row 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 mm. Secretory elements andcambial variants: 132: Iaticifers or tanniferous

tubes. Mineral inclusions: 152: crystals of othershapes (mostly small); 154: more than onecrystal of about the same size per cell or chain-her.

(R. Shanda, E. Uetimane, P. Baas & P. E. Gas-son)

Growth and development Seedlings areshade tolerant and can be found under the

mother tree in the forest. However, naturalregeneration is often poor. Once established,young trees may grow fairly fast. In a planta-tion of 54 years old the size of the holes showedconsiderable variation, with a maximum holediameter of 68 cm.

Ecology Cepholosphoero usQinborensis oc-curs in evergreen rainforest at 200-1500 inaltitude, often on steep slopes and along rivers

Propagation and planting Seeds of Cepho-10sphoero usumborensis are recalcitrant. Theyare heavy, with about 35 seeds per kg, andhave a thin seed coat and a small embryo em-bedded in a massive endosperm. After conec-tion, they should be packed in cotton bags inquantities of less than 10 kg. They loose viabil-ity quickly and should be sown within 6 weeksafter collection. The germination rate of freshseed without pre-treatment is high, up to 97%6-7 weeks after sowing. Scarification of theseeds can hasten germination, and when scan-fled seeds are sown in warm moistened soil

litter under shade they germinate within 3weeks. The seeds cannot withstand desiccation

below 25% moisture content, but they can bestored at temperatures below O"C. Seedlingestablishment is usually very low when seedsare sown directly into the field because theyare often damaged by rodents. It is preferableto sow them into polyethylene bags and plantseedlings out at the start of the rainy seasononce they have reached a height of 30-50 cm

Management Cepho!OSphoero usamberren-sts usually occurs scattered and in low densi-ties in the natural forest. It has been plantedin plantations. In the East Usambara moun-tains in Tanzania, Moesopsis eminii Engl. hasbeen planted as a nurse tree in young Cepho!0-sphoero usQinborensis plantations

Harvesting The bole usually does not splitduring felling operations and is quite easy to

fell with hand tools such as the two-inari cross-

cutsaw, bow saw and axeHandling after harvest Freshly harvested

logs are susceptible to blue-stain and borerattacks and should be converted immediatelyor treated with preservatives. Treatment of thetimber with an anti-stain solution before stack-

Ing for air drying is recommended. However,felling shakes may occur when large trees areharvested, and compression failures can bepresent in their buttlogs

Genetic resources Cepholosphoero usQin-borensis is threatened because of over-exploita-tion and low regeneration rates. It is endemicto a comparatively smallarea in eastern Kenyaand Tanzania, where it is subjected to serioushabitat degradation, and has been included inthe IUCN Red List as vulnerable.

Prospects Cepholoshoero usQinborensis isgreatly valued as a source of multi-purposetimber, and especially for plywood production.Moreover, it has good prospects for agroforest-ry systems and for commercial timber planta-tions. Research on growth rates, propagationand appropriate management systems is need-ed. Cepholosphoero usQinborensis should beincluded in domestication programmes becausesustainable and economicalIy viable timberproduction from natural forest seems no longerpossible within its limited area of distribution

Major references Anonymous, 1960a; Bolza& Keating, 1972; Chudnoff, 1980; Msanga,1998; Mugasha, 1978a; Mugasha, 1978b; Mu-gasha, 1983; Roe at a1. , 2002; Takahashi, 1978;Verdcourt, 1997

Other references Bryce, 1966; Bryce, 1967;Greenway, 1941; Lovett & Clarke, 1998; Lovettat a1. , 2007; Malde, 1972; Watt & Breyer-Brandwijk, 1962

Sources of illustration Verdcourt, 1997Authors W. Mojeremane

CHAETACHME 201

CHAETACHMEARISTATA E. Mey. ex Planch

Protologue min. Sci. Nat. , Bot. , s6r. 3, 10:341 (1848).

