indonesia andMalaysia) is important on the internationaltimber market, AIStonio timber is only occa-SLonally exported from African countries, andthen probably in mixed consignments of light-weight timber. It is mostly used locally.

Properties In trade the wood of AIStoniocongensis is not distinguished from that ofAIStonio boonei De Wild. ; it has similar proper-ties. The heartwood is creamy white and iridis-tinctly demarcated from the up to 20 cm widesapwood. The grain is straight, occasionallywavy, texture moderately coarse. Growth ringsindistinct or absent.

The wood is lightweight. The density is 340-400kg/ina at 12% moisture content. The shrinkagerates are moderately low, from green to ovendry 3.2-4.0% radial and 5.1-5.5% tangentialThe wood dries rapidly, with slight risk of dis-tortion and checking. Once dry, it is stableAt 12% moisture content, the modulus of rup-ture is 48-70 N/mm2, modulus of elasticity5800-8800 N/min2, compression paraUelto grain23-36 N/min2, shear I-2 N/min2, cleavage 7-16N/mm and Janka side hardness 1825 N

The wood is easy to saw, although the presenceof latex may cause clogging of sawblades. Thepeeling characteristics are good. Nailing andscrewing usually do not cause problems, andgluing, painting and varnishing give good re-SUIts. The surfaces produced are generallysmooth if sharp cutting edges are used. Thesteam-bending properties are poorThe wood is not durable. It is easily attackedby fungi and very prone to blue stain. It is sus-ceptible to dry-wood borers, powder-post bee-tles and termites. Consequently, it is onlysuited for indoor use and cannot be used in

contact with the ground. However, it is easy totreat wlth preservatives, absorbing over 480kg/ms using open tank or pressure systems.A methanolic leafextract showed some antima-

Ianal activity in tests on Piersmodium bergheiin mice; It suppressed early Infection, butshowed no effect when the infection was al-

ready established. The bark and leaves showedcardio-activity in animal tests. Several alka-10ids have been isolated from the bark andleaves. One of these is the indole alkaloid echi-

tamine, which showed several pharmacologicalactivities (e. g. hypotensive activity and relax-ing activity of smooth muscles) in animal testmodels. However, this compound showed onlyvery limited antiplasmodialeffect


of the closely related AIStonio booneiis used in-discriminateIy from that of AIStonio congensis.

Description Small to medium-sized tree upto 15(-25) in tall, with latex; hole up to 30(-100) cm in diameter, fluted at the base or withsteep buttresses; bark smooth or rough, usuallywith large lenticels, greyish or pale brown,

.

$3-F

AIStonio congensis - wild

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3

and an elongate cleft stigmoid apical partFruit composed of 2 linear follicles 20-45 cmlong and 2-4 min in diameter, glabrous, many-seeded. Seeds oblong, flattened, 6-9 min x 2-2.5 mm, brown, with long hairs up to 2 cm longon the ends

Other botanical information AIStonio

comprises about 40 species and has a pantropi-cal distribution. Only two species are indige-nous in Africa, both belonging to section AISto-itto. AIStonio boonei differs from AIStonio con-

gensis in its stalked leaves, pubescent sepalsand pubescent follicles, and also in surfacesculpturing of the lower leaf epidermis (sto-mata exposed in AIStonio congensis, concealedbelow wax chains in AIStonio boonei). In muchof the older literature concerning West Africa,the name AIStonio congensis has been errone-ously used for AIStonio boonei.

anatomy Wood-anatointcal description OAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; (10:vessels in radial multiples of 4 or more coin-mon); 13: simple perforation plates; 22tervessel pits alternate; (23: shape of alternatepits polygonal); (24: intervesselpits minute (s 4prn)); 25: intervessel pits small (4-7 pin); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;(43: mean tangential diameter of vessellumina;a 200 prn); 46: s 5 vessels per square minime-tre; (47: 5-20 vessels per square minimetre).Tracheids and fibres: (61: fibres with simple tominutely bordered pits); (62: fibres with dis-tinctly bordered pits); 66: non-septate fibrespresent; 68: fibres very thin-walled. Axial pa-Tenchyma: 78: axial parenchyma scanty para-tracheal; 86: axial parenchyma in narrowbands or lines up to three cells wide; (87: axialparenchyma reticulate); (92: four (3-4) cells perparenchyma strand); 93: eight (5-8) cells perparenchyma strand; (94: over eight cells perparenchyma strand). Rays: 97: ray width I-3cells; 106: body ray cells procumbent with onerow of upright and/or square marginal cells;107: body ray cells procumbent with mostly 2-4rows of upright and/or square marginal cells;115: 4-12 rays per min. Secretory elements andcambial variants: 132: Iaticifers or tanniferous

tubes. Mineral inclusions: (136: prismatic crys-tals present); (137: prismatic crystals in up-right and/or square ray cells); (141: prismatic

V

AIStonio congensis - I, port of bronch withIeoues, . 2, inflorescence, . 3, flower, . 4, fruit, ' 5,seed.

Redrown und oddpted by Achingd Sotiri Nur-hornon

4

2

,,

\\

ALSTONiA 69

inner bark creamy or pale yellow, with copiouslatex; crown layered. Leaves in whorls of 4-6(-8), simple and entire; stipules absent; petioleup to 0.5(-I) cm long, flattened and groovedabove, with narrowly triangular coneters atbase; blade obovate to spatulate, 8-26 cm x3.5-11.5 cm, curieate at base, rounded toshortly acuminate at apex, occasionally Tetuse,leathery, glabrous, many-veined with lateralveins joining near the margin. Inflorescence aterminal, compound, umbel-like cyme up to 25cm long; peduncle 3.5-12 cm long, stout; bractsup to 2 min long. Flowers bisexual, regular, 5-merous, fragrant; pedice1(2-)3-5(-9) min long;sepals up to 3 mm long, fused at base, glabrousbut slightly pubescent at margins; corollacream-coloured, pale yellow or pale pink, pu-bescent, with tube 4-7.5 mm long, wideningnear the end, lobes obliquely obovate, 4-10 mmlong; stamens Inserted in upper part of corollatube, filaments c. 05 mm long, arithers up to1.5 mm long; ovary superior, ovoid, glabrous oroccasionally slightly pubescent, 2-celled, styleI-3.5 mm long, ending in a pistil head coin-posed of a basal ring, a cylindrical central part

5

In-

.

70 TIMBERSl

crystals in non-chambered axial parenchymacells); (154: more than one crystal of about thesame size per cellor chamber)(E. Ebanyenle, A. A. Oteng-Amoako & P. Baas)

Growth and development The seeds aredispersed by wind

EcologyAlstonio congensis usuaUy occursinswampy localities (e. g. Rophio swamps) up to500 in altitude, but it is sometimes also foundon drier, rocky soils. In Equatorial Guinea it ismost common in a zone near the coast, with onaverage almost 2 trees over 30 cm in hole di-ameter per ha

Propagation and planting Propagation ofAIStonio congensis by stem cuttings usinggrowth regulators was successful(up to 63%rooting) in Nigeria

Handling after harvest The logs float eas-11y in water, and can be transported by rivers.They are very susceptible to insect and fungalattack and should be dried quickly or treatedwith preservatives after felling. They are notwell suited for rotary peeling because they areoften deeply fluted

Genetic resources AISton!a congensis Is

considered locally endangered, e. g. in Nigeria,where the bark is commonly collected for me-dicinalpurposes

Prospects Export prospects are poor for A1-stonto congensis because of the poor durabilityand limited uses of the wood and the usuallysmall size of the tree. It may be useful as anauxiliary plant in agriculture because it is easyto propagate, grows moderately fast and devel-ops a taproot, which does notinterfere with thesuperficial root system of cropsThe effectiveness of AIStonio andcongensis

other AIStorrio species in the treatment of ina-Iana Is controversial, and more research is stillneeded to confirm the claimed medicinal activi-

ties

Major references Burki11, 1985; Chudnoff,1980; CIRAD Forestry Department, 2003;Neuwinger, 2000; SIdiyasa, 1998; World Agro-forestry Centre, undated.

Other references Awe & Foeke, 1990; deJong, 1979; InsideWood, undated; Ilesanmi,A1adesanmi & Adeoye, 1988; Jayeola, 1998;Pauwels, 1993; Raponda-Walker & Sillans,1961; Rudjiman at a1. , 1993; Takahashi, 1978;Te0, 2001

Sources of illustration de Jong, 1979.Authors R. H. M. J. Lemmens

AMBLYGONOCARPUSANDONGENSIS(WeIw.ex 011v. ) Exell & Torre

Protologue Bol. SOC. BTOt. , ser. 2, 29: 42(1955)



Synonyms Am61ygonocorpus o6tusongu!us(WeIw. ex 011v. ) Harms (1899), Am 61ygonocor-pus schweinfurthiiHarms (1899).

Origin and geographic distribution Am 61y-gonocurpus gridongensis occurs in the savannazone from northern Ghana east to Sudan,through Uganda and Tanzania, south to Bot-swana, Zimbabwe and Mozambique.

Uses The wood, known in southern Africa as'bangawanga', is used for joinery and furniture.It is considered excellent for heavy duty floor-ing and for railway sleepers, and is also suit-able for construction, mine props, vehicle bod-Ies, sporting goods, agricultural implements,poles and piles, and vats. The wood is also usedas firewood and for charcoal production.Various plant parts are used in local medicinea root decoction is administered as an emetic to

treat food poisoning, against colic and coughand as a vermifuge, the bark is used as an an-tidote for snakebites, a bark decoction is ap-plied to sores, a leaf extract Is used to treatstomach-ache, and pulverized pods are appliedto ulcers and also used as a fish poison

Roasted seeds are eaten. Boiled and fermented

seeds are used as a condiment, e. g. in Camer-o0n, although they are hard to prepare andused in smallamounts.


wood of Am 61ygonoco, pus gridongensisis mainlylocally used; small quantities of sawn woodhave been exported from Mozambique

Properties The heartwood is dark brown orred-brown, darkening on exposure, and dis-tinctly demarcated from the narrow grey-whitesapwood. The grain is wavy or straight, some-times slightly interlocked, texture fine andeven.

The wood has a density of 910-1090 kg/in3 at15% moisture content. Shrinkage rates are2.3% radial and 2.7% tangential from green to12% moisture content. The wood air-dries

slowly, with some surface checking, but it isstable in service. The wood is difficult to saw

and work; blunting of cutting edges is commonIt can be finished to an excellent surface, isresistant to abrasion, and has good gluingproperties. Pre-boring is necessary for screws

^:

and nails. The wood is durable and termite-

resistant; it is resistant to impregnation withpreservatives. Charcoal made from the wood isconsidered excellent for iron-forge workThe seeds contain about 12% oil, with a highproportion of linoleic acid. The inner bark,roots and seeds have been recorded to be poi-sonous, but are used in various medicinal orfood preparations.

Botany Small to medium-sized tree up to20(-25) in tall, glabrous; hole straight, branch-less for up to 10 in, up to 90 cm in diameter,without buttresses; bark scaly, greyish brownto blackish. Leaves alternate, hipinnately coin-pound, with 2-5(-6) pairs of pinnae; petiole 4--9 cm long, rachis 2-18 cm long; leaflets alter-nate, shortly stalked, 11-21 per pinna, ellipti-cal to obovate-elliptical, I-3 cm x 0.5-2 cm,usually notched at apex. Inflorescence an axil-Iary raceme (3-)6-18 cm long; peduncle I-4.5cm long. Flowers bisexual, regular, 5-merous,white; pedice1 1.5-3.5(-5) min long; calyx 0.5-Imm long, toothed; petals free, elliptical, 3-5min x I-1.5 mm; stamens 10, with filaments 5-6 min long; ovary superior, oblong, shortlystalked, I-celled, style slender, curved nearapex. Fruit an oblong indehiscent pod 8-17(-20) cm x 2-3.5 cm, usually bluntly tetragonalin section, woody, brown, glossy, septate be-tween the seeds, c. 10-seeded. Seeds 10-13 minx 7-8 min, slightly flattened, hard, brownAm 61ygonocorpus comprises a single species. Itresembles Tetropleuro, which differs in its podvalves that are only thickened in a band and inits glandular arithers

Ecology Am61ygonocorpus undongensis oc-curs in deciduous woodland and savanna up to1350 in altitude.

Genetic resources and breeding Am61y-80nocorpus Gridongensis is widespread andlocally common. There Is no reason to considerit endangered

Prospects Am 61ygonocorpus gridongensis isa multipurpose tree with favourable wood andcharcoal qualities. Its applications as food andin traditional medicine merit more research

Major references Bolza & Keating, 1972;Brenan, 1970; Burki11, 1995; Chevassus & Pas-caud, 1972.

Other references Aubr6ville, 1950; Brenan,1959; Bryce, 1967; CTFT, 1962b; Fanshawe,1962; Neuwinger, 2000; Oxford Forestry Insti-tute, 1997-2004; Tanzania Forest Division,1967; Vivien, 1990a; Zambia Forest Depart-merit, 1979a.


AMPHIMAS FERRUGINEUS Pierre ex Fellegr

Protologue Notul. Syst. (Paris) 2: 293 (1912)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Synonyms Amphimos Momeonus Pierre ex

Penegr. (1912)Origin and geographic distribution Amphi-

inOS Ierrugineus occurs in south-western Cam-eroon, Equatorial Guinea, Gabon, Congo, west-ern DR Congo and northern Angola.

Uses The wood (trade names: Iati, bokanga)is used for interior construction, carpentry,flooring, planks, Interior trim, joinery, furni-ture, frames, crates, boxes, toys, novelties, ve-neer and plywood. A bark decoction is used intraditional medicine to treat dysmenorrhoea,blennorrhoea and as poison antidote: bark pulpIs applied to mumps

Production and international trade Am-

phimos Ierrugineus and Amphimus pterocor-potdes Harms are both traded as Iati. In 2003Cameroon exported about 70 ina of Iatilogs and130 ina of sawnwood. In 2004 Gamero0n ex-

ported about 2500 ina of Iatilogs and 110 in3 ofsawnwood

Properties The heartwood is yellowishbrown, often with whitish markings thatdarken with age, and more or less distinctlydemarcated from the 5-8 cm thick, yellow-white to pale brown sapwood. The grain isusually straight, sometimes wavy, texturemoderately coarse. The wood has a coarse SII-ver figureThe wood is moderately heavy. At 12% ino1s-ture content, the density is 690-750 kg/ina. Therates of shrinkage during drying are moder-ately high, from green to oven dry 3.4% radialand 8.5% tangential. The wood dries slowly,with severe risk of deformation. It is recoin-

mended to quarter-saw the timber before dry-ing. Some pre-drying is recommended beforekiln drying. After drying, the wood is moder-ately stable in service.At 12% moisture content, the modulus of rup-ture is 133-182 N/min2, modulus of elasticity11,300-13,600 N/mm2, compression parallel tograin 55-67 N/mm2, cleavage 21-30 N/min,and Chalais-Meudon side hardness 3.9-7.2

The wood saws and works well with standard

equipment, and can be planed to a smooth sur-face, but with some dulling effect on cuttingedges. The nailing and screwing properties aregood, with satisfactory holding properties, butpre-boring is often needed. The gluing, stainingand painting properties are satisfactory. The

AMPHiMAS 71

72 TIMBERSl

wood is suitable for sliced veneer and plywoodproduction. It is moderately durable to non-durable;it is susceptible to dry-wood borer andmarine borer attack, but often moderately re-sistant to fungal and termite attacks. Theheartwood is moderately resistant to impreg-nation by preservatives, the sapwood is perme-able.

Botany Large deciduous tree up to 45(-50)in tall; hole straight, cylindrical, branchless forup to 25(-30) in, up to 120(-150) cm in diame-ter, with thick buttresses; bark surface smoothto slightly rough, finely fissured or with rec-tangular scales, grey to greyish brown, innerbark thick, soft, fibrous, creamy to yellowish,exuding a red exudate; crown hemispherical;young twigs densely reddish brown hairyLeaves arranged spiraUy in tufts at the ends ofbranches, imparipinnately compound with 11-19 leaflets; stipules leafy, up to 2.5 cm long,caducous; petiole 2.5-4.5 cm long, rachis up to30 cm long but sometimes longer, denselyhairy; leaflets alternate to opposite, withthread-like stipels at base of 2-3 mm longpetiolules, (3-)6-18 cm x (1.5-)2-6 cm, ovate tooblong or elliptical, rounded to shortly acumi-nate at apex, densely short-hairy below, pin-nately veined with veins distinct below. Inflo-rescence a lax terminal or axillary compoundraceme c. 20 cm long, strongly branched, many-flowered. Flowers bisexual or male, regular, 5-merous, fragrant; pedicel c. I mm long; calyxcampanulate, c. 3 min long, with short lobes,densely brown hairy; petals free, equal, c. 5min long, deeply 2-lobed, whitish; stamens 10,fused at base, c. 4 min long; ovary superior,stipitate, densely to sparsely hairy, style SIen-der, c. 3.5 mm long. Fruit a pendulous, flat,elliptical to oblong pod, 15-22 cm x 5-7 cm,with large papery wing all around, reticulatelyveined, golden brown, indehiscent or slowlydehiscing with 2 valves, I(-2)-seeded. Seedkidney-shaped, 2-2.5(-3) cm long, brown.The winged fruits of Amphimos forrugineus arespread by wind.Amphimos comprises 2 or 3 species, and isconfined to West and Central Africa. The affin-

ity of the genus is still uncertain. It is usuallyclassified within the Sophoreoe tribe of Popil-ionoceoe, but has also been classified in Cues-o1pinioceoe because of its corolla which lacksthe typical papllionaceous structure. Amphi-mus pterocorpoides differs from Amphimosforrugineus in its almost glabrous leaflets hav-Ing Inconspicuous velns

Ecology Amphimos Ierrugineus occurs in

dense evergreen forest up to 500 in altitudeManagement In Cameroon the minimum

diameter for felling is 50 cm. After felling, logsshould be removed rapidly from the forest be-cause they are liable to blue stain attack. Whenleft for longer periods, they should be treatedwith preservatives. Usually, logs float in waterand thus can be transported by river.