Family Geltidaceae (APG: Cannabaceae)Chromosome number it = 15

Synonyms Choetochme mudogoscoriensis Ba-ker (1885), Choetochme microcorpo Rendle(1916)

Vernacular names Thorny elm (En)Origin and geographic distribution Choe-

tochine onstoto is found from C6te d'TVoire

eastwards to Sudan and Kenya and it extends

202 TIMBERS2

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Chuetochme onstoto- wild

southward to Angola, Botswana, Zimbabwe,Mozambique, South Africa and Swaziland. Italso occurs in Madagascar.

Uses In DR Congo the wood of Choetochmeonstoto is used to make musical instruments

like guitars and tam-tams. It has been reportedto be used in Ghana to make the 'talkingdrums' of the Asante people. In Kenya thewood is used for clubs, and in Madagascar forspears. In South Africa the wood is used fortool handles and small ornaments. The Maasai

people in Kenya use the spiny branches to con-struct fences.

In C6te d'TVoire and Madagascar leaves andbark are applied in traditional medicine totreat coughs, colds and throat problems. In DRCongo an infusion of the roots is drunk as apurgative. The Zulu people of South Africa usethe bark to treat haemorrhoids and the POW-dered roots to reheve toothache. In the Central

African Republic a filtrate of young leavesground in water is drunk to treattuberculosis

Properties The wood is yenow-white. It isrecorded as heavy in Kenya, but elsewhere aslight-weight. In Madagascar it is considerednot durable, but is appreciated because itworks well. Leaves and stems contain 0.05%

and 0.08% of alkaloids, respectively. The seedcontains 6.4% of oil of which 82% is linoleic

acid

Botany Dioecious or monoecious, deciduousshrub to small tree up to 13 in tall, spiny; holeup to 30 cm in diameter; bark smooth, grey,becoming fibrous and longitudinalIy grooved;branches spreading, drooping, zigzag, withspines up to 3 cm long; twigs short-hairy toglabrous. Leaves alternate, simple; stipules up

to 2 cm long, united, soon falling; petiole 3-6min long; blade elliptical to elliptical-ovate, 3-9cm x 1.5-5 cm, base usually rounded andslightly asymmetrical, apex acute to short-acuminate, with distinct bristle up to 7 minlong, margins entire, occasionally toothed, gla-brous and glossy dark green above, sparselyshort-hairy and paler below, pinnately veinedwith c. 10 pairs of indistinct lateral veins. In-norescence an axillary cyme up to 1.5 cm long,often branched and congested, up to 30-flowered. Flowers unisexual, 5-merous, green-ish white; pedice1 I-3 min long; male flowerswith tepals 2-3.5 min x 1.5-2 mm, 5 stamensand a rudimentary ovary; female flowers oftensolitary in upper leafaxils, with tepals I-2 mmx I-1.5 mm and superior, sparsely hairy ovary3-5 mm long, styles 2, up to 2(-3) cm long.Fruit an ovoid to depressed-globose drupe up to3.5 cm x 2 cm, glabrous, yellowish orange whenripe.

Choetochme comprises a single species. Tradi-tionally, it has been treated as part of the raini-Iy Ulmoceoe, but later it was often consideredto belong to a separate family Celtidoceoe,whereas from most recent research it was pro-

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posed to take up the latter family into Coring-60ceoe.

Choetochme artstoto shows a lot of variation in

hairiness. In East Africa the species appears tobe mostly dioecious and in South Africa veryoften monoecious.

Ecology Choetochme artstoto is found inhumid evergreen forest to dry deciduous forest,but also in riverme and coastal forest, woodedgrassland and on sand dunes, up to 2100 inaltitude

Genetic resources and breeding As Choe-toehme onstoto is very widespread and notintensiveIy utilised, it is not liable to geneticerosion

Prospects Choetochme onstoto is not ofinterest for the timber trade because the hole is

too small. Research to identify the alkaloids inleaves, stem and roots is recommended in viewof their applications in traditional medicine.

Major references Burki11, 2000; Hanman,1948; Polhi11, 1966; Sytsma at a1. , 2002.

Other references Bussmann at a1. , 2006;Dale & Greenway, 1961; Hegnauer, 1973;Neuwinger, 2000; Noad & Birnie, 1989; Schatz,2001; van Wyk & Gencke, 2000; Wilmot-Dear,1991b; Wilmot-Dear, 1999.

Sources of illustration Ako6gninou, vander Burg & van der Maesen (Editors), 2006

Authors C. H. BOSch

I.

CHIDLowiA 203

CHIDLOWIASANGUINEA Hoyle

Protologue Bull. Misc. Inform. Kew 1932(2):101 (1932)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Origin and geographic distribution Chad!0-wig songuineo is restricted to West Africa,where it occurs from Guinea and Sierra Leoneto Ghana.

Uses In C6te d'Ivoire the wood, known as'bala', is locally used for joinery, stakes andrifle butts. It is suitable for heavy constructionand probably occasionally used in building 10-cal houses, and is also suitable for sliced ve-neer. The seeds are reportedIy eaten in C6ted'Ivoire.

Properties The heartwood is dark greenishgrey with a bronze tinge to dark brown anddistinctly demarcated from the narrow yellow-ish white to pale greyish brown sapwood. Thegrain is interlocked, texture moderately fineand even. Radialsurfaces show a ribbon figureThe wood is moderately heavy and hard. It is

Chidlowio songuineo - wild

difficult to saw due to its hardness and the

presence of interlocked grain. It is liable tosplitting upon nailingThe purine base triacanthine (chidlovine) hasbeen isolated from roots, twigs and leaves

Botany Small to medium-sized tree up to25(-30) in tall; hole often twisted, crooked, flut-ed and low-branching, up to 80(-100) cm indiameter, with many adventitious stems, occa-SIonally with steep buttresses; bark surfaceirregularly flaky and scaly, greyish to brown-ish, with many lenticels, inner bark fibrous,soft, pink to reddish brown; crown widelyspreading; twigs glabrous, with lenticelsLeaves alternate, panpinnately compound with4-6 pairs of leaflets; stipules small, caducous;petiole up to 2.5 cm long, rachis up to 25 cmlong; leaflets ovate to elliptical or obovate, 4-12cm x 2-5 cm, acuminate at apex, leathery, gla-brous. Inflorescence a slender, pendulous pani-cle up to 30 cm long, usually on older branches,many-flowered. Flowers bisexual, regular, 5-merous, deep red; pedicel slender, c. 3 minlong; calyx campanulate, c. 2 min long, withvery short teeth; petals free, ovate-elliptical tooblong-elliptical, 6-7 mm long, slightly fleshy;stamens 10, free, c. 2.5 cm long; ovary superior,slender, 4-5 mm long, with stipe c. 2.5 mmlong, style slender, c. 2 cm long. Fruit a strap-shaped pod up to 60 cm x 6 cm, flattened, gla-brous, glossy dark brown, dehiscing with 2woody valves becoming spiralIy twisted, up to15-seeded. Seeds nearly round, flattened, 2-2.5cm in diameter, shining red-brown and finelypitted. Seedling with epigeal germination; by-pocotyl c. 3 cm long, epicoty1 8-10 cm long;cotyledons thick and fleshy, rounded, c. 2.5 cm

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long; first 2 leaves opposite, pinnately coin-pound with 2 pairs o11eaflets.The flowers develop just after the appearanceof new leaves, in Sierra Leone between Febru-ary and May. It has been suggested that theyare pollinated by birds. Fruits ripen about oneyear after flowering. They are explosrvely de-hiscent, dispersing the seeds over some dis-tance. The presence of nitrogen-fixing rootnodules has been reportedChidlowio comprises a single species. Its affini-ty is stilluncertain

Ecology Chidlowio sanguineo occurs in ev-ergreen and moist semideciduous forest. It isusually found in the middle storey of primaryforest, but occasionally also in secondary forestand gallery forest. It was very abundant in theupper basins of the Sassandra and Cavallyrivers, locally occurring in nearly pure standsin the understorey. It also occurs in forest inthe mountains, e. g. in the Mount Nimba re-gion.

Management The boles of large trees areoften hollow, which diminishes the commercialvalue and may cause danger during felling

Genetic resources and breeding Chid-low, o songuineo Is locally common, e. g. inwestern C6te d'TVoire and in parts of SierraLeone and Liberia. However, with the split-ting-up of undisturbed forest in West Africa, itmay be liable to genetic erosion

Prospects Chidlowio songuirteo will likelyremain ofno commercial importance because ofits usually poorly shaped and short hole and itshard wood that is difficult to saw and work.