Genetic resources Amphimos Ierrugineusmay be threatened by genetic erosion becauseit has a rather limited area of distribution and

specific ecological requirements. It does notseem to occur in high densities, and is uricom-mon or absent in several regions, e. g. in thesedimentary basin of western Gabon

Prospects Amphtmos Ierrugineus occurs inan area where many timber species with betterwood are still available, and therefore it haspoor prospects as a timber tree of commercialimportance in the near future. However, it ispoorly known, and research may reveal unex-pected possibilities

Major references Bolza & Keating, 1972;CIRAD Forestry Department, 2003; Fouarge &G6rard, 1964; Takahashi, 1978; Vivien &Faure, 1985.

Other references ATIBT, 1986; ATIBT,2004; ATIBT, 2005; Aubr6ville, 1970; Bouquet,1969; de Saint-Aubin, 1963; Neuwinger, 2000;Normand & Paquis, 1976; Witczek at a1. , 1952;Wilks & Issemb6, 2000


AMPHIMASPTEROCARPOIDES Harms

Protologue Repert. Spec. Nov. Regni Veg.12: 12 (1913)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)


Origin and geographic distribution Amphi-ings pteroco, potdes is widespread from Guineaeast to DR Congo and Sudan, and south tonorthern Gabon

Uses The wood (trade names: Iati, yaya, bo-kanga) is used for interior construction, car-pentry, flooring, planks, interior trim, Joinery,furniture, frames, crates, boxes, toys, novelties,veneer and plywood; it is also locally used forcanoes, huts, mortars and as frames to supportyam. It has been used for railway sleepers andpoles, although use in contact with soilis dis-couraged.The reddish resin from the bark is used in tra-

ditional medicine to treat dysentery, anaemia,

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Amphimospterocorpoides- wild

haematuria, dysmenorrhoea, blennorrhoea,schistosomiasis and mumps, and as poisonantidote. The inner bark is administered to

treat cough and in a steam bath against yawsand sores on the feet. A decoction of twig barkis applied to prevent threatening abortion, anda wood decoction is drunk to treatimpotence.

Production and international trade Am-

phimos pterocorpoides and Amphimos Ierrugi-iteus Pierre ex Penegr. are both traded as Iati.In 2003 Garnero0n exported about 70 in3 of Iatilogs and 130 ina of sawnwood. In 2004 Gamer-o0n exported about 2500 ina of Iatilogs and 110ing of sawnwood. The wood of Amphimos ptero-coreoides can be found on a very smallscale asveneer in Europe, under the name 'whitewerige', but it is now mainly used locally inAfrica. In C6te d'TVoire it is sold for local utili-zation, but it is considered of lesser quality.

Properties The heartwood is yellowishbrown, often with whitish markings thatdarken with age, and more or less distinctlydemarcated from the 5-9 cm thick, yellow-white to pale brown sapwood. The grain Isusually straight, sometimes wavy, texture me-dium to coarse. The wood has a coarse silver

figure.The wood is moderately heavy. At 12% ino1s-ture content, the density is 670-880 kg/ing. Therates of shrinkage from green to 11% moisturecontent are 2.9% radial and 5.6% tangential,and from green to oven dry 5.4-6.4% radial and9.7-10.8% tangential. The wood dries slowly,with severe risk of deformation, checking orcase hardening. It is recommended to quarter-saw the timber before drying. Some pre-dryingis recommended before kiln drying. After dry-

ing, the wood is moderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 80-129 N/min2 modulus of elasticity11,600-16,300 N/mm2, compression parallel tograin 50-64 N/min2, shear 16 N/mm2, cleavage12-16 N/min, Janka side hardness 5800 N andJanka end hardness 6300 N

The wood normally saws and works well withstandard equipment, but stellite-tipped saw-teeth are recommended for sawing large logs.It can be planed to a smooth surface, but withsome dulling effect on cutting edges. The nail-ing and screwing properties are good, withsatisfactory holding properties, but pre-boringis often needed. The gluing properties are sat-ismctory, and painting and varnishing areeasy. The wood is suitable for sliced veneer andplywood production. It is moderately durable tonon-durable; it is susceptible to dry-wood borerand Lyctus beetle attack, but often moderatelyresistant to fungal and termite attacks. Theheartwood is poorly to moderately permeable topreservative treatment, the sapwood is perme-able. The pulping characteristics for paper pro-duction are rather poor and the wood pulp canonly be used in combination with better pulpThe presence of alkaloids has been demon-strated for the bark and pods; the bark is poi-sonous to mice and fish. The heartwood con-tains the isoflavone afromosin.


of Amphimos ferrugineusis very similar to thatof Amphimos pterocorpoides and mixed intrade in regions of Central Africa where bothspecies occur, e. g. in Cameroon. In Ghana Am-phimos pterocorpoides wood is considered asuitable substitute for the wood of Miltcio ex-eelso ayelw. ) CG. Berg. It has some resem-blance to the wood ofSterculiu oblongo Mast.

Description Large deciduoustree up to 45(-50) in tall; bole straight, cylindrical, branchlessfor up to 25(-30) in, up to 130(-180) cm in di-ameter, with thick buttresses; bark surfacerough, with rectangular scales, greyish brownto dark brown, inner bark thick, soft, fibrous,orange to brown, exuding a red exudate; crownhemispherical; young twigs densely reddishbrown hairy. Leaves arranged spiralIy in tuftsat the ends of branches, Imparipinnately coin-pound with 11-21 leaflets; stipules leafy, up to2.5 cm long, caducous; petiole 2.5-4.5 cm long,rachis up to 30 cm long but sometimes longer,densely hairy; leaflets alternate to opposite,with thread-like stipels at base of 2-5 mm longpetiolules, (2-)5-13 cm x (1.5-)2.5-5 cm, ovateto oblong or erriptical, rounded to shortly acu-

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74 TIMBERSl

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genus is still uncertain. It is usually classifiedwithin the Sophoreoe tribe of Poptlionoceoe,but has also been classified in Goesolpinioceoebecause of its corolla that lacks the typicalpapilionaceous structure. Amphimos Ierrugi-iteus differs from Amphimos pteroco, :potdes inmore hairy leaflets having more conspicuousveins. Amphimus tessmonnii Harms has beendescribed on the basis of material from Equa-tonal Guinea;its status is uncertain.

Anatomy Wood-anatomical description CAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 27: intervesselpits large G 10 prn); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 F1m;43: mean tangential diameter of vessellumina2 200 prn; (45: vessels of two distinct diameterclasses, wood not ring-porous); 46: s 5 vesselsper square minimetre. Tracheids and fibres:61: fibres with simple to minutely borderedpits; 66: non-septate fibres present; (69: fibresthin- to thick-walled); 70: fibres very thick-walled. Axial parenchyma: 83: axial paren-chyma confluent; 85: axial parenchyma bandsmore than three cells wide; 92: four (3-4) cellsper parenchyma strand. Rays: 98: larger rayscommonly 4- to 10-senate; 104: all ray cellsprocumbent; 1/5: 4-12 rays per min. Stonedstructure: 1/8: all rays stoned; (1/9: low raysstoned, high rays non-stoned); 120: axial pa-renchyma and/or vessel elements stoned; 122:rays and/or axial elements irregularly stonedMineral inclusions: 136: prismatic crystalspresent; 142: prismatic crystals in chamberedaxial parenchyma cellsre. Derienne & P. E. Gasson)

Growth and development Natural regen-eration is generally abundant. Amphimospterocorpoides is classified as a non-pioneerlight demander, and regeneration is common insmall gaps in the forest. Seedlings toleratesome shade, but saplings require more light forproper growth, developing straight and un-branched stems untilthey are at least 70 cmtall. In the semi-deciduous forest zone in C6te

d'Ivoire, 14-year-old Amphimos preroco, potdestrees planted in full sun have a mean hole di-ameter of 19.6 cm, but the diameter growth innatural forest is only 0.2-0.4 cm per yearTrees usually flower when they are leaness in

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minate at apex, sometimes srightly notched,glabrous except inIdrib below, pinnately veinedwith rather indistinct veins below. Innores-

cence a lax terminal or axiUary compound ra-ceme c. 20 cm long, strongly branched, many-flowered. Flowers bisexual, regular, 5-merous,fragrant; pedicelc. I mm long; calyx campanu-late, c. 2 mm long, with short lobes, denselybrown hairy; petals free, equal, c. 6 mm long,deeply 2-10bed, whitish; stamens 10, fused atbase, c. 5 min long; ovary superior, stipitate,glabrous, style slender, c. 3.5 min long. Fruit apendulous, flat, elliptical to oblong pod, 8-22cm x 3-7 cm, with large papery wing allaround, reticulately veined, golden brown, in-dehiscent or slowly dehiscing with 2 valves, I(-2)-seeded. Seed kidney-shaped, 1.5-2(-3) cmlong, brown. Seedling with epigeal germina-tion; hypocoty1 3-4 cm long, epicoty1 5-6 cmlong; cotyledons thick and fleshy, 1.5-2 cmlong; first two leaves opposite, simple andstipulate

Other botanical information Amphimoscomprises 2 or 3 species, and is confined toWest and Central Africa. The affinity of the

4

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I

the dry season, in Sierra Leone, C6te d'Ivoireand Ghana from October to December. Fruits

take about 3 months to ripen. The wingedfruits are spread by wind. The roots have nod-ules containing nitrogen-fixing bacteria.

Ecology Amphimos prerocorpoides occurs indifferent types of forest, from dense evergreenforest to deciduous forest, sometimes also insecondary forest and bush fallow. It may be-come common in agricultural land as the treesare difficult to fell. In Ghana it is most commonin moist evergreen forest, but is also fairlycommon in moist semi-deciduous forest. It pre-fors high rainfall areas, and its abundancedecreases with decreasing soil fertility. Am-phimos pterocorpoides is poorly resistant toforest fires.

Propagation and planting The 1000-seedweight of Amphimos pterocorpoides is about800 g. Seeds germinate within 5-15 days, butseedlings stop growth for a few weeks whenthey have developed the first 2 leaves.

Management In Liberia, where Amphimospterocorpoides is rather frequent, the averagestock (more than 50 cm hole diameter) is esti-mated at 43 in31km2 and the total volume ofsuch trees at 1,700,000 ina. In Sierra LeoneAmphimos pterocorpoides is considered a weedin production forest.

Diseases and pests In Liberia and SierraLeone ambrosia beetles often attack the livingtree and create holes in the wood

Harvesting The minimum diameter for fell-ing is 50 cm in Liberia, 60 cm in Ghana and 70cm in C6te d'Ivoire

Handling after harvest After felling, logsshould be removed rapidly from the forest be-cause they are liable to blue stain attack. Whenleft for longer periods, they should be treatedwith preservatives. Usually, logs float in waterand thus can be transported by river.

Genetic resources There are no indicationsthat Amphimospterocorpoides is threatened bygenetic erosion. It is not only widespread, butalso locally common with abundant regenera-tion. Moreover, it has a wide ecological ampli-tude

Prospects Until recently the wood of Am-phimospterocorpoides was considered of ratherpoor quality, but with the depletion of manycommercial timber species it is becoming ofmore interest, especially for local constructionand joinery, but presumably also for export inthe future. However, much research is stillneeded on growth rates, ecological require-merits and management of naturalstands on a

sustainable basis.

Major references Ayarkwa, 1994; Bolza &Keating, 1972; Burki11, 1995; CIRAD ForestryDepartment, 2003; Dudek, F6rster & mis-senbauer, 1981; Richter & Danwitz, 2000;SIepel, Poorter & Hawthorne, 2004; Takahashi,1978; Vivien & Faure, 1985; Voorhoeve, 1979

Other references Adjanohoun at a1. , 1996;A1<6 Assi at a1. , 1985; Mobgninou, van derBurg & van der Maesen, 2006; ATIBT, 1986;ATIBT, 2004; ATIBT, 2005; Bouquet, 1969;Bouquet & Debray, 1974; de in Mensbruge,1966; Durrieu de Madron at a1. , 1998a; Fou-quet, 1984; G6rard at a1. , 1998; Hawthorne,1995; Hawthorne & Jongkind, 2006; Inside-Wood, undated; Neuwinger, 2000; Normand,1960; Normand & Paquis, 1976; SOSef at al. ,2006; Wilczek at a1. , 1952; Wilks & TSSemb6,2000

Sources of illustration Vivien & Eaure,1985; Wilczek at a1. , 1952

AuthorsA. T. Tchinda & P. Tan6

ANTIARiS 75

ANTIARISTOXICARIALesch.

Protologue min. Mus. Natl. Hist. Nat. 16478, PI. 22 (1810).

Family MoraceaeChromosome number2n = 24, 28Vernacular names Antiaris, bark cloth

tree, false IToko (En). Ako (Fr). P6 de bitcho, p6de leite (Po). Mkunde (Sw)

Origin and geographic distribution Antio-ris toxicorio is extremely widespread, beingfound throughout the Old World tropics. InAfrica it occurs from Senegal east to southernEthiopia, and south to Zambia and hagola; it is

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also found in Madagascar. It occurs in tropicalAsia, islands of the Pacific Ocean (east to Fijiand Tonga) and northern Australia.

Uses The wood is used for interior joinery,panelling, moulding, shuttering, furniture,strip flooring, boxes and crates, tool handles,toys, carvings, peeled and sliced veneer forinterior and exterior parts of plywood, fibreand particle board, and blockboard. It is fairlycommonly used domestically for light construc-tion and canoes. It is locally popular for drummaking, e. g. in Uganda. The wood from theroots is sometimes used as a cork substitute

The bark yields a latex which is one of theprinciple components of most dart and arrowpoisons in South-East Asia. In Africa the latexis applied to cuts, wounds and skin complaintssuch as eczema and leprosy, and is taken in-ternally as a purgative. It Is also reported to beused as a fish poison and birdlime. Seeds,leaves and bark are used as a febrifuge and theseeds also as an antidysenteric. The bark isused as an anodyne and vermifuge, and totreat hepatitis. It has also been used for dye-ing. The inner bark is used to make roughclothing, hammocks, sandals, hut walls, cord-age, sacks, mats and paper. The fruit is edible.The leaves are used as fodder. Antioris tort-

cor, o Is sometimes planted as a roadside treeProduction and international trade The

export of Antioris tonedrio timber from WestAfrica started in 1959, when 1600 ms of logswere exported. It increased to 40,000 ina in1963; in 1973 66,000 ina was exported fromC6te d'TVoire and in 1983 165,000 ina. After1983, the export decreased rapidly. At present,Antioris tonedrig Is most important on theinternational market for Its veneer and ply-wood. In 2001 Ghana exported 4000 ms of ve-neer at an average price of Us$ 5241m3, and in2002 it exported 2000 ina at an average price ofUs$ 4631m8; the export of plywood from Ghanawas 13,000 ms in 2001 and 9000 ing in 2002 atan average price of Us$ 2681m3. The wood ofAntioris tonedrig is often traded in mixed con-

signments of lightweight hardwoodProperties Heartwood whitish to pale yel-

low or pale yellow-brown, indistinctly demar-cated from the sapwood, which is up to 8 cmthick. The grain is interlocked, texture moder-ately coarse. The wood has a ribbon-like aspecton quarter-sawn faces, and is lustrous. Freshwood has woolly surfaces.The wood of Antioris toxicorio is a lightweighthardwood. The density is 370-480(-660) kg/in3at 12% moisture content. The shrinkage rates

are moderate, from green to 12% moisture con-tent 2.2% radial and 4.3% tangential, and fromgreen to oven dry 3.3-4.6% radial and 5.8-8.2%tangential. The timber dries moderately easilyand rapidly, with rather high risk of distortionfor flat-sawn boards and slight risk orcheckingOnce dry, the wood is moderately stable inservice.

At 12% moisture content, the modulus of rup-ture is 42-98 N/min2, modulus of elasticity5700-10,000 N/mm2, compression parallel tograin 28-53 N/min2, shear 3-9 N/mm2 cleav-age 10-17 N/min, Janka side hardness 1690-5610 N and Janka end hardness 2270-6630 N.

The wood works easily with hand and machinetools; ordinary saw teeth and cutting tools canbe used, but these should be kept sharp to pre-vent crumbling, particularly along the edges. Asmooth finish can be obtained, but with sometearing due to interlocked grain. The peelingand slicing properties are good, but rotary-peeled veneer is somewhat brittle. The woodstains and polishes well. Filling is recoin-mended to obtain a good finish. The nailingand screwing properties are satisfactory. Glu-ing does not cause problems.The wood is not durable. It is liable to fungalattack (e. g. blue stain), and susceptible to ter-mites and dry-wood borers; the sapwood is Ii-able to powder-post beetle attack. The woodshould not be used in contact with the groundor exposed to the weather. It is easy to treatwith preservatives using either open tank orpressure systems; a retention of 455 kg/ina hasbeen determined for heartwood and 540 kg/in3for sapwood. The sawdust may cause skin irri-tation and occupational asthmaThe latex is reported to be a mild circulatoryand cardiac stimulant when used in very smallamounts, but in large amounts it Is a myocar-dial poison. The active principles are cardiacglycosides (cardenolides), e. g. u-antiarin, p-antiarin and Tantiarin, which have digitalis-like effects on the heart. In larger amountsthey lead to cardiac arrest and secondary ef-fects such as vomiting and convulsions. Re-ports on lethal dosage, administered intrave-nously, specify 0.3 ing as lethal within 12 min-utes for a rabbit, and I ing within 3-9 minutesin dogs. Tests with animals suggest that thecardiac glycosides affect Na+K+ATPase activityof the heart muscle-cells. The poison must en-ter the bloodstream to be effective; the latexcan be ingested without any effects. Reportsclaim that the latex from African trees is less

poisonous or even innocuous. It is possible that

these reports refer to the latex being used dif-forently, not as a dart or arrow poison as inSouth-East Asia, and thus not entering thebloodstream. All aqueous ethanol extract of thebark exhibited cytotoxic activity against tu-mour celllines

Adulterations and substitutes The woodof Antioris toxicorio resembles that of Tr, -PIOchiton seieroxylon K. Sebum. and can beused as a substitute for the latter. It can alsobe used as a substitute for Terminalto superboEngl. & Diels and Pterygoto inocrocorpoK. Schum. wood.

Description Deciduous, monoecious, smalltolarge tree up to 45(-60) in tall; hole straight,branchless for up to 25(-33) in, up to 180 cm indiameter, sometimes with steep buttresses upto 3 in high; bark surface smooth becomingslightly fissured, greyish white to greyishgreen, with numerous lenticels, inner bark softand fibrous, exuding a creamy latex soon dark-ening to dirty brown; crown fairly small, dome-shaped; twigs hairy. Leaves alternate, more orless distichous, simple; stipules free, up to I(-1.5) cm long, caducous; petiole up to I cm long,hairy; blade elliptical to oblong or obovate, (4-)6-20(-30) cm x 3-12 cm, obtuse to slightly

cordate and slightly unequal at base, obtuse toshortly acuminate at apex, entire to slightlytoothed, thinly leathery to leathery, hairy, pin-nately veined with lateral veins in (5-)7-14pairs. Inflorescence on a short shoot in or belowthe leaf axils, I-8 together, male ones 0.5-I(-2) cm in diameter, many-flowered, female ones3-4 min in diameter, I-flowered, subtended byinvolucral bracts. Flowers unisexual; maleflowers with (2-)3-5(-7) free tepals and 2-4stamens; female flowers with 4-lobed perianthand I-celled ovary adnate to the perianth,styles 2, long. Fruit forming a drupe-like, ellip-sold to ovoid or globose entity together with theenlarged, fleshy orange to scarlet receptacle, I-1.5(-2) cm long, I-seeded. Seed globose to ellip-sold, 7-9 min long, with thin seed coat, veinednear hilum. Seedling with hypogeal germina-tion; cotyledons thick, fleshy; epicotyl with afew scale leaves, followed by spiralIy arranged,toothed leaves

Other botanical information Antioris coin-

prises a single variable species, which is subdi-vided into 5 subspecies. Two of these occur intropical Asia and islands of the Pacific, two inMadagascar: subsp. ingdogoscoriensis (H. Ferner)C. C. Berg (synonym: Antioris inodogoscoriensisH. Perrier) and subsp. humbertii (Leandri)C. C. Berg (synonym: Antioris humbertti Lean-dri), and one in mainland Africa: subsp.weiwitschii(Engl. ) C. C. Berg (synonym: Antio-ris weiwitschii Engl. ). Within the latter sub-species 3 varieties are distinguished based ondifferences of texture, indumentum and vena-tion of the leaves: var. weiwttschii(Engl. ) Cor-ner widely distributed in rainforest, var. ofri-cong Achev. (synonym: Antioris of neono Engl. )widely distributed in drier habitats, and varusumborensis (Engl. ) C. C. Berg (synonym: An-tioris usumborensis Engl. ) from eastern DRCongo, Kenya, Uganda and Tanzania. How-ever, intermediates are rather common.

anatomy Wood-anatomical description OAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;22: intervessel pits alternate; (23: shape of al-ternate pits polygonal); 26: intervessel pitsmedium (7-10 prn); 27: intervessel pits large (^10 pin); 31: vessel-ray pits with much reducedborders to apparently simple: pits rounded orangular; 42: mean tangential diameter of ves-sellumina 100-200 prn; 43: mean tangentialdiameter of vessellumina z 200 pin; 46: S 5vessels per square minimetre; 47: 5-20 vesselsper square minimetre. Tracheids and fibres

ANTIARIS 77

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Antioris toxicorio - I, buse of 601e, . 2, twig Mitthmole inflorescences, 3, twig with fernole milo-rescences, . 4, port o117uiting twigRedrown und udopted by Ishoh Syomsudin

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78 TIMBERSl

61: fibres with simple to minutely borderedpits; 65: septate fibres present; 69: fibres thin-to thick-walled. Axial parenchyma: 79: axialparenchyma vasicentric; 80: axial parenchymaanform; 81: axial parenchyma lozenge-allform;83: axial parenchyma confluent; 92: four (3-4)cells per parenchyma strand; 93: eight (5-8)cells per parenchyma strand. Rays: 98: largerrays commonly 4- to 10-senate; (102: rayheight > I min); 106: body ray cells procumbentwith one row of upright and/or square marginalcells; (107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells); (110: sheath cells present); 1/5: 4-12 rays per min. Secretory elements and cain-bial variants: 132: Iaticifers or tanniferoustubes. Mineral inclusions: (136: prismatic crys-tals present); (137: prismatic crystals in up-right and/or square ray cells); (141: prismaticcrystals in non-chambered axial parenchymacells)(M. Thiam, P. 06tienne & E. A. Wheeler)

Growth and development Antioris tort-curio is a non-pioneer light demander. Seed-lings are usually abundant near the mothertree, but experience high mortality in the firstyear. In the shade of the forest, seedlings up to40 cm tall are common, but exposure to fulllight is required for further growth. Underexposed conditions, the tree can grow rapidly;growth rates of 50 cm/year in height are coin-mon in abandoned farmland. Mean annualdiameter growth can reach I cm. Antioris tort-curio has a good self-pruning ability. In WestAfrica trees lose their leaves between Novem-

her and February, and in this period floweringoccurs. However, in wetter types offorest, treescan be more or less evergreen. Fruits are ripein February-March. Trees usually do not fruituntilat least 40 cm in hole diameter. The seedsare dispersed by animals such as birds, mon-keys and antelopes, which relish the fruits

Ecology Antioris toxicorio is found from thewettest to dry forest types, from wet evergreenforest to dry deciduous forest, and even inwooded grassland. It is often common in secon-dary forest, and is an emergent tree of the highforest. In the wetter types of forest, it seems toprefer well-drained sites. Antioris toxicorio canbe found from sea-level to 1800 in altitude. Ithas no special soil requirements. In the driestforest types (e. g. in southern Mali and BurkinaFaso) it is frequently associated with Minetoexcelso ayelw. ) CG. Berg and Catbo pentondro(L. ) Gaertn.

Propagation and planting Fresh seed has

a high germination rate, up to 94% in 2.5-13weeks. Under natural conditions, the seeds loseviahinty rapidly, but when stored in wet sandat low temperatures they still may have a ger-ruination rate of 82% after 5 months. In an

experimental plantation in northern 06ted'TVoire, the survival rate of seedlings after 3.5years was only 49% and the average heightonly 60 cm because of high grazing pressure ofcattle and wild animals; planting of 2 in-tallseedlings wasrecommended

Management Stands of Antioris toxicorioare managed under selective logging systems,and removal of trees is based on prescribedSIIvicultural parameters such as minimumfelling diameter, cutting cycle and felling in-tensity, which depend on stocking in the van-ous countries

Diseases and pests The psyllid Triozomiolamborniis the most important pest of Antioristoxicorio. All stages of the insect can attack theplant, but the greatest damage is done by thenymphs, which killthe apical points of suckersor seedlings, causing dieback, shedding ofleaves, and sometimes death

Harvesting Trees are harvested by powerchain saw and the logs are subsequently crosscut. The log parts are skidded to log yards,from where they are transported to processingmills or to the port for exportIn tropical Asia the latex is tapped by makingscores in the bark with a knife. It is usuallycollected when required as arrow poison or formedicinal purposes, as it is generally usedfresh and only occasionally dried for later useThe bark is harvested by stripping from thetree.

Yield The total stock volume above 30 cm

diameter at breast height was estimated in2001 in Ghana at 825 in31km2 and that above

the minimum felling diameter of 70 cm at 166in8thin2. The latex yield of a scarred tree maybe 100-500 g in 2 days

Handling after harvest Rapid conversionand the application of anti-stain chemicalsupon felling and immediately after sawing areessential, as the wood is liable to sap-stain; itcan become stained up to 15 cm deep. The woodmay also be seriously damaged by various in-sects if it ls not quickly converted or treatedwith preservativesIn tropical Asia the latex from the bark ismixed with other ingredients such as bark orroots of Strychnos and Derris spp. The mixtureis boiled over a fire to obtain a thick paste inwhich the dart and arrow points are dipped

Bark cloth is obtained by shaving off the outerpart from bark stripped from the tree, andbeating and washing the inner fibrous partCareful preparation Is required because traceso11atex may irritate the skin

Genetic resources Antioris toxicor, a Is ex-

tremely widespread and consequently not eas-11y liable to genetic erosion. However, in tropi-cal Asia it is not common, with usually onlylow densities in the forest. In Africa it is gen-erally much more common, but local exploita-tion has severely reduced populations. Thegreat vanability of the species should be stud-ied in more detail, also in relation to its woodand chemical properties and ecology.

Prospects Antioris toxicorio is still a poorlyknown timber tree; in Ghana it is on the list oflesser-used species which are currently beingpromoted. Studies of its technological and In-vestment profile may pave the way for ln-creased utilization as a commercial timber. Its

fast growth and ease of propagation make it apotential plantation species

Major references Agyeman at a1. , 2003;Boer, Brink & SOSef, 1999; Boer & SOSef,1998a; Burki11, 1997; Chudnoff, 1980; CTRADForestry Department, 2003; Katende, BITnie &Tengnas, 1995; Richter & Danwitz, 2000;SIepel, Poorter & Hawthorne, 2004; Voorhoeve,1979.

Other references Akanbi, 1980; Aubr6ville,1959c; Beentje, 1994; Berg, 1977; Berg, 1991;Berg & Hijman, 1989; Berg, Hijman & Weer-denburg, 1984; Berg, Hijman & Weerdenburg,1985; InsideWood, undated; Irvine, 1961; Ker-baro & Adam, 1974; Raponda-Walker & SII-Ians, 1961; Takahashi, 1978; Wagenfuhr, 1979;Wilks & TSSemb6, 2000; World AgroforestryCentre, undated.

Sources of illustration Berg, 1977; Wilks& TSSemb6, 2000

Authors P. P. BOSu & E. Krampah

have been established within these regions andalso in Indonesia, Malaysia, Thailand, the Phil-IPPines, India, the Solomon Islands, CostaRica, Venezuela, Argentina, and several Mri-can countries. Large-scale plantations exist InSouth Africa, and small-scale or trial planta-tions have been established in C6te d'TVoire,

Nigeria, Congo, central Uganda, Zimbabwe,Madagascar and Mauritius, whereas the tree isoccasionally planted as an ornamental else-where marrica

Uses Arguedrig cunninghornii yields an ex-cellent timber, which is used for all kinds oflight construction and interior work, includingmouldings, linings, panelling, domestic floor-ing, shelves, cupboards, general joinery, furni-ture and cabinet work. Special applications arematches, chopsticks and implements. The tim-her is suitable for high-quality plywood and forpremium grade pulp. The tree is also plantedas an ornamental. The seeds are edible andeaten ln tropical America

Production and international trade Aruu-curio timber is commercially Tinportant. Ply-wood was a majorexportitem from Papua NewGuinea until 1980, but declining supplies oflogs from the natural forests resulted in a de-cline in plywood production. The plantationarea of Aroucorio curtnLnghomiiis largest inAustralia, amounting to 45,000 ha with a tim-ber production of 350,000 ing in 2002. In 2001about 1000 ms of Argueorio plywood was ex-ported from Papua New Guinea, at an averageprice of Us$ 3721m3. The timber from Africanplantations is only used locally.

Properties The heartwood is pale yellowishbrown, occasionally with a pinkish tinge andnot sharply demarcated from the straw-coloured sapwood, which may be up to 15 cmthick. The wood is lightweight and soft. It hasa natural sheen and lustre. The density isabout 530 kg/ms at 12% moisture content. Thegrain is straight, texture fine and even.At 12% moisture content, the modulus of rup-ture is 82-90 N/min2, modulus of elasticity8920-13,000 N/min2, compression parallel tograin 39-49 N/mm2, shear 9 N/min2, cleavage39 N/min radial and 56 N/min tangential,Janka side hardness 3200-4230 N and Jankaend hardness 5430-5910 NThe rate of shrinkage is small, from green to12% moisture content 2.2% radial and 3.8%

tangential. The timber seasons satisfactorilywith little or no degrade, although precautionsmust be taken to prevent blue stain. The woodis easy to work with hand and machine tools, it

ARAUCARIA 79

ARAUCARIACUNNINGllAMllAiton ex D. Don

Protologue Lamb. , Descr. Pinus ed. 2, 3: t.79 (1837).

Family AraucariaceaeChromosome number2n=26

Vernacular names Hoop pine, colonial pine,Richmond River pine, Moreton Bay pine (En).Pin de Hoop, araucaria (Fr)

Origin and geographic distribution Arau-carta cunninghornii occurs naturally in NewGuinea and eastern Australia. Plantations

.

80 TIMBERSl

finishes well and takes paint, stains, varnishesand lacquers uniformly without requiring theuse of a filler. It nails and glues well, and it iseasy to peel, making excellent veneer and ply-wood.

The wood is considered to be non-durable incontact with the ground and is susceptible totermite, pinhole borer and marine borer attack.It is reportedIy variable in its resistance toImpregnation.The major constituents of the essential oil ob-tamed from the leaves of Argueorio cunning-hornii grown in south-western Nigeria were orpinene (14.8%), terpinen-4-o1(14.7%), shyobunol(8.9%) and spathulen01(8.8%).

Botany Large evergreen tree up to 60(-70)in tall; hole straight, cylindrical, branchless forup to 45 in high and up to 200 cm in diameter;bark reddish brown to blackish brown, trans-versally wrinkled, fissured, ridged or plate-like,peeling either horizontally or in slabs;branches in whorls, horizontally displayed,with UPCurved tips. Leaves arranged spiralIy,crowded, simple and entire, sessile and broadlyattached, lanceolate to triangular, curved,pointed at apex, many-veined. Cones unisex-ual; male cones up to 8 cm long, hanging; fe-male cones 6-10 cm x 5-8 cm when matureand later disintegrating, with spiny, wingedcone scales, each scale with I ovule partlyfused with it. Seed triangular, 2-3 cm x I cmexcluding the membraneous wings. Seedlingwith epigealgerminationArgueorio comprises 19 species and occurs inNew Guinea, eastern Australia, some islands ofthe Pacific Ocean and South America. NewCaledonia is richest in species (13). Aruucoriohunsteinit K. Schum. , originating from NewGuinea, and Arguedrig origustifolio (Bertol. )Kuntze from South America have been plantedin experimental plantations in Uganda. Somespecies are occasionally planted in tropicalAfrica as ornamentals, e. g. Aruucorio colum-

Hook. and Argueorio heterophyllanor, s

(Sansb. ) Franco, in addition to Aruucorio cun-rimghomiiTrees of Arguedrio cunninghornii usually startbearing cones at 15-25 years of age. Femalecones take about 2 years to ripen after pollina-tion. The winged seeds are dispersed by windJuvenile growth is generally slow, but treesmay reach a height of 33 in and a hole diameterof 42 cm in 30 years, In Congo 12-year-old treesplanted at a density of 1000 treesA1a had anaverage height of 18.5-20.5 in, and an averagediameter at breast height of 20 cm. The aver-

age annual volume increment in plantations inPapua New Guinea and Australia is 11-14msftia, but in successful plantations 20-30ing/ha can be reached.