Little is known about this and re-species,

search is needed on phytochemistry and possi-ble toxic effects of the seeds, which are reput-edly edible.

Major references Burki11, 1995; Hoyle,1932; Kryn & Fobes, 1959; Savill & Fox, 1967;Tra Bi, 1997.

Other references Aubr6ville, 1959b; Bou-quet & Debray, 1974; de to Mensbruge, 1966;Hawthorne & Jongkind, 2006; Irvine, 1961;Keay, Hoyle & Duvigneaud, 1958; Kunkel,1965; Lewis at a1. , 2005; Normand, 1950a;Normand & Paquis, 1976.

Authors R. H. M. J. Lemmens

Synonyms Cleistonthus mildbroedii Jabl(1915), Cleistonthus inichelsortii J. Leonard(1955)

Origin and geographic distribution Cleis-tonthus COMdotus occurs from Cameroon east

to the Central African Republic and south toGabon and DR Congo

Uses The wood is used in carpentry andjoinery. It is suitable for heavy construction,heavy flooring, interior trim, mine props, shipbuilding, hydraulic works, vehicle bodies, rail-way sleepers, toys, novelties and agriculturalimplements

Properties The heartwood is dark red-brown, often with a purple tinge, and distinctlydemarcated from the pale pinkish brown, up to5 cm wide sapwood. The grain is straight, tex-ture fine and even. Quarter-sawn surfacesshow a slight figure of darker streaks and palersilver grain. A reddish resinous substance isoften present in the woodThe wood is heavy, with a density of 920-1080kg/in3 at 12% moisture content, hard and verystrong. The rates of shrinkage during dryingare high, from green to oven dry about 4.3%radial and 9.8% tangential. It is recommendedto quarter-saw logs before drying to avoid ex-cessive splitting. After drying, the wood ismoderately stable to unstable in service. At15% moisture content, the modulus of ruptureis 178-230 N/mm2, compression parallel tograin 69-86 N/mm2, cleavage 26 N/min andChalais-Meudon side hardness 11.1

The wood saws slowly and is rather difficult toplane due to its hardness and abrasiveness, butit can be finished to a nice surface. It holds

nails and screws well, but pre-boring is needed.The wood is durable, being resistant to fungal

CLEISTANTHUSCAUDATUS Pax

Protologue De Wild. & T. Durand, min. Mus.Congo, Bot. , s6r. 2, I: 49 (1899).

Family Euphorbiaceae(APG: PhyUanthaceae)

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01eistonthus COMdotus - wild

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attacks, but it has been reported to be slightlysusceptible to insect attacks. It is very re-sistant to impregnation with preservatives.

Botany Shrub or small to fairly large treeup to 35(-40) in tall; hole branchless for up to20 in, usually straight and cylindrical, up to90(-150) cm in diameter, usually without but-tresses; bark surface flaky with elongatescales, greyish brown, inner bark fibrous, pinkto red, with a reddish or whitish exudate;crown rounded, large; twigs hairy, but soonbecoming glabrous. Leaves alternate, simpleand entire; stipules c. I cm long, early cadu-cous; petiole up to I cm long; blade ovate toelliptical or oblong, 3.5-22 cm x 1.5-10 cm,base curieate to slightly cordate, apex long-acuminate, glabrous to slightly halry below,pinnately veined with up to 8 pairs of lateralveins. Inflorescence an axillary raceme up to 20cm long, glabrous or slightly hairy, wlth smallbracts or small caducous leaves. Flowers uni-sexual, regular, (4-)5(-6)-merous, greenish orreddish; pedicel up to Tel. 5) cm long; sepalsoblong-lanceolate to ovate-lanceolate, 4-9 mmlong, glabrous; petals linear, I-4 mm long; diskannular; male flowers slightly smaller thanfemale ones, with 5(-6) stamens fused at baseinto a column of 2-4 min long, ovary rudimen-tary; female flowers with superior, roundedovary, glabrous or slightly hairy, (2-)3(-4)-celled, styles (2-)3(-4), 2-4 mm long, 2-branched. Fruit a (2-)3(-4)-celled, nearly glo-hose capsule 1.5-3 cm in diameter, reticulatelyveined, splitting into 2-valved, (1.2-seededsegments, with central column persistentSeeds c. I cm long, brownish, shiny. Seedlingwith epigeal germination; hypocoty1 6.5-10 cmlong; cotyledons leafy, broadly ovate, c. 3 cmlong; first leaves alternate.Cleistonthus occurs throughout the tropics ofAsia, Australia and Africa, and comprisesabout 150 species. About 20 species occur inmainland tropical Africa, mainly in CentralAfrica, and about 6 in Madagascar. Studies ofleaf anatomy and pollen, as well as a phyloge-netic study using DNA sequence data, showedthat Cleistonthi, s is heterogeneous; a criticalreview of the genus is needed.01eistonthus sch!echteri(Pax) Hutch. is a shrubor small to medium-sized tree up to 20 in tall,with hole up to 40 cm in diameter. It occurs inmixed deciduous forest, woodland and thicketsfrom Kenya south to northern South Africa. Itsdark brown and hard wood is used in construc-