Ecology Under natural conditions in NewGuinea and Australia, Aroucorio cunning-horniiis most common above 1000 in altitude

(up to 2750 in)in areas with high rainfalland atemperature range of 9-26'C. It occurs on avariety of rainforest soils, from acid to neutralAroucorio cunninghornii can tolerate mild DC-casionalfrosts, but does not tolerate fire

Management Arguedrig cunninghornii canbe propagated by seed sown in beds of well-rotted sawdust or friable sandy loam, under70-90% shade. Pre-germination and tubing aswellas direct sowing may be feasible. To obtainthe seeds, cones are collected before they disin-tegrate on the trees. Cones that do not disinte-grate within 10 days of harvesting are jinma-ture. Often, less than I% of the seeds collectedfrom a tree are viable. Filled and unfilled seedsmay be separated by flotation. The 1000-seedweightis 330-500 g. Fresh viable seeds give upto 90% germination. The seeds can be storedfor a few years at temperatures below 3'C. Theseeds can also be dried to 2% moisture content

without damage and be kept at temperaturesof -18'C or lower, resulting in a viahinty ofabout 75% after 6 years of storage. In-vitropropagation has been practised successfullyStern segments with 3-5 leaf axils, excisedfrom the upper portion of the main stem of 2-year-old seedlings, produce orthotropic budsfrom the concealed axillary menstems whencultured on a medium of half-strength Mura-shige and Skoog inorganic salts. Up to 80%rooting is obtained, and after 2 weeks theplantlets can be transferred to a mixture ofpeat and perlite and maintained at a relativehumidity of 90-95%. The young plants are sub-sequently transferred to normal greenhouseconditions and then to the field, with less than5% mortality. Inoculation for mycorrhiza for-mation is necessary. Seedlings reach plantablesize in 18-24 months. When planted out intothe field, spacing is usually 3 in x 3 in, butwider spacings of up to 7 in x 7 in have alsobeen practised.The main form of planting is in monocultureplantations. Underplanting in Pinus planta-tions has been tried with varying results. Weedcontrol during juvenile stages Is essential andthe plants respond well to fertilizers. Pruningis usually first done when the trees are about 6years old. In a 35-year-old plantation about 100

stems per ha are maintained. A height of 30 inand a hole diameter of 50 cm are considered

satisfactory in plantations in Australia; thisrequires a rotation of 50-60 years on goodsites. The most significant fungal disease ofplantation-grown Aroucorio cunninghorniiinPapua New Guinea and Australia is root andheart rot caused by Phellinus itoxius. Armil-10no attack has been recorded for plantationsin Zimbabwe, and Porio root disease in EastAfrica. In Papua New Guinea the most seriouspests in plantations are the branchlet-miningbeetle Hylurdrectonus uruucorioe and the wee-vil Vonop0 o6erthueri. Termites may alsocause serious damage. Attacks by groups offruit bats (Bidolon heIuum), stripping the treesof their leaves and bark, have been recordedduring long dry seasons in C6te d'Ivoire, result-ing in almost complete destruction of 8-12 intallornamentalAroucorio cunninghorniitrees.The timber is prone to blue stain infection andshould be treated with fungicides and removedrapidly from the forest after logging.

Genetic resources and breeding In NewGuinea large stands of Araucorio cunning-hornii have been depleted due to heavy tlinherexploitation, but a large number of smallstands still exist in the wild. Extensive geneticimprovement research has been conducted inAustralia and Papua New Guinea, encompass-ing provenance and progeny trials.

Prospects Much is known about propaga-tion and SIIviculture of Aruucorio cunning-hermit. Extensive plantations have alreadybeen established and they produce largeamounts of timber, especially in Australia. Inplantations rapid growth is possible, withoutaffecting the wood quality. The results of theexperimental plantations in different regionsin tropical Africa should be evaluated before adecision can be taken on the economic feasibil-

ity of larger-scale plantations.Major references Arentz, Keating & Inc.

1993; CAB International, 2005; Katende,Birnie & Tengnas, 1995; World ConservationMonitoring Centre, 2000.

Other references Anonymous, 1982; Beinat a1. , 1996; Byabashaija & BSegu, 2003; Diet-ers, Nikles & Johnson, 2003; Malagnoux &Gautun, 1976; 01awore & Ogunwande, 2005;Takahashi, 1978; Telllier, 1988.


Based on PROSEA 5(I): Timber trees: Majorcommercial timbers.

ADBREVILLEAPLATYCARPAPellegr.

Protologue Bull. SOC. Bot. France 80: 467(1933)


Vernacular names Dabema noir(Fr).Origin and geographic distribution AMbre-

vineo plotycorpo occurs from Guinea and Si-erra Leone to Gabon and DR Congo.

Uses The timber of Aubreuil!eop!otycorpo isknown as 'k16k16' in C6te d'Ivoire and 'dabema

noir' in Guinea. The wood is suitable for con-

struction purposes. In Gabon the wood is usedfor joinery. Probably it is also used elsewhereas a substitute for the wood of Piptodeniostrumo17iconum (Hook. f. ) Brenan, traded as 'dabema'or 'daboma'.

Properties The heartwood is pale brown orgreyish brown with a violet tinge and darkerstreaks, distinctly demarcated from the palersapwood. The grain is often interlocked, tex-ture moderately coarse. The heartwood Is rela-tively hard. The wood dries slowly and is un-treatable with preservatives.

Botany Large tree up to 50 in tall; holestraight, cylindrical, branchless for up to 25 in,up to 100(-150) cm in diameter, with thickbuttresses up to 3 in high often extending inflares along the lower part of the hole; barksmooth but in large trees becoming scaly, greywith red-brown patches; crown dome-like,dense, dark green. Leaves alternate, hipin-nately compound with (3-)4-8 pairs of oppositepinnae; stipules inlnute, early caducous; petiole4-12 cm long, rachis 6-26 cm long, groovedabove; leaflets in (6-)8-15(-21) pairs perpinna, opposite, sessile, asymmetricalIy oblongto obovate, 1.5-5 cm x 0.5-2 cm, obtuse tonotched at apex. Inflorescence a spikelike ra-ceme up to 22 cm long, several togethergrouped in a terminal panicle, many-flowered;peduncle densely rusty hairy. Flowers bisexual,regular, 5-merous; pedicel up to 2.5 min long;calyx cup-like, I-1.5 min long, shortly toothed;petals fused at base for c. I mm, lobes 2-2.5min long, hairy; stamens 10, united at base,filaments c. 3 mm long; ovary superior, oblong,shortly stiped, flattened, sparsely hairy, style c.2 mm long. Fruit an indehiscent, papery, ob-long pod 13-22 cm x 3.5-5.5 cm, with a thinstipe up to 2 cm long, twisted at base, reticu-late, pale yellow when ripe, I-3-seeded. Seedsround to kidney-shaped, flat, up to I cm x 1.5cm. Seedling with epigeal germination; hypo-cotylup to I cm long, glabrous, epicoty1 5-8 cm

AUBREVILLEA 81

82 TIMBERSl

long; cotyledons kidney-shaped, fleshy, notspreading; first 2 leaves oppositeAubreuilleo comprises 2 species. It seems tohave an isolated position in the tribe Mi-inOSeoe. Aubreuil!eo resembles Piptodenios-trum, which has a similar hole and crownshape but differs particularly in its leaves, withmore numerous pinnae and leaflets, stamenswith glandular arithers and dehiscent podscontaining up to 12 seeds. The wood of Aubre-un!eo herstingit (Harms) Fellegr. is probablyalso used for joinery and as a substitute of Pip-todentostrum @1nconum wood. AMbreui!ledherstingit occurs from Sierra Leone to DRCongo and differs from Aubreuil!eo plotycorpoin its more numerous and more narrowly ob-long leaflets. A bark decoction is used in tradi-tional medicine in Congo as an anodyne and toprevent abortion. In C6te d'Ivoire a leaf decoc-tion Is administered in enema as a laxative and

crushed leaves are applied externalIy as ananodyne. A root maceration in palm wine isdrunk to treat gonorrhoeaGermination of AMbreui!leg pinkycorpo is usu-ally within 8-15 days after sowing. Trees areshortly deciduous at the time the tree is fruit-ing; when new leaves develop the crowns showa brilliant red colour. However, they may alsobe evergreen. In Sierra Leone and C6te d'TVoireflowering is in DecemberJanuary, and fruit-ing from February to May. Fruits are usuallymature at the end of the dry season. Wholefruits are dispersed by wind. The roots of AM-byeui!!eoplotycorpo are nodulated

Ecology Aubreuilleo plotycorpo occurs indense evergreen and semi-deciduous lowlandforest.

Management The 1000-seed weight of AM-67eutl!eo plotycorpo is about 67 g. In generalAubreuilleo plotyco, :per trees occur scattered inthe forest, although seedlings may be abundantnear the mother tree.

Genetic resources and breeding AMbre-uilleo plotycorpo as well as Aubreuilleo herst-ingii are widespread in the forest zone of Westand Central Africa, but they occur scatteredThere is little reason to consider them threat-

ened, although some concern has been ex-pressed, e. g. in Guinea

Prospects There is little information onwood properties and ecological requirements ofAubreu, Ileo, but it apparently occurs too scat-tered to be of future economic importance. Itseems to be barely exploited, but the amountsmay be underestimated because it may bemixed with the wood of similar Mimosaceoe

species.

Major references Burki11, 1995; Lewis ata1. , 2005; Normand & Paquis, 1976; Viniers,1989; Voorhoeve, 1979

Other references Aubr6ville, 1959c; de inMensbruge, 1966; Diabate at a1. , 2005; Dudek,Forster & missenbauer, 1981; GIIbert & Bou-tique, 1952; Hawthorne, 1995; Keay, 1958c;Neuwinger, 2000; Saville & Fox, 1967.


AUCOUMEAKLAINEANAPierre

Protologue Bull. SOC. Linn. Paris, n. s. I1241 (1896).

Family BurseraceaeChromosome number2n=26

Vernacular names Okoum6, Gaboon ina-hogany (En). Okoum6 (Fr). Ocum6 (Po).

Origin and geographic distribution Au-coumeu hloineono occurs naturally in westernand central Gabon and in continental Equato-rial Guinea, south to Congo, where it is re-stricted to the Chainu and Mayombe MassifsThere are small natural stands in southern

Garnero0n, near the border with EquatorialGuinea. Its natural occurrence in Nigeria, nearthe border with Cameroon, needs confirmation.Okoum6 has been planted for timber bothwithin its natural range in Gabon and Camer-o0n, and elsewhere in Cameroon and C6ted'Ivoire. Small-scale planting trials have beenmade in Congo, DR Congo, Ghana, Madagas-car, Indonesia, Malaysia, Sunname andFrench Guiana.

Uses Okoum6 is one of the best timber spe-cies for plywood. It is a major commercialtim-

.

,

Aucoumeo hidineuno - wild

L. .

o

,^

..

her in Gabon and Equatorial Guinea, repre-senting more than 60% of timber production,while it is of lesser importance in Congo. It ismade into blockboard, particle board and ve-neer, and is widely used in boat building fordecorative interior paneUing and for exteriorapplications. The wood is also suitable for lightinterior construction, carpentry, furniture,sports equipment, cigar boxes and packingcases. Logs are traditionally used to make ca-noes. The wood can be used as firewood and issuitable for the production of pulp for paper-making.Bark resin is used for torches and o111amps inGabon and Equatorial Guinea, and also forcosmetic applications in skin care products andnail polish. It is also applied to treat superficialwounds and abscesses, and as a water disinfec-tant. The astringent bark is used to treat diar-rhoea

Production and international trade Okou-in6 timber is mainly exported as logs to Asia(69% in 2002) and Europe (24% in 2002). In the1960s, Gabon exported 1.1-1.4 million ing ofokoum6 per year, Congo about 250,000 in31yearand Equatorial Guinea 60,000-120,000in31year. According to ATIBT (AssociationTechnique Internationale des Bois Tropicaux),exports of logs, sawn timber, veneer and ply-wood from Gabon in 2002 were 1.0 million in3,60,000 ina, 108,000 ms and 30,000 ing respec-tively. The average export prices of okoum6sawn timber, veneer and plywood in 2001 wereUs$ 176 per ms. Us$ 360 and Us$ 233 respec-tively. China is the largest importer of okoum6with 820,000 ina in 2002, followed by Francewith 230,000 ingSmall quantities of resin are exported fromGabon to France for cosmetic applications

Properties The heartwood is salmon pink topale pinkish brown or reddish brown, darken-ing upon exposure to light to a mahogany-likecolour, and is distinctly or indistinctly demar-cated from the white to pale grey sapwood. Thegrain is usually slightly interlocked, texturemedium to moderately fine. Quarter-sawn sur-faces with a striped or inottled figure. Growthrings distinct due to alternating paler anddarker layers.Okoum6 is a lightweight hardwood. At 12%moisture content, the density is (320-)350-450(-580) kg/in, . Okoum6 wood dries "apidly,with little degrade and few defects in either airor kiln drying. The shrinkage rates from greento oven dry are 3.4-7.0% radial and 5.3-11.3%tangential.

The wood is comparatively soft. At 12% inois-ture content, the modulus of rupture is 51-109N/mm2 modulus of elasticity 5500-10,800N/min2, compression parallel to grain 25-44N/mm2, shear 4-8 N/mm2, cleavage (6-)9-22N/min tangential, Janka side hardness 1690-2510 N andJanka end hardness 2790-3560 NThe wood is easy to work with both hand- andmachine tools, but worked surfaces tend to bewoolly, so cutting edges must be kept sharp. Itcontains up to 0.3% silica and sawing requirestungsten-carbide-tipped cutters as saw teethblunt rather quickly. When planing, an angleof 20' is necessary to prevent tearing. Thewood responds well to sanding, can be nailedwithout pre-boring, and glues and stains well.It can be polished to a lustrous surface. Dimen-SIonal stability is rated as fair, and seasonedwood is reported to move moderately aftermanufacture. Okoum6 is often rotary-cut forthe production of plywood; figured wood issliced into highly decorative veneers for panel-ling and cabinet makingOkoum6 has little natural resistance to decayLogs are susceptible to forest longhorn beetleattack. While the sapwood is readily attackedby Lyetus beetles, the heartwood is resistant,but susceptible to termltes, marine borers andfungal attack. The heartwood is Impermeableto preservatives. Glues with fungicides areused for plywood which may come into contactwith moisture. The energy value of the wood is29,970 kJ/kg.Several terpenoids are present in the barkresin, including orterpineol and p-phellandreneand several tetracyclic and pentacyclic triter-penes

Description Dioedous, medium-sized to largeevergreen tree up to 50(-60) in tall; hole cylin-drical, often contorted and bent, up to 110(-240) cm in diameter, with buttresses up to 3 inhigh, and clear of branches up to 21 in; bark05-2 cm thick, greyish to orange-brown, de-taching in more orlessthick rectangular brownscales revealing orange bark in adult trees,with lenticels, slash strongly resinous, pinkish-red, fibrous; crown rather open structuredLeaves alternate, imparipinnate; stipules ab-sent; rachis up to 40 cm long; leaflets 7-13,with stalks up to 4 cm long, ovate to oblong,10-30 cm x 4-7 cm, rounded at base, acumi-nate at apex, entire, leathery. Inflorescence anaxillary or terminal panLcle up to 20 cm long;male inflorescence comprising up to 5 timesmore flowers than female one. Flowers unisex-

ual, regular, 5-merous; sepals lanceolate, up to

AUGOUMEA 83

.

84 TIMBERSl

6\

5

11

I

^ <,.

, -II~\,,/I

-,

I

I

Iygona1; (26: intervessel pits medium (7-10F1m)); 27: intervessel pits large Q 10 F1m); 31:vessel-ray pits with much reduced borders toapparently simple: pits rounded or angular; 32:vessel-ray pits with much reduced borders toapparently simple: pits horizontal(scalariform,gash-like) to vertical(palisade); 42: mean tan-gential diameter of vessellumina 100-200 prn;46: s 5 vessels per square minimetre; 47: 5-20vessels per square minimetre; 56: tyloses coin-mon. Tracheids and fibres: 61: fibres with SIm-

PIe to minutely bordered pits; 65: septate fibrespresent; 68: fibres very thin-walled. Axial pa-renchyma: 75: axial parenchyma absent orextremely rare; 78: axial parenchyma scantyparatrachea1; 92: four (3-4) cells per paren-chyma strand; 93: eight (5-8) cells per paren-chyma strand. Rays: 97: ray width I-3 cells;106: body ray cells procumbent with one row ofupright and/or square marginal cells; (107:body ray cells procumbent with mostly 2-4rows of upright and/or square marginal cells);1/5: 4-12 rays per mm. Mineral inclusions:159: silica bodies present; 160: silica bodies inray cells(M. Thiam, P. 06tienne & E. A. Wheeler)

Growth and development Okoum6isshal-low-rooted, with virtually no taproot. At 6-12years, populations become organized into 'bio-logical cells' through joining root grafts. When10-25 years old, natural and planted popula-tions segregate into dominant and suppressedtrees. Trees reach an average height of about10 in in the first 4 years; from 10-50 years, thegrowth rate of dominant trees is about I in peryear, and eventually they reach a height ofabout 50 in. Suppressed trees grow very slowlyand reach 15 in in 50 years, Growth in holediameter is variable and depends on age, standdensity, site fertility, hierarchical position ofthe tree and thinning rates. In coastal Gabon,hole diameters of 30-80 cm after 50-60 yearsfor dominant trees and 10-40 cm for sup-

pressed trees have been recorded. The meanannual diameter increment for dominant trees

gradually decreased from 1.4 cm (7-12 yearsold) to 0.7 cm (50-60 years old) and for sup-pressed trees from 06 cm (5-20 years old) to0.1 cm (10-40 years old). Other studies at dif-forent sites in the interior of Gabon recordedmean bole diameter growth rates of 0.5-0.7cm/year, without taking into account the pres-ence of dominant and suppressed individuals.New leaves appear from September to Decem-her and are bright red for one week. Trees startto flower when they are about 10 years old, but

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I,

AMcoumeo Mumeono - I, base of 601e, . 2, led/: 3,fernole /lower (front sepols undpetols reinoued),4, mole flower ifront sepals undpetols reinoued),.5, fruit with opening uolues, . 6, fruits ofter drop-ping ofuo!Des.Redrown und udopted by MM. Spitteler

5 min long, hairy, greenish; petals spatulate,5-6 min long, hairy on both sides, whitish; diskconsisting of 2-10bed nectaries; male flowerswith 10 stamens and rudimentary pistil; re-male flowers with 10 staininodes and a supe-nor, 5-celled ovary, style columnar, with head-like stigma. Fruit a capsule up to 5 cm x 3 cm,opening with 5 valves from the base, 5-seeded.Seeds enclosed by endocarp ('pyrenes'), ovoid,extending into a wing 2-3 cm x 0.5 cm. Seed-ling with epigeal germination; cotyledonsrounded, leafy.

Other botanical information Aucoumeocomprises a single species, and is characterizedby its extra-staminal disk and dry, dehiscentfruit fuseudocapsule), which after opening re-leases 5 seeds covered by a winged endocarp

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-

I I

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fruiting only begins after 15 years. Floweringstarts in August and lasts for I-2 months de-pending on weather conditions. Okoum6 isdioecious; male and female flowers are foundon different trees. Individual flowers last for afew days and are pollinated by bees and fliesFruiting starts in September; fruits are fullygrown in about 40 days, but mature after about80 days. Fruiting is annual, but large quanti-ties of seeds are produced only every 2-3 yearsA healthy, dominant mature tree can produceup to 20,000 seeds. Seeds are wind-dispersedup to 80 in from the parenttree

Ecology Okoum6 occurs naturally from sea-level to about 600 in and occasionally to 1400in altitude in areas with a mean annual rain-fallof 1200-3000 mm and a marked dry season(less than 50 min/month) from June to Au-gust/September, during which evaporation Isreduced by the relatively low temperatures andhigh relative humidity. Mean annual tempera-tures range from 23-26'C, with mean maxi-mum temperatures of the hottest month 32-36'C, and mean minimum temperatures in thecoldest month 10-15"C. The natural distribu-tion of AMcoumeu hintneono is directly relatedto rainfall. In the north-east of Gabon, a seconddry season (during the fruiting period) ac-counts for the absence of the species there,whereas in the south of Gabon the annual rain-fallofless than 1200 mm is the limiting factorOkoum6 is a long-lived pioneer of, in particu-Iar, large forest clearings and fire-protectedsavanna edges, where it often becomes mono-dominant. It requires full sun to grow wellSeedlings and saplings can, however, survivein shade for some years, Huge lridividuals oc-cur in what seems to be virgin forest, but is infact old secondary forest.Okoum6 grows well on a wide range of acidsoils (such as ferralitic arenosols, ferraliticsoils, podzoluvisols) developed on various sub-strates. It can grow on infertile sandy soils butprefers fertile, deep sandy-clay-loams. It cantolerate a certain degree of impeded drainage,but notlong periods of waterlogging

Propagation and planting Seeds lose vi-ability within I month in the field, but can bestored at 4'C in airtight containers for up to 3years after being dried to 8% moisture content.The mean 1000-seed weight is 98 g. Propaga-tion by seed is the preferred method. Seeds donot require pre-treatment and can be sowndirectly in polythene bags (20-30 cm high, 10-15 cm in diameter) filled with a mixture ofsand and clay. The addition ofNPK fertilizer is

recommended. Two seeds are placed per bag;one seedling is selected after 3-4 weeks. After2.5 months, the seedlings are 20-25 cm tall,have 5-7 simple leaves and are ready fortransplanting. Propagation by grafting andcuttings is possible.

Management Good regeneration requlres asufficient number of seed trees, large canopyopenings or clearings (> 2500 in2) and clean soilduring the fruiting season. These conditionsare found in shifting cultivation or in loggedareas (e. g. log yards and wide logging tracks)Selective logging or natural tree fall events donot produce sufficiently large canopy openingsIf site conditions are favourable, okoum6 doinI-nates regrowthTwo main site preparation methods are usedfor establishing plantations. The first ismechanized clear felling, in which existingwoody vegetation is cleared using bulldozersand placed into windrows and burned. Seed-lings are planted between the windrows. Thesecond consists of cutting the vegetation to 50cm above the ground to allow resprouting andregrowth. Seedlings are planted in lines cutthrough the regrowth. In both methods, exist-ing large trees are killed. Recommended spac-ings for seedlings are from 625-950 trees/ha. Itis necessary to eliminate climbers, especiallyMikonio species, and trees such as Musongocecropioides R. Br. ex Tedlie, which compete forlight and space, for up to 5 years after plant-Ing

Thinning in both natural and artificial standsis advisable, but should be conducted carefullyto avoid increased sensitivity to black cankerattack resulting from lateral illumination ofthe stem. In both mixed and almost purestands, thinning is beneficial for diametergrowth. Suppressed trees are much more re-sponsive to thinning than doinlnant ones. Inalmost pure stands, thinning should be re-stricted to those young stands of less than 15

because in older stands thinning willyears,

remove potentially commercial treesIn plantations, a thinning regime to reducestern density to 350 stems/ha after 5 years,200-250 stemsftia after 10 years and 150stemsftia after 15 years is recommended. Ifresources only allow 2 thinning operations,stand density can be reduced to 250-300stems/ha after 5 years and 150 stemsftia after13 years. If thinning is not by cutting but bykilling standing trees, it involves girdling ok-oum6 trees, or girdling and poisoning otherspecies, taking care to avoid intense hole illu-

AucouMEA 85

.

86 TIMBERSl

mination of lower strata trees. Okoum6 trees

should not be poisoned because of the risk ofaffecting adjacent trees via the connected rootsystems.

Diseases and pest Okoum6 is attacked byvarious pathogens, which are only serious inpure stands. The most important disease isblack canker, a complex infection which beginswith a primary infestation of scale insects(mainly Astero1ecunittm pustulans) spread byants (Cremotogoster, Oecophyllo). Subsequentbark injuries are then infested by a secondaryfungal pathogen, Botryodip!odio theo670moe,resulting in an external proliferation of smutwhich blackens the bark and causes abnormal

resin secretion. Very dense or severely thinnedplantings and regrowth on abandoned humansettlements are particularly prone to blackcanker. The problem can be avoided by goodSIIvicultural practices, such as selecting a suit-able planting site, adequate spacing and care-ful thinning. Seedlings and young plants areattacked by psyllids (Pseudophocopteron spp. )and centipedes, although serious infestationsonly occur in badly managed nurseries andonly affect young plantings. During Decemberand January, the foliage of both young andadult trees is often attacked by caterpillars ofthe moth PIeuroptyo bolteotu, sometimes re-SUIting in total defoliation of pure stands, buttrees only suffer a reduction in growth. Locally,elephants cause serious destruction in youngand pure stands

Harvesting Only I or 2 okoum6trees per haare usually extracted in natural forest. Logsfloat and are commonly transported by riverDuring seasonalIy low river levels or wherethere are no rivers, logs are transported byroad or rail

Yield In established plantations, the esti-mated potential yield varies from 7 ms/ha/yearat 43 years on a moderately fertile site withoutthinning, to 11 in8tha/year at 32 years on afertile site with thinning. However, actual pro-duction is lower because 30% of trees above

minimum felling diameter (70 cm) are usuallynot suitable for timber due to their poor shapeThe yield potential in pure natural stands canbe 5.5-7.5 in3ftia/year, but commercial yieldsare I-2 in8tha/year due to selective logging

Handling after harvest 80% of okoum6timber is peeled for plywood production. It isconsidered one of the best woods for this pur-pose. Peeling can be performed with or withoutcontrolled steaming or seasoning, although thelatter Increases the overall quality of end prod-

UCtS.

Sawn timber is becoming important as bothGabon and Congo are attempting to developtheir wood-processing capacities. While good-quality logs are exported, or locally processedfor plywood, lower grades are now used forsawn timber

Genetic resources In order to safeguardthe genetic diversity and sustained productionof okoum6, stands should be identified andprotected throughout the natural range of thespecies. Special attention should be given tonatural stands not having a long history ofexploitation. Field experiments indicate a poorperformance of provenances from sites havinga long history of exploitation

Breeding There are currently no major se-Iection and breeding programmes as all plant-ing has ceased, the current trend favouringnatural regeneration. However, the few studieson provenance and progeny testing revealedlarge vanability. This diversity at the prove-nance/progeny levelis confirmed by recent ge-netic studies, indicating that there are possi-hinties for selecting better planting materialOn the other hand, a study of chloroplast DNAvariation showed a relatively low level of dif-forentiation between okoum6 populations inGabon

Prospects Okoum6 will remain a majorcommercial timber for Gabon and EquatorialGuinea, but its climatic requirements restrictits importance elsewhere. Breeding and selec-tion programmes are needed.

Major references Brunck, Grison & Maitre,1990; CTFT, 1988a; de Kam at a1. , 1996; Fuhr,Nasi & Delegue, 2001; Fuhr, Nasi & Minkou6,1998; Koumba Zaou at a1. , 1998; Koumba Zaou,Mapaga & Verkaar, 1998; Leroy-Deva1, 1976;Muloko-Ntoutoume at a1. , 2000; Nasi, 1997

Other references Aubr6ville, 1962a;Biraud, 1959; Biraud & Catinot, 1960; Cabal16,1978; de Saint-Aubin, 1963; Grison, 1978a;Grison, 1978b; Grison, 1978c; InsideWood, un-dated; Lebacq & Deschamps, 1964; Leroy-Deva1, 1973; Leroy-Deva1, 1974; Leroy-Deval,1975; Raponda-Walker & Sillans, 1961;Reltsma, 1988; Richter & Danwitz, 2000;Tessier, Delaveau & Piffault, 1982; Touzard,1964; van Valkenburg, Ketner & Wilks, 1998;White & Ahemathy, 1997; Wilks & TSSemb6,2000

Sources of illustration Aubr6ville, 1962a;Grison, 1978b; Tailfer, 1989; Wilks & Issemb6,2000

Authors J. L. C. H. vanValkenburg

I

AUTRANELLACONGOLENSIS (De Wild. )Achev.

Protologue Veg. Ut. Mr. Trop. Franc. 9: 271(1917)

Family SapotaceaeChromosome number2n=26

Synonyms MimusopsletestuiLecomte (1920)Origin and geographic distribution AMtro-

itel!o congo!errsis occurs from south-easternNigeria east to the Central African Republic,and south to Gabon and DR Congo.

Uses The wood (trade name: inukulungu)isused for heavy construction, heavy-duty floor-ing, bridges, sluice gates and other water-works, railway sleepers, stairs, poles, vats,vehicle bodies and quarter-sliced veneer. It islocally used to make canoes and paddles. It issuitable for joinery, marquetry, turnery, Inte-nor trim, furniture and cabinet work, sportinggoods, toys and novelties, musical instruments,draining boards, agricultural implements, shipand boat building and mine propsThe fruits are reported to be edible. An edibleo111s extracted from the seeds; it is used forcooking in Gabon. Dry ground bark is appliedto cuts and sores. A bark decoction Is taken totreat gonorrhoea, syphilis, dyspepsia, colic andcolds, and applied externalIy to treat fever,pain, skin diseases and wounds. The brokenseed coats are strung together to make rattles

Production and international trade AMtro-

itel!o congo!errsis timber Is exported in smallamounts from Central Africa. According toATIBT figures, in 2001 the log export fromCameroon was 235 in3 and from Gabon 105 in3.In 2003 about 930 ing of sawnwood was ex-

ported from Gamero0n

Properties The heartwood is reddish brownwith dark brown streaks, and usually distinctlydemarcated from the greyish or brownish sap-wood, which is I-3 cm wide. The grain Isstraight or sometimes interlocked, texture fineand even. The wood is heavy, with a density of910-990 kg/in3 at 12% moisture content. Itdries slowly, with severe risk of deformation; itneeds to be dried with care and quartersawingis recommended. The shrinkage rates are inod-erate to high, from green to oven dry 3.9-9.0%radial and 4.1-9.1% tangential. Once dry, thewood is often unstable in service.

At 12% moisture content, the modulus of rup-ture is 120-199 N/min2, modulus of elasticity11,800-21,900 N/mm2, compression parallel tograin 66-105 N/mm2, shear 6.4-13.2 N/min2,cleavage 18.1-22.5 N/min, Janka side hardness10,230 N and Janka end hardness 12,370 N.The wood saws easily but slowly, with seriousblunting effect on tools due to the high densityand the presence of silica. Stentte-tipped saw-teeth and tungsten-carbide-tipped cutting toolsare recommended. The wood planes fairly eas-11y with a smooth finish. It holds nails andscrews well, but pre-boring is needed. The glu-ing properties are poor to satisfactory. It can beused for sliced veneer, but rotary peeling isdifficult. It is resistant to acids, making it suit-able for chemical containers and laboratorytables.

The wood is durable and resistant to fungi anddry-wood borers, but sometimes there is slighttermite attack. It is considered resistant tomarine borers, but in tests in Italy it was notresistant. It is resistant to impregnation withpreservatives. The wood dust may cause seri-ous irritation to nose, throat and eyes. Thepresence of saponins has been reported for thewood, which contributes to the resistance tofungi.


of Autrunello congolensis is similar to that ofBanjonel!o tortspermo Pierre (inoabi), which isused for similar purposes. It also resemblesTieghemello wood, but this has a lower density.