tion for posts, beams and roof laths, and forrailway sleepers and walking sticks. In South

Africa powdered bark is applied to burnsEcology 01eistonthus edudutus occurs in

lowland primary evergreen or semi-deciduousrainforest and riverme forest, also in swampyor periodically flooded forest, up to 700 in alti-tude

Management Cleistontht, s cowdotus Is 10-cally common in the forest. Logs should be con-verted soon after felling to avoid checking;quarter-cutting is recommended. A log of 13 inlong and 76 cm in diameter yielded 5.3 ing ofwood

Genetic resources and breeding C!eis-tonthus eaudotus is fairly widespread and 10-cally common. Although it is unknown to whatextent it is exploited, there are no Indicationsthat it is subject to genetic erosion at present

Prospects The wood of Cleistonthus COMdu-tus and other Cleistonthus spp. is likely to re-main of limited Importance, mainly for localconstruction because of its durability. Thegrowth rates are probably low, as reported forCleistonthus schlechteri with an average annu-al hole diameter growth of little more than I

Major references Bolza & Keating, 1972;Breteler, 2011; Fouarge, G6rard & Sacr6, 1953;Leonard, 1962; Tailfer, 1989

Other references Coates Palgrave, 1983;Gaugris, van RDOyen & Bothma, 2008; Gaugrisat a1. , 2007; Grace at a1. , 2002a; Kathriarachchiat a1. , 2005; Leonard, 1955; Leonard, 1960;Neuwinger, 2000; Radcliffe-Smith, 1987a; Rad-cliffe-Smith, 1996.

Authors L. FA. Oyen

CLEisTOPHOLiS 205

min.

CLEISTOPHOLiS PATENS (Benth. ) Engl. &Diels

Protologue Engl. , Monogr. afrik. Pnanzen-Fam. a 35 (1901).

Family AnnonaceaeChromosome number2n= 14

Origin and geographic distribution C!eis-topholis pote?!s is widely distributed from Sen-egaleastward to Uganda, and southward to DRCongo and Cabinda (Angola)

Uses The wood, known as 'avom', is used forjoinery, door frames, roof-beams, drums, floatsand canoes. It is suitable for furniture, sportinggoods, boxes, crates, Implements, food contain-ers, veneer, plywood, hardboard, particle board,wood-wool and pulpwood. Tree holes are usedto float heavy timber in rivers. Wood ash issometimes used as food preservative

206 TIMBERS2

.

C!eistopholispotens - wild

The fibrous bark is used for matting, basketry,cordage and hut walls. Several plant parts arecommonly used in local medicine. Bark decoc-tions are taken to treat stomach-ache, diar-rhoea, tuberculosis, bronchitis and hepatitis.Bark pulp is applied against swellings, oedemaand whitlow, and bark sap is dropped into thenose to treat headache and rubbed in to treatrickets in children. In Uganda crushed bark isused in preparations to treat malaria and mea-SIes. In Nigeria the bark is used to treat ty-phoid fever and bark extracts are used in thetreatment of menstrual irregularities. The rootbark is used as vermifuge. Leaf infusions ordecoctions are administered against hepatitis,fever, trypanosomiasis and rheumatic arthritis,and as vermifuge. Roasted and ground seedsare applied against headache. Seeds have beenused as beads.