Description Medium-sized to large tree upto 40(-50) in tall; hole straight and cylindrical,branchless for up to 30 in, up to 150(-200) cmin diameter, often slightly buttressed at base;bark surface brown, fissured, inner bark red tobrown, fibrous, exuding latex; crown umbrella-shaped, flattened; branches with numerous leafscars. Leaves arranged spiraUy, clustered atends of branches, simple and entire; stipulestriangular, early caduceus; petiole slender, 4-5

AUTRANELLA 87

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Autronel!o congo!errsis - wild

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10 cm long, leafy but leathery;leaves alternatefrom the beginning

Other botanical information Autronello

comprises a single species. It seems to be re-lated to Mimusops and neghemello, the formerof which differs in lacking stipules and havinga shorter corolla tube and smaller fruits, thelatter in lacking stipules and having stamensand staininodes inserted near the base of the

corolla tube and larger seed scar. Boillonellotortspermo resembles AMtronello congo!errsis,but differs in its shorter corolla tube, distinctstamen filaments and larger seed scar

Anatomy Wood-anatointcal description OAWAhardwood codes)Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;7: vessels in diagonal and/or radial pattern; (8:vessels in dendritic pattern); (10: vessels inradial multiples of 4 or more common); (11:vesselclusters common); 13: simple perforationplates; 22: intervessel pits alternate; 26: in-tervesselpits medium (7-10 prn); 30: vessel-raypits with distinct borders; similar to intervesselpits in size and shape throughout the ray cell;31: vessel-ray pits with much reduced bordersto apparently simple: pits rounded or angular;32: vessel-ray pits with much reduced bordersto apparently simple: pits horizontal(scalari-form, gash-like) to vertical (palisade); 33: ves-sel-ray pits of two distinct sizes or types in thesame ray cell; (34: vessel-ray pits unilateralIycompound and coarse (over 10 prn)); 42: meantangential diameter of vessellumina 100-200I'm; 47: 5-20 vessels per square minimetre; 56:tyloses common. Tracheids and fibres: 61: fi-bres with simple to minutely bordered pits; 66:non-septate fibres present; 70: fibres verythick-walled. Axial parenchyma: (76: axial pa-renchyma diffuse); 86: axial parenchyma innarrow bands or lines up to three cells wide;87: axial parenchyma reticulate; 93: eight (5-8)cells per parenchyma strand; 94: over eightcells per parenchyma strand. Rays: 97: raywidth I-3 cells; 107: body ray cells procumbentwith mostly 2-4 rows of upright and/or squaremarginal cells; (108: body ray cells procumbentwith over 4 rows of upright and/or square mar-ginal cells); 115: 4-12 rays per mm. MineralInclusions: 136: prismatic crystals present; 142:prismatic crystals in chambered axial paren-chyma cells; 159: silica bodies present; 160:silica bodies in ray cells; (161: silica bodies inaxial parenchyma cells)(H. Beeckman & P. 06tienne)

Growth and development The fruits take

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AMtronello congole, 1sts - I, buse of 601e, 2, flow-ering bronch, ' 3, fruit, . 4, seeds.Redrawn grid oddpted by Ishoh Syomsudin

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cm long, channelled above; blade elliptical-oblong to slightly obovate, 10-15 cm x 4-5 cm,rounded to curieate at base, rounded to shortlyacuminate at apex, leathery, glabrous, pin-nately veined with c. 15 pairs of indistinct Iat-eral veins. Inflorescence an axillary fascicle atthe ends of branches. Flowers bisexual, regu-Iar; pedicelc. 2 cm long; sepals free, in 2 whorlsof 4, oblong, c. I cm long, obtuse at apex, hairy;corolla with c. 7.5 min long tube and 8 lobes c2.5 min long, divided to base into 3 segmentswith median segment erect and clasping sta-men, and lateral segments much larger andspreading or reflexed, segments shortly hairyinside; stamens inserted at top of corolla tubeopposite corolla lobes, almost sessile, alternat-Ing with short staininodes; ovary superior, el-lipsoid, large, hairy, 8-celled, gradually nar-rowed into the short style. Fruit a fleshy, ovoidto ellipsoid I(-2)-seeded berry 5-10 cm long,warty, yellowish green. Seed obovoid, slightlyflattened, c. 4.5 cm long, glossy brown, withfairly large rectangular scar. Seedling withepigeal germination; hypocoty1 14-22 cm long,epicoty1 I-4.5 cm long; cotyledons elliptical, 6-

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4

10-14 months to ripen. It has been reportedthat in the dry season many young fruits fall,long before they are ripe. Elephants eat thefruits and are probably the main seed dispers-ers. It is unknown whether seeds that have

passed through elephants' guts germinatemore easily, but seeds that have not been eatenby elephants do germinate although the ger-mination rate is low. Seedlings grow slowly.After germination, growth of aerial parts stopsfor about 2 months, after which seedlings growat a rate of I cm per month, reaching an aver-age height of 25 cm after 18 months and 36 cmafter 29 months. Autrone!IQ congolensis formsarbuscular mycorrhizae with Glomeromycetespp.

Ecology Autrone!10 congo!errsisprimary evergreen rainforest, usually scat-tered, rarely abundant.

Propagation and planting The seeds withtheir thick and hard seed coat show a dor-

inaricy of 2.5-12 months before germinating.The germination rate may be up to 40%, but isoften lower; the germination rate of depulpedseeds is lowest, up to 9%. In the forest, seed-lings of AMtrone!IQ congo!errsis are often rare.Under experimental nursery conditions, thesurvival rate of seedlings was 100% after 18months. The seedlings are shade tolerant, butare classified as non-pioneer light demanders,preferring smallgaps in the forest canopy.

Management In general AMtronello congo-lensis occurs in low densities in the forest, butlocally in DR Congo 3-4 large trees per ha havebeen recorded.

Diseases and pests The seeds with theirthick and hard seed coat are not as easily at-tacked by insects as those of inoabi(Boillonel!otortspermo). The seedlings with their leafycotyledons are also less commonly eaten bybush pigs, antelopes and porcupines than thoseof inoabiwith fleshy cotyledons

Harvesting Logs may split in the centre atfelling and they also may show other defects inthe central part

Yield A large Autronello congolensis treemay yield up to 20 in3 of usable wood

Handling after harvest Logs sink in waterand cannot be transported by river. They havea good durability and can be left in the forestfor considerable time without serious degrade

Genetic resources Autronello congolensisis classified as criticalIy endangered in the2006 TUGN Red list of threatened species, withheavy exploitation for its timber being themain threat.

Prospects It will be difficult to managenatural forest in such a way that sustainableand economicalIy sufficient production of Aut-runello congolensis timber is possible. Largertrees generally occur scattered in low densitiesNatural regeneration is poor, and the low ger-mination rate and dormancy of the seeds ham-per large-scale production of seedlings fortransplanting. Moreover, growth is slow andvery long cutting cycles are probably requiredfor sustainable harvesting. This makes Autro-itello congo!errsis a tree wlth few prospects forcommercial timber production, and attentionshould concentrate on lts protection

Major references ATIBT, 1986; Bolza &Keating, 1972; Chudnoff, 1980; 01RAD For-estry Department, 2003; CTFT, 1954; Debrouxat a1. , 1998; Normand & Paquis, 1976; Richter& Danwitz, 2000; Takahashi, 1978; Wilks &Issemb6, 2000.

Other references African Regional Work-shop, 1998a; Aubr6ville, 1961; Aubr6ville,1964; Burki11, 2000; 060n, Chadenson & Hau-teville, 1980; Keay, 1989; Neuwinger, 2000;Origuene & Kuyper, 2001; Pennington, 1991;Phongphaew, 2003; Raponda-Walker & Sillans,1961; Terashima & Ichikawa, 2003.

Sources of mustration Aubr6ville, 1964;Wilks & Issemb6, 2000.


AwcENNiA 89

Occurs In

AVICENNIAGERMiNANS (L. ) L

Protologue Sp. PI. ed. 3, 2: 891 (1764).Family Avicenniaceae (APG: ACanthaceae)Synonyms Autoennio nitido Jacq. (1760),

Auicennio of neono P. Beauv. (1809).Vernacular names White mangrove, black

mangrove, olive mangrove (En). PalerUVier blanc,mangle blanc, faux palerUVier (Fr). Mangyeamare10 (Po).

Origin and geographic distribution Auicen-itto germinons is found along the AtlanticOcean, in tropical Africa from Mauritania toAngola. It is widespread along the coasts ofCentral and South America and the Caribbean

from the southern Unitedregion, occurringStates to Brazil along the Atlantic coast and toPeru along the Pacific coast.

Uses The wood is used for marine construc-tion, boat construction (especially ribs),wharves, piles, house building, furniture andimplements. It is suitable for heavy flooring,mine props, railway sleepers, joinery, vehiclebodies, handles, ladders, toys, novelties and

90 TIMBERSl

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Auicenniogerminons - wild

carving. The wood is also used as firewood andfor charcoal production. It is commonly usedfor smoking fishThe cotyledons of the seed are eaten duringfamines, but only after careful preparation toremove toxic compounds. The leaves and rootsare used to prepare a vegetable salt. The barkhas been used for tanning and for dyeing red-dish colours. Ash is added to water as a soapsubstitute. Smoke from the wood is reported todispel mosquitoes effectiveIy. The flowers are asource of nectar for honey bees.The powdered bark made into a paste is ap-plied to skin complaints and to dispelfleas andlice. Decoctions of bark and leafy twigs aretaken or added to a bath to promote childbirth.The leaves are applied as an enema to treatpiles. A root decoction is taken to treat intesti-nal problems. In tropical America the barkresin is used in traditional medicine to treattumours, diarrhoea, haemorrhage, haemor-rhoids, rheumatism, swellings, wounds andsore throat

Properties The heartwood is pale brown topinkish brown, darkening upon exposure, andrather indistinctly demarcated from the whit-ish sapwood. The grain is often interlocked,texture fine.

The wood is heavy, with a density of about 950kg/ina at 12% moisture content. It is difficult toair dry, being liable to twisting and checking.The rates of shrinkage are high, from green tooven dry about 7.1% radial and 10.2% tangen-tial. Once dry, it is stable in service. The woodis difficult to split radially, but not tangen-tially. It saws satisfactorily, but is difficult towork by hand tools. It has some tendency to

split upon nailing and screwing. The wood isvery durable, even under water, and resistantto termite and Lyetus attacks. It has been re-ported that the wood is suitable for particleboard and pulp production, but for the latterpurpose it should be mixed with wood of otherspecies that have longer fibres.The bark contains about 12.5% tannin. All ex-

tract of leafy twigs of ADicennio germinonsfrom southern Florida (United States) showedcytotoxic activity in several human cancer celllines, with the naphthoquinone 3-chlorodeoxylapacholas active compoundEthanol solutions of smoke from burningADicennio germ, nuns wood inhibited thegrowth of pathogenic bacteria and fungi, suchas ESCherichio coli, Stophylococcus oureus andSOCchoromyces cereuisiue, which supports theusefulness of the wood for smoking fish andother foods. Moreover, the smoke significantlydeterred ACroeo deeroto, a pest of sweet potato,from depositing eggs.

Description Evergreen shrub or small tomedium-sized tree up to 17(-25) in tall; rootswith many vertical breathing roots above soillevel; bole usually low-branching, up to 40(-100) cm in diameter, sometimes with smallprop roots; bark surface smooth or slightly fis-

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sured, dark grey to dark brown, inner barkyellowish; crown dense and rounded; twigsfine-hairy, terete, with 2 grooves. Leaves de-cussately opposite, simple and entire; stipulesabsent; petiole I-2.5 cm long; blade elliptical toobovate or lanceolate, 5-18 cm x 1.5-8 cm,curieate at base, obtuse to acute at apex, Ieath-ery, powdery white and short-hairy below, inI-nutely pitted, pinnately veined with 6-15 pairsof lateral veins. Inflorescence an axillary orterminal cyme, short-hairy, with flowerscrowded near top of Inflorescence. Flowers bi-sexual, slightly zygomorphic, sessile, with 3bracts up to 2.5 min long at base; sepals 5,broadly elliptical to orbicular, 4-4.5 mm long,fine-hairy outside; petals 4, fused up to half-way, c. 6.5 min long, lobes oblanceolate,spreading or reflexed, hairy, white; stamens 4,2 slightly longer than other 2, Inserted on co-rolla tube and alternating with corolla lobes;ovary superior, ellipsoid-globose, c. 2 min long,hairy, I-celled, style c. 3 mm long, stigma 2-lobed. Fruit a slightly asymmetrical, ellipsoidcapsule 2-3 cm long, with short lateral beak,leathery, short-hairy, dehiscing with 2 valves,I'Seeded. Seed without seedcoat. Seedling withepigeal germination, vrviparous; hypocotylelongated; cotyledons thick and fleshy, folded;radicle hairy.

Other botanical information AUIcennio

comprises about 8 species and occurs in alltropical regions, IOCaUy extending into the sub-tropics; 2 species are found in tropical AfricaIn the past Auicennio has been included in thefamily Verbenoceoe, but it differs in wood andleaf anatomy and seedling and pollen inorphol-ogy, and has been placed in a separate famlIyAntcennioceoe. In a recent molecular analysis itwas nested withinAconthoceoe.In the literature, Auicennio germinons plantsfrom Africa are sometimes distingulshed fromthose from Central and South America asADicenniu of neono P. Beauv. Although the twotaxa were merged in 1963 and the Africanplants have since usually been called AUIcenniogerminons, recent phytochemical and molecu-Iar studies showed significant genetic differen-tiation between the African and American

populations, and more elaborate biosystematicstudies have been recommended. A molecular

analysis showed that regional differentiationbetween the Pacific coast and Atlantic coast

populations in America was even greater thanbetween west and east Atlantic populations,suggesting at least some dispersal across theAtlantic Ocean, and providing no support for

the treatment of the African populations as aseparate species

Anatomy Wood-anatomical description (IAWAhardwood codes)Growth rings: 2: growth ring boundaries Iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(7: vessels in diagonal and/or radial pattern);(10: vessels in radial multiples of 4 or morecommon); 13: simple perforation plates; 22intervessel pits alternate; (23: shape of alter-nate pits polygonal); 24: intervesselpits minute(S 4 prn); 30: vessel-ray pits with distinct bor-ders; similar to intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumlna 50-100 pin;47: 5-20 vessels per square minimetre; 48: 20-40 vessels per square minimetre. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;69: fibres thin- to thick-walled; 70: fibres verythick-walled. Axial parenchyma: 78: axial pa-Tenchyma scanty paratrachea1; 79: axial paren-chyma vasicentric; 85: axial parenchyma bandsmore than three cells wide; 86: axial paren-chyma in narrow bands or lines up to threecells wide; (91: two cells per parenchymastrand); 92: four (3-4) cells per parenchymastrand; 93: eight (5-8) cells per parenchymastrand. Rays: 97: ray width I-3 cells; 105: allray cells upright and/or square; 109: rays withprocumbent, square and upright cells mixedthroughout the ray; 1/5: 4-12 rays per min;1/6: Z 12 rays per mm. Secretory elements andcambial variants: 133: included phloem, con-centric. Mineral inclusions: 152: crystals ofother shapes (mostly small); 154: more thanone crystal of about the same size per cell orchamber

(P. 06tienne & P. E. Gasson)Growth and development In experiments

in Ghana, seedlings reached a height of up to26 cm at 4 months after planting. Under opti-inal growing conditions, trees may grow 60cm/year in height. The leaves excrete excesssalt through pores, and salt crystals are coin-monly present on the leaves. The minute hairsthat cover the lower surface of the leaf play arole in the water balance, but more studies arestill needed to understand the exact processes.In each cluster of flowers there is often only a

single open flower at a time. The flowers areprotandrous and attract short-tongued insectssuch as bees, which receive poUen on theirbacks while correcting nectar. After some shriv-elling of the stamens, the 2 lobes of the stigmaexpand, enabling pollination by insects that

AVICENNIA 91

92 TIMBERSl

previously visited a flower in the male stage.The seeds start germinating while still at-tached to the tree, but the embryo stays withinthe fruit until fruit fall

Ecology Auicennio germinons occurs in man-grove vegetation in tidal estuaries, often ongraveny or sandy soils. It is often common,especially in secondary vegetation types, some-times forming almost pure stands or mixedstands with Rhizophoro spp. and Loguncu!onurocemoso (L. ) CF. Gaertn. It is most commonon the landward side of mangrove swamps, insheltered mud fiats of tidal zones and at the

mouths of rivers. In mangroves in Nigeria,Auicenn, o germinons dominates, together withthe introduced palm Nypo Iruticons Wurmb, inlocalities where calcium is in plentiful supplyand where soils are less acidic than in localities

dominated by Rhizophoro spp. In Senegal andGambia Auicennio germirtons often grows indense stands in mangrove localities where thesoil has become drier and sandier in coinpari-son to localities where the mud is most recentand least consolidated and which are doini-

nated by Rhizophoro rocemoso G. Mey. It mayalso invade abandoned paddy fields affected bytides. In Gambia ADicennio germtnons is saidto be more tolerant to sannity than Rhizophorospp. , probably because it is able to excrete saltADicennio germinons occurs in regions with amean annual temperature of 25-27'C andmean annual rainfallof500-2000 min.

Propagation and planting Natural regen-eration is often abundant, and wildlings can becollected to serve as planting stock. Experi-merits with seedlings resulted in approxi-mately 90% survival. The seedlings grow bestwhen they are in contact with fresh waterFruits with germinating seeds may float in saltwater for one year without rooting. They arevery susceptible to desiccation

Management In many mangrove regions,the potential of ADicennio germinons is consid-ered limited and other mangrove species likeRhi20phoro spp. are often more highly valuedfor timber, firewood, charcoal, dye and tanninproduction. Although mangroves are oftenheavily exploited, ADicennio germinons is oftenleft. In Gambia a standing volume of 106 mathahas been recorded, corresponding to a produc-tivity of about 3.5 in3ftia per year. In mixedADicenn, o1Rhizophoro mangrove the standingvolume was 29 in3ftja

Diseases and pests The germinating seedsare often attacked by crabs and snails, and theleaves of trees can also be damaged by crabs

Genetic resources ADicennio germinons isone of the most common species in mangrovevegetations, being a pioneer species with greatpower of natural regeneration. As such, it doesnot seem liable to genetic erosion. In manyregions of tropical Africa, however, mangroveshave been or are being converted into agricul-tural land, especially into rice fields, or aresubJect to overharvesting for timber or fuel,and this may eventually affect the genetic di-versity of Antcennio germinuns. Locally thepopulations are also threatened by petroleumproduction and resulting pollution, as in someareas in Nigeria.

Prospects Mangroves are socio-economicallyimportant ecosystems for the inhabitants ofcoastal regions, but they are under high pres-sure worldwide. As one of the main constitu-ents of mangrove vegetations in Africa, ADicen-rim germinons deserves protection. Its role intimber production in tropical Africa shouldtherefore be limited to local applications forheavy construction, with emphasis on sustain-ability. Moreover, the holes are often too smalland of too poor shape to be interesting forcommercial timber exploitation

Major references Berhaut, 1971; Bolza &Keating, 1972; Burki11, 1985; Dodd, ofzal Rafti& Bousquet Melou, 2000; Dodd at a1. , 2002;Normand & Paquis, 1976; Tetteh, 1999;Tomlinson, 1986; Ukpong, 2000; Viniers,1975a

Other references Abbiw, 1990; Ako6gni-nou, van der Burg & van der Maesen, 2006;Asita & Campbe11, 1990; Aubr6ville, 1959d;Bousquet Melou & Fauve1, 1998; Chua, 1998;Coinp6re, 1963; Duke, 1991; Fagbami, Udo &Odu, 1988; Hawthorne & Jongkind, 2006;Hubbard, 1981; InsideWood, undated; Jones ata1. , 2005; Keay, 1989; Neuwinger, 2000; Nor-mand, 1960; 0gbalu at a1. , 2004; Raponda-Walker & Sillans, 1961; Saville & Fox, 1967;Schwarzbach & MCDade, 2002; Viniers, 1973a.

Sources of illustration Hepper, 1963.Authors N. S. A1varez Cruz

AVICENNLAMARINA (Forssk. ) Vierh.

Protologue Denkschr. KaiserI. Akad. WISS. ,Math. -Naturwiss. in. 71: 435 (1907)

Family Avicenniaceae (APG: ACanthaceae)Chromosome number2n=36Synonyms ADicennio officinglis auct. non L.Vernacular names White mangrove, grey

mangrove, olive mangrove (En). PalerUVier

blanc (Fr). Mangue branco, mangue nero, sal-gueiro (Po). Mchu, rutsu, nsusi, nsuti(Sw)

Origin and geographic distribution ADicen-itto morino is extremely widespread along thecoasts of eastern Africa, islands of the IndianOcean, tropical Asia, Australia, New Zealandand islands of the Pacific Ocean to Fiji. Intropical Africa it occurs from Egypt to SouthAfrica, and in most islands of the IndianOcean

Uses The wood is used for poles in housebuilding, for boat construction, especially forthe ribs, for furniture and handles, and tomake beehives. The branches serve as stakesfor fences. The wood is also used as firewoodand for charcoal production; it is especiallyused for lime burningThe bark has been used for tanning and fordyeing reddish and brownish colours. In Mo-zambique the bark of ADicennio morino wasformerly utilized commercially for tanning, butthe tannin content is rather low. The smoke ofburning wood is considered very efficient as amosquito repellent. The cotyledons of the seedare occasionally eaten, but may contain toxiccompounds. The foliage serves as fodder forlivestock. Leafy branches are used for makingfish kraals. The tree is useful for preventingcoastal erosion and as a windbreak. Honeybees collect nectar from the flowersThe resin from the bark is used to treat snakebites and to remove the placenta after child-birth. In Madagascar a leaf decoction has beenused as antidote after eating poisoned fish,whereas leaf and bark decoctions are appliedagainst scabies. In traditional medicine in AUS-tralia, leaves, young shoots and bark are ap-plied as an anodyne, and wood ash to treat skin

complaintsProperties The wood is greyish to yellowish,

with fine and even texture. It is heavy anddurable, and the wood properties are coinpara-ble to those of ADicenniogerminons (L. ) L.Several indoid glucosides and navonoids havebeen isolated from the aerial parts of AUIcenniomanner. Several naphthoquinone derivativeshave been isolated from the twigs, and some ofthese (avicequinone A, avicequinone C, steno-carpoquinone B, avicennone D and avicennoneE) showed strong antiproliferative and moder-ate cytotoxic activities as well as antibacterialeffects

The leaves contain 10.5% crude protein, 21.5%crude fibre, 3.5% ether extract and 21% ashThe in-vitro organic matter digestIbility is60.5%, indicating that the leaves can be usedas a forage to meet maintenance requirementsof camels. The bark contains 0.6-2.2% tannins.

ADicennio morino tolerates heavy metalsin thesoil very well. The roots may be employed as abiological indicator of environmental exposureto copper, lead and zinc

Description Evergreen shrub or smalltreeup to LOG15) in tall; roots with many verticalbreathing roots above soillevel; bole usuallylow-branching, up to 40(-50) cm in diameter,sometimes with small prop roots; bark surfacesmooth to slightly fissured or flaky, brownishor yellowish green, inner bark greenish; crowndense and rounded; twigs fine-hairy, slightlyangular. Leaves decussately opposite, simpleand entire; stipules absent; petiole 0.5-Tel. 5)cm long; blade elliptical to ovate-elliptical or

1.5-5 cm,elliptical-lanceolate, 3-12 cm x

curieate at base, acute or acuminate at apex,leathery, minutely whitish halry below, some-times minutely pitted, pinnately veined with8-15 pairs of lateral veins. Inflorescence anaxillary or terminal head-like cyme, short-hairy. Flowers bisexual, regular, sessile, with 3bracts up to 4 min long at base; sepals 5, ovateto elliptical or nearly orbicular, 3.5-4 mm long,hairy outside; petals 4, fused halfway, 4.5-7mm long, lobes ovate, hairy outside, yellow toorange, becoming blackish after flowering;stamens 4, inserted on corolla tube and alter-nating with corolla lobes, filaments very short;ovary superior, conical, c. 2.5 mm long, hairy inupper part, I-celled, style c. I min long, 2-lobed. Fruit a slightly asymmetrical, broadlyellipsoid to ovoid capsule I-3 cm long, leathery,scaly hairy, yellowish green, dehiscing with 2valves, I'Seeded. Seed compressed. Seedlingwith epigealgermination, viviparous; hypocotyl

AVICENNiA 93

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nate pits polygonal); 24: intervesselpits minute(S 4 F1m); 30: vessel-ray pits with distinct bor-ders; similar to intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumina 50-100 prn;42: mean tangential diameter of vessellumina100-200 prn; 48: 20-40 vessels per square inn-limetre; 58: gums and other deposits in heart-wood vessels. Tracheids and fibres: 61: fibres

with simple to minutely bordered pits; 66: non-septate fibres present; 69: fibres thin- to thick-walled; 70: fibres very thick-walled. Axial pa-renchyma: 76: axial parenchyma diffuse; 78axial parenchyma scanty paratrachea1; 79axial parenchyma vasicentric; (80: axial paren-chyma alitorm); (81: axial parenchyma lozenge-anform); 91: two cells per parenchyma strand;92: four (3-4) cells per parenchyma strand; 93:eight (5-8) cells per parenchyma strand. Rays(97: ray width I-3 cells); (98: larger rays coin-monly 4- to 10-senate); 105: all ray cells up-right and/or square; 109: rays with procum-bent, square and upright cells mixed through-out the ray; 1/5: 4-12 rays per min. Secretoryelements and cambial variants: 133: included

phloem, concentric. Mineral inclusions: (144druses present); (145: druses in ray paren-chyma cells); 152: crystals of other shapes(mostly small); 154: more than one crystal ofaboutthe same size per cellor chamber(P. 06tienne & P. E. Gasson)

Growth and development The leaves ex-crete excess salt through pores, and salt crys-tals are commonly present on the leaves. Theminute hairs that cover the lower surface of

the leaf play a role in the water balance, butmore studies are stillneeded to understand the

exact processes. In southern Africa trees flowerfrom August to October, in Kenya in October-November. Individual flowers are open for 2-5days. The flowers are protandrous, and thismakes self-pollination unlikely, but pollinationfrom flowers of the same plantis likely. Partialself-compatibility has been demonstrated bybagging flowers. The flowers attract short-tongued insects such as bees, which receivepollen on their abdomen while collecting nec-tar. After the stamens have turned black, the 2lobes of the stigma expand, enabling pollina-tion by insects that previously visited a flowerin the male stage. Fruits take 2-3 months tomature in regions around the equator, but upto 10 months in temperate climates such as inNew Zealand. However, in general the wholeprocess from bud initiation to abscission ofmature fruit is completed within a year. In

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elongated; cotyledons thick and fleshy, folded;radicle usually glabrous, with short hairy col-Iar

Other botanical information Auicenniocomprises about 8 species and occurs in alltropical regions, locally extending into the sub-tropics; 2 species are found in tropical Africa.In the past ADicennio has been included in thefamily Verbenoceoe, but it differs in wood andleaf anatomy and seedling and pollen inorphol-ogy, and has been placed in a separate familyADicennioceoe. In a recent molecular analysis itwas nested withinAconthoceoe

ADicennio morinu is variable over its largearea of distribution, and 3 varieties (also con-sidered as subspecies) have been distinguishedOnly one of these (var. marina) occurs in tropi-cal Africa. However, the varieties show muchinorphologicaloverlap

Anatomy Wood-anatointcal description OAWAhardwood codes)Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(7: vessels in diagonal and/or radial pattern);(10: vessels in radial multiples of 4 or morecommon); 13: simple perforation plates; 22intervessel pits alternate; (23: shape of alter-

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south-eastern Australia only 3% of floral budsdeveloped into viable seeds, with insect attackson developing fruits as the main cause of inor-tanty. The average annual supply of viableseeds per tree is approximately 250. The seedsstart germinating while still attached to thetree, but the embryo stays within the fruit un-til fruit fall. Fruits with germinating seedsmay float in salt water for 5 months withoutlosing their viahinty. Most of these strandwithin I kin of the mother tree, and very fewwere observed to disperse more than 10 kin.

Ecology AUIcennio indrino occurs ln man-grove vegetation, usually at the landward sideof the mangrove, where it is often doinLnant or

stands. It has a wideOccurs even In pure

physiological tolerance to satinity, being able tosurvive in fresh stagnant water as well as inseasonalIy dry conditions with very high sann-ity; it also has a wide tolerance to intertidalposition and to temperature. It can be foundacross the entire intertidal profile above meansea level, and occupies offshore reef lagoons aswell as sandy or rocky sheltered embayments.In KenyaAuicennio morino typically displays adouble zoriation pattern, occurring on thecoastal edge of mangroves, often as a tree up to15 in tan, as well as on the inland side, often asa shrub. In South Africa tree height frequentlydecreases from up to 10 in in the fringe zone toless than 1.5 in inland at a slightly higher ele-vation, as a result of hydro-edaphic factorscontributing to high soil satinities, low waterpotentials, water stress and ion imbalancewithin tissues in the inland sites. Auicenniomorino is often a pioneer in sandy habitats,but may also invade mud flats. It tolerates soilswith a pH of 6 to 8.5. It is intoIerant of shadeAuicennio morino occurs in regions with amean annual temperature of 17-26'C andmean annual rainfallof (200-)1000-4500 min

Propagation and planting Natural regen-eration is often abundant, and wildlings can becollected to serve as planting stock. The seedsare recalcitrant. They are very susceptible todesiccation. There is progressive deteriorationof the internal tissues of the seeds associatedwith fungal infection during hydrated storageExperiments with seedlings resulted in ap-proximateIy 90% survival. Young seedlingsgrow best when they are in contact with freshwater, but growth diminishes soon under theseconditions and is best in water with 10-50% offull seawater sannity for older seedlings. Thereappears to be no restriction to establishment ofseedlings within mangrove stands, but re-

CTuitment to the sapling stage appears to berestricted by light and sediment resourcesPropagation by air layering and root suckershas been successful.

Management In many mangrove regions,the potential of Antcennio inuring is consideredlimited and other mangrove species such asRhizophoro spp. are often more highly valuedfor timber, firewood, charcoal, dye and tanninproduction. Although mangroves are oftenheavily exploited, ADicennio morino is oftenleft. Trees suffer little from removal of branch-wood because they can resprout rapidly frombuds along the stems.

Diseases and pests In Australia the leaves,flowers and seedlings are affected by leafblightcaused by Alternorio alternoto.Crabs consume the propagules and may be themain reason for the absence of Auicennio ina.ring in certain areas. The mangrove decapodcrab Neosormottum memerti, which is verycommon in the ADicennio indrino zone alongthe East African coast, feeds heavily on seed-lings and fallen leaves

Genetic resources ADicennio mar, rid ls oneof the most common species in mangrove vege-tations, being a pioneer species with greatpower of natural regeneration. As such, it doesnot seem liable to genetic erosion. However, inmany regions of tropical Africa mangroveshave been under much pressure because ofclearing for other land uses and exploitation forfuel.

In populations in tropical Asia and Australia, itwas shown that a very high level of geneticstructure and inbreeding exists and that popu-Iations are functioning as Independent evolu-tionary units more than as components of ametapopulation system connected by gene flow.This makes peripheral populations likely todevelop local adaptations and therefore to be ofparticular interest for conservatlon strategiesas well as for adaptation to possible futureenvironmental changes.

Prospects Mangroves are socio-economicallyimportant ecosystems for the inhabitants ofcoastal regions, but they are under high pres-sure worldwide. As one of the main constitu-ents of mangrove vegetations in Africa, Auicen-itto inuring deserves protection. Its role in fu-ture timber production in tropical Africa seemsvery limited because the boles are often toosmall and of too poor shape to be interestingfor commercial timber exploitation. It can beplanted for mangrove restoration, because itexhibits wide physiological tolerance and it

AVICENNIA 95

96 TIMBERSl

creates a suitable environment for other man-

grove species after it has become well estab-lished.

Major references Amaud, 2006; Beln at al. ,1996; Chua, 1998; Coates Palgrave, 1983; Hanat a1. , 2007; Maundu & Tengnds, 2005; Naidoo,2006; Palmer & Pitman, 1972-1974; Tomlinson,1986; Verdcourt, 1992

Other references A1 Dosan, 2001; Beentje,1994; Boiteau, Bolteau & NIOrge-Boiteau,1999; Bousquet Melou & Fauve1, 1998;Chandrashekar & Ball, 1980; Clarke, 1992;Clarke, 1993; Clarke & Allaway, 1993; Clarke& Myerscough, 1991; CIOugh, 1984; Dahdouh-Guebas at a1. , 1997; Decary, 1946; Downton,1982; Duke, 1991; InsideWood, undated;Moldenke, 1956; Neuwinger, 2000; Schatz,2001; Schwarzbach & MCDade, 2002; Thulin,2006; vanWyk & Geneke, 2000

Sources of illustration Bein at a1. , 1996;Maundu & Tengnas, 2005; Verdeourt, 1992

Authors N. S. A1varez Cruz

struction. Its uniform reddish brown colour and

fine grain make it popular for veneer, furni-ture, decorative uses and flooringThe fruit pulp is eaten fresh although it con-tains latex. The seed kernelcontains a fatty oilthat is used in cooking, for cosmetic purposesand in traditional medicine. The oilis appliedas a pomade to the hair and used in soapmanufacture, and it is applied externalIy totreat rheumatic pains. The residues of oil ex-traction are sometimes used as fish poisonBark decoctions are used to treat kidney prob-Iems, toothache, rachitis, vaginal infectionsand affections of the respiratory and digestivetracts. The tree has several ritual uses.

Production and international trade Coin-

mercial exploitation of inoabi started in theinId 1960s. In 1960 the total trade of logsamounted to 3000 in3, in 1973 already to47,300 ina and additionally almost 1000 in3 ofprocessed wood. In 2003 Garnero0n exported16,000 ms processed wood and Gabon 54,000in 310gs.Moabi bark is regularly sold in markets inGarnero0n for medicinal purposes; in 2000 theamounttraded was estimated at 3.2 t. The seed

oilis sold on local markets, but no statistics areavailable

Properties Heartwood pinkish brown to red-dish brown, fairly distinctly demarcated fromthe pinkish white or greyish brown sapwood.The grain is straight to slightly interlocked,texture fine. Wood with satiny lustre on quar-ter-sawn surfaces. Growth rings more or lessdistinct

The density is 820-940 kg/in3 at 12% moisturecontent. The timber dries slowly, but with careit seasons without checking and warping. Theshrinkage rates are moderate, from green tooven dry 5.8-6.5% radial and 6.9-8.6% tangen-tial. Slow drying to a moisture content of 10-12% is needed for use in joinery or carpentryOnce dry, the wood is stable.At 12% moisture content, the modulus of rup-ture is 148-218 N/mm2, modulus of elasticity15,100 N/mm2, compression parallel to grain57-84 N/min2, cleavage 1324 N/min andChalais-Meudon side hardness 5.6-7.0.

The wood is difficult to work due to the pres-ence of silica (0.2-0.3%) and because it is hardStellite-tipped sawteeth are needed. The woodglues, nails and finishes well, and has goodsteam-bending properties. Painting and var-nishing with the principal industrial productsdo not cause problems. After steaming for 4360 hours, rotary peeling and slicing give veneer

BAILLONELLATOXISPERMAPierre

Protologue Not. bot. L 14 (1890).Family SapotaceaeChromosome number2n=24

Synonyms Mtmusops digue Eng1. (1897)Vernacular names Moabi, African pearwood

(En). Mochi(Fr). Muabi(Po)Origin and geographic distribution Moabi

occurs from southern Nigeria to Gabon, Congoand south-western DR Congo

Uses Moabi wood is used for joinery and car-pentry in exterior applications, e. g. for doors,windows, garden furniture and marine con-

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and plywood of satisfactory qualityMoabi wood is durable. It is resistant to fungi,Lyetus beetles and termites. It is fairly resis-tant to marine borers. Impregnation by pre-servatives is difficult or impossibleThe allelopathic compound 3-hydroxyuridinehas been isolated from leaves, stems and rootsof inoabi; it inhibited the growth of seedlings ofsome test plants

Adulterations and substitutes Moabiwood

can be confused with that of other Sopotoceoespecies having heavy and reddish brown wood,particularly inukulungu (Autronel!o congo!erusis (De Wild. ) A. Chev. ), makore (Tieghemel!oheekelii (Achev. ) Roberty) and douka(negheme!IQ Girtcono Pierre). The fatty seedoilis similar to that of neghemello and sheabutter tree (Vitellorto parodo"o CF. Gaertn. )and used for similar purposes

Description Very large tree up to 60(-70) intall; hole up to 300(-500) cm in diameter,straight and cylindrical, sometimes swollen inlower part, reaching up to 30 in to the firstbranches, without buttresses; bark 4-5 cmthick, surface red-brown to dark grey, deeplylongitudinalIy furrowed, red-brown in outerpart of section, yellow-pink in inner part, littlefibrous, exuding a sticky latex; crown urn-

brella-shaped, very large, up to 50 in In diame-ter, heavy branches spreading and sinuate;ultimate branches very thick, with numerousscars of fallen leaves. Leaves arranged spiralIyin tufts at the ends of branches, simple; stip-ules lanceolate, large, persistent; petiole 3-4cm long, slender; blade narrowly obovate, 15-30 cm x 5-10 cm, curieate at base, shortlyacuminate at apex, margin entire, initiallyreddish hairy below but glabrescent, lateralveins numerous, distinct, curving and joinednear leaf margin. Flowers in dense fascicles atthe ends of branches, bisexual, regular; pedicel2-3 cm long, pubescent; calyx with 2 whorls of4 lobes c. I cm long, pubescent outside; corollawith c. 2.5 mm long tube and 8 lobes c. 4 minlong, each lobe with 2 large lateral segments c5.5 mm long, creamy white; stamens 8, In-serted on the corolla tube in front of the corollalobes, free, filaments short, 8 larger staininodesalternating with the stamens; ovary superior,long-hairy, 8-celled, each cell with I ovule,style short. Fruit a large, globose, smooth berry5-8 cm in diameter, grey-green, becominggreenish yellow when ripe, containing I-2(-3)seeds in a pale yellowish pulp. Seeds ellipsoid,slightly lateralIy compressed, c. 4 cm long,testa thin, smooth and shining in dorsal part,rough and bullate in ventral part (scar); en-dosperm thin or absent. Seedling with epigealgermination, hypocotyl short, 0.5-1.5 cm long,epicoty1 15-26 cm long, reddish to greyishbrown hairy, cotyledons thick, sessile, c. 4 cm xICm, green

Other botanical information Boillonellocomprises a single species and resemblesneghemellu and Mimusops, the first differingin the absence of stipules and its thicker seedcoat, the latter in its seed with small and basal

BAILLONELLA 97

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Anatomy Wood-anatointcal description (IAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; (7vessels in diagonal and/or radial pattern); (10:vessels in radial multiples of 4 or more coin-mon); 13: simple perforation plates; 22: in-tervessel pits alternate; (23: shape of alternatepits polygonal); 26: intervessel pits medium (7-10 pin); 27: intervessel pits large G 10 prn); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; 31: vessel-ray pits with much re-duced borders to apparently simple: pitsrounded or angular; 32: vessel-ray pits with

3

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98 TIMBERSl

much reduced borders to apparently simple:pits horizontal(scalariform, gash-like) to verti-cal (palisade); 33: vessel-ray pits of two distinctsizes or types in the same ray cell; 42: meantangential diameter of vessellumina 100-200urn; 47: 5-20 vessels per square minimetre; 56:tyloses common. Tracheids and fibres: 61: fi-bres with simple to minutely bordered pits; 66:non-septate fibres present; 69: fibres thin- tothick-walled. Axial parenchyma: (77: axial pa-renchyma diffuse-in-aggregates); 86: axial pa-renchyma in narrow bands or lines up to threecells wide; 87: axial parenchyma reticulate; 93:eight (5-8) cells per parenchyma strand; (94:over eight cells per parenchyma strand). Rays:97: ray width I-3 cells; (100: rays with multis-eriate portion(s) as wide as uniseriate por-tions); 106: body ray cells procumbent with onerow of upright and/or square marginal cells;107: body ray cells procumbent with mostly 2-4rows of upright and/or square marginal cells;108: body ray cells procumbent with over 4rows of upright and/or square marginal cells;1/5: 4-12 rays per min. Mineral inclusions159: silica bodies present; 160: silica bodies inray cells; (161: silica bodies in axial paren-chyma cells)re. 06tienne & E. A. Wheeler)

Growth and development Germinationtakes I-4 weeks. Shade is required for germi-nation, and seeds germinate in abundance un-der or close to mother trees. However, youngplants of I in tall are rare as a result of preda-tion by animals, mainly elephants, and thesurvival rate of inoabi seedlings after 18months is practically nil under the mother treeand very low (less than 2%) elsewhere. In for-est undergrowth seedlings grow 4-5 cm/yearThey respond rapidly to opening of the forestcover and direct sunlight, and under these cir-cumstances may grow up to 40 cm/year. Innatural forest, inoabi trees of 10 cm in stem

annual diameterdiameter have an average

growth of I mm. When the tree bole has at-tamed I in in diameter and the crowns have

reached or passed the forest canopy, diametergrowth is up to 9.5 min/year. It has been esti-mated from growth rings and Ci4 dating that atree with a hole diameter of I in was 260 yearsold and one with a diameter of 2.8 in 600-700

years' In an 11-year-old plantation in Ekouk(Gabon) trees grown in fullsun showed a meanannual height growth of 130 cm and underpartial shade 165 cm. A dense (570 stemsftia)plantation of 67 years old near Libreville had amean annual diameter Increment of 3.5 min,

with the most vigorous tree reaching 9 mm; theannual volume increment was estimated at 4.2

matha, with an average clear hole of 15 in and23% of the trees having a straight and cylindri-calbole

The tree starts flowering at an age of 50-70years, when it has reached the light in the up-per storey of the forest, but regular fructifica-tion starts still later, when the hole hasreached 70 cm in diameter. Fruits are producedannually, but mass production occurs once in 3years. About 6000 seeds per tree are producedwithin a cycle of 3 years; a single tree produced2460 kg of fruits, including 327 kg of seed ker-nels.

In Cameroon flowering starts at the end of thelong dry season and beginning of the shortrainy season, in February to April. It is accom-panied by complete defoliation of the tree;when the tree does not flower, it maintainspart of its foliage. Fruits ripen just before thestart of the long rainy season, from June toAugust. The fruiting season lasts up to 4 weeksfor an individual tree, but may last 3 monthsfor the whole population. In Gabon inoabifruits become ripe from December to FebruaryElephants are the main seed dispersers. Theyeat the fruits, and intact seeds can be found inthe droppings. It is not necessary for the seedsto pass through the intestines for germination,but it does accelerate germination slightly.Other fruit-eating mammals such as giantpouched rats (Cricetomys emini) and monkeysmay also disperse seeds, as wellas man.

Ecology Moabioccurs in primary rain forestin humid and warm climates, wlth a mean an-rinaltemperature o123-26'C and mean annualrainfall of 1500-3000 mm. It usually occursscattered, with about I adult tree per 20 ha,but occasionally loose groups of 5-50 trees arefound. Some forests are rich in inoabitrees, e. g.Dja forest in Cameroon, with on average 0.6tree over 70 cm in hole diameter per ha. In Djaforest there is a maximum number of trees in

the diameter class 10-40 cm and a secondarymaximum at 160-190 cm. At the northern 11m-

its of the primary Congolean rain forest inCameroon large inoabitrees are abundant, butregeneration is absent. MDabican be found onallsoiltypes except on marshy soil

Propagation and planting The germina-tion capacity offresh seeds is high (> 85%), butdrops rapidly to 5% after 6 weeks of storagedue to rapid degradation of the fatty acidsHowever, fresh seeds stored at 4'C and humidconditions still showed a germination rate of

about 50% after 11 months.

In the nursery the lower halfofthe seed is bur-ied, with the scar pointing downwards. Themortality of seedlings in the nursery Is verylow. Partial shading is favourable for seedlingdevelopment. Under optimal nursery condi-tions seedlings may reach 35 cm in height after6 weeks, 45 cm after 3 months and 1.5 in after18 months. The seedling develops a taproot andit is recommended to sow into polythene potsAt the beginning of the short rainy season InCameroon 4-5-month-old seedlings are plantedinto the field under the light shade of about 10retained adult trees per ha. The shade shouldbe removed gradually when a reduction in di-ameter growth is observed

Management At present, the only manage-merit measure for natural forest Is the mainte-

nance of a minimum diameter limit for exploi-tation, i. e. 70 cm in Gabon, 80 cm in Congo and100 cm in Gamero0n. The Pygmy people highlyvalue large inoabi trees for their fruits andconflicts arose in Garnero0n between the Pyg-inIes and companies entitled by the govern-merit to exploit the forest for timber. This hasresulted in a law endowing the village chiefwith the power to authorize the cutting of ainoabitree within 5 kin distance from a village

Diseases and pests The sapwood is suscep-tible to attack by insects and fungi. In felledlogs some of the insects affecting the sapwood(e. g. Plotypes and SCOlytes spp. ) may reach theheartwood

Fruits still attached to the tree are attacked bylarvae of the beetle Corpophilus sp. and themoth Mussidio sp. When they reach the em-bryo, the seed will not germinate; otherwisethe development of the seedling may bestrongly retarded. Bush plgs and porcupineseat the seeds, and bush pigs and antelopesdestroy seedlings while eating the fleshy coty-ledons and young shoots

Harvesting In Garnero0n commercial exploit-tation of inoabiimplies the cutting of 90% oftrees above 100 cm in diameter, leaving onlypoorly shaped trees. It has been estimated thatafter 300 years 84% of the harvested volume isre-established and after 500 years 92%. Sus-tainable production of inoabi in natural forestis therefore not possible. Cutting cycles of 30years, as often practised, threaten inoabi popu-Iations seriously.In Cameroon fruit collection starts in July orAugust. Pygmy families move to localitieswhere clusters of trees are found and staythere for 2-8 weeks. Locally in Gabon people

move into temporary camps in January to Feb-when the inoabitrees are fruiting. Theruary,

collected fruits are depulped and the seed ker-nels dried. After heating the oil is gathered bypressing with a traditional wooden press. Theoilis used domestically or sold. It can be storedfor I-2 years

Yield The commercial wood volume of indi-vidualtrees is about 3 ing in trees with a holediameter of 50 cm to about 26 ina in trees witha hole diameter of 1.8 in

In 1992, which was a very good year for fruitproduction, the yield of oil was estimated at135-165 IPer adult tree, which was equivalentto Us$ 165-200. The price on markets inYaound6 and Douala (Cameroon)is more thanUs$ 3 perlbecause demand exceeds supply

Handling after harvest The sawdust maycause irritation to throat, nose and eyes, andalso dermatitis

Genetic resources According to the IUCNclassification, inoabiis vulnerable because ofoverexploitation and habitat degradation. Thelow growth rate and often poor natural regen-eration make it even more liable to geneticerosion or even extinction in forest liable totimber exploitation

Prospects Proper propagation and plantingtechniques have been developed for inoabi andconsequently it is possible to include the spe-

plantation forestry. Although theCies In

growth of planted trees can be faster than innatural forest, timber exploitation can not beexpected within a century. Moabi trees areimportant for local populations because of theirvaluable fruit, and those close to villagesshould be protected, taking into account thatthey start fruiting many decades after germi-nation. Moabi deserves protection also becauseit is the largest and perhaps most linpressivetree of the African forest

Major references Aubr6ville, 1964; Burkill,2000; Chudnoff, 1980; CTFT, 1956; CTFT,1976b; Debroux, 1998; Debroux at a1. , 1998;Mapaga, Ingueza & Louppe, 2002; Richter &Danwitz, 2000; Schneemann, 1995

Other references Agom & Ogar, 1994;ATIBT, 1986; Aubr6ville, 1961; Berti at al. ,1982; Bierna, 1963; Bokdam, 1977; Brown,1977; Fouquet, 1984; Inc. 1991; InsideWood,undated; Koumba Zaou at a1. , 1998; Monet ata1. , 1995; Moss, 1995; Normand & Paquis,1976; Objgashiet a1. , 1989; Raponda-Walker &Sillans, 1961; Takahashi, 1978; Vivien &Faure, 1988c; White & Ahemathy, 1997; Wilks& Issemb6, 2000.

BinLLONELIA 99

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100 TIMBERSl

Sources of illustration Aubr6ville, 1961;White &Ahemathy, 1997

Authors D. Louppe

BAMBUSAVULGARIS Schrad. ex J. C. Weridl.

Protologue Coll. PI. 2(2): 26, t. 47 (1810).Family Poaceae (Gramineae)Chromosome number2n= 72

Vernacular names Common bamboo (En)Barnbou, bambou de Chine (Fr). Bambu vulgar(Po). Mwanzi, inwazi(Sw)

Origin and geographic distribution Born-buso uulgoris originated in the Old World,probably in tropical Asia. It is only known fromcultivation, although escaped and naturalizedpopulations exist. Common bamboo is the mostwidely grown bamboo throughout the tropicsand subtropics. In tropical Africa it is widelygrown and also occurs subspontaneously

Uses The sterns are used for construction

and fencing, but also for tool handles, furni-ture, handicrafts, stakes, pots, weapons, Irriga-tion pipes and bows for fishing nets. In Kumasi(Ghana) the sterns are used in construction astemporary support material. A more recentdevelopment in Ghana is that bamboo stemsare used for flooring, panelling and furnitureIn Tanzania the stems serve as poles to sup-port banana plants and they are used as distil-Iation pipes for liquor making. Split stems aremade into baskets, fences, roofs and roof tiles.The stems are also used as fuel. They providegood quality pulp and are used for paper inak-ing, e. g. in India. The young shoots are edible.In Rodrigues onauntius) a decoction of thegrowing point of the plant mixed with roots of

Job's tears (Cotx IOCrymo:jobi L. ) gives a re-freshing drink. The leaves are browsed bygoats. Stem sheaths are used as covers for bee-hives. Bombuso uulgoris is planted for erosioncontrol and as an ornamental. In Sierra Leone

it is planted to mark boundaries. In Nigeria adrink of macerated leaves is taken against ve-nereal diseases. In DR Congo the leaves formpart of preparations used for treatment of mea-SIes


worldwide production and trade of Bombusouulgoris products is considerable, but no statis-tics are available

Properties For both green-stem and yellow-stem cultivars the density of the stem wantsabout 0.63 g/cm3 at 12% moisture content.Shrinkage from green to 11.3% moisture con-tent is 9.7-14.0% radial and 6.0-11.9% tangen-tial. For green-stem cultivars at 17% moisturecontent, the modulus of rupture is 84 N/mm2,compression parallel to grain 25 N/mm2 andshear 6.6 N/mm2. For yellow-stem cultivars at16% moisture contents the modulus of ruptureis 86 N/mm2, compression parallel to grain 32N/min2 and shear 4.3 N/min2.

The working and machining properties of thestems are poor. The stems are not straight, noteasy to split, and inflexible, but they are thick-walled and initially strong.Among bamboos, the carbohydrate (starch andsugar) content of the stems mainly determinesthe susceptibility to insects and fungi. AsBombuso uu!80ns has a high starch content, itis more easily attacked than other bamboo spe-cies, such as Dendrocolomus gigonteus Wall. exMunro. The stems are very susceptible to at-tack by powder-post beetles. Preservation toprotect against these beetles and other biologi-cal degradation is of vital importance for long-term service. The inner and outer walls of

bamboo stems are rather resistant to penetra.tion by preservative liquids, and the uptake ismainly restricted to the ends of the stems.Stem fibres from Bombuso uulgoris from tropi-calAfrica (C6te d'Ivoire, Garnero0n, Gabon andCongo) were on average 2.6-2.9 min long, witha diameter of 17.1-20.0 prn and a lumen di-ameter of 3.5-7.3 prn. The average chemicalcomposition was: cellulose 41-44%, pentosans21-23%, Iignin 26-28%, ash 1.7-1.9%, silica0.6-0.7%. The solubility in hot water was 3-5%, in alcohol-benzene 2-3%, in I% NaOH 20-22%. Papermaking studies have shown thatpulp obtained from the stems of Bombuso uu!-goris has an exceptional tear strength, coinpa-

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Cl - ITTO€¦ · (Project PD 264104 Rev. 3 on, I) 2006-2008) ITTO, Plant Resources of Tropical...

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