Production and international trade The

wood is only used locally and not or rarelytraded on the international market.

Properties The heartwood is greyish white,sometimes with a pinkish tinge, indistinctlystreaked, indistinctly demarcated from thesapwood. The grain is straight, texture coarsebut even. The wood is lustrous.

It is lightweight, with a density of 290-500kg/in3 at 12% moisture content, and rather soft.The wood air dries rapidly, but should bestacked properly to avoid degrade; the rates ofshrinkage are rather high, from green to ovendry 5.0% radial and 9.6% tangential. Once dry,the wood is moderately stable in service. At12% moisture content, the modulus of ruptureIs 29-38 N/mm2, modulus of elasticity 6080-8230 N/mm2, compression parallel to grain 38

N/mm2, shear 11 N/min2 and Janka side hard-ness 1410 N.

The wood Is easy to saw and work. It planeswell and finishes smoothly. It does not spliteasily upon nailing and the nail-holding poweris satisfactory. The wood peels well. It is notdurable, being liable to fungal, termite, Lyetusand marine borer attacks. Tests showed inod-

erate pulping characteristics for paper produc-tion

Methanol extracts of the bark showed distinctantimicrobial activity. The steroidal fraction ofthe extract showed much more potent activityagainst Boci!jus subtilis and Klebsiello piteu-moniue than penicillin and chloramphenicol,the glycoside and alkaloid fractions showedsignificant activity against Klebsiello prteu-monme, and the saponin fraction against Sol-monel!o typhimi, rium. The alkaloids eupolauri-dine, onychine and 3-methoxysampangine havebeen isolated from the bark; these compoundsshowed potent antifungal activity against Con-did0 o16icons. Additionally, bark extracts exhi-bited arithelmintic activity against Rhobdttispseudoelongoto.The essential oilfrom the leaves contains (E)-^-ocimene as main constituent, that from thebark myrcene, P-cymene and germacrene D,and that from the fruits Iinalool and (E)- and(Z)-11nalool oxides. The essential oils from barkand leaves showed distinct in-vitro activityagainst Pigsmodium foldporum. Safety testsshowed no inutagenic activity against Solmo-itello typhimurium and Escherichia colistrains.

Administration of methanol extracts of the

bark to female rats showed a dose-dependentIncrease in the levels of several hormones in-

cluding progesterone and oestrogen. Severalpartially acetylated ongorhamnosides havebeen Isolated from the leaves; some of theseshowed significant antibacterial activityagainst methicillin-resistant Stophylococcusoureus. Fruit extracts exhibited in-vitro an-tileishmanial activity. Powdered bark showedprotection against Indian meal moth (PIOdiointerpunctello), which is a pest in stored foodproducts.

Botany Smallto medium-sized tree up to 20(-30) in tall; hole branchless for up to Toe20)in, usually straight, cylindrical and slender, upto 80(-90) cm in diameter, sometimes slightlyfluted at base; bark surface smooth, shallowlyfissured, greyish white to grey, inner barkstrongly fibrous, peelable in long strips, whiteto pale orange-brown, scented; crown with hor-

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glabrous; cotyledons leafy, elliptical, c. 6 cm x 4cm; first leaves alternateC!eistopholis patens is a pioneer which growsrapidly. In Sierra Leone 7-year-old treesreached 13 in tall and over 20 cm in hole diam.eter, and in Ghana the same height and diame-ter have been reached on 4-year-old loggingtracks. In Guinea trees have been reported tostart fruiting already 5 years after planting. InC6te d'TVoire and Ghana ripe fruits occur inAugust-November. In Uganda rotten materialof dead Cleistopho!is potens trees is an Tin-portant food for chimpanzees; they chew it,extracting the juiceC!eistopho!is comprises 3 species, all in tropicalAfrica, and seems related to Cottongo and Let-towionthus

Cleistopholis gigueo Pierre ex Engl. & Diels isa small to medium-sized tree up to 35 in tallwith hole up to 80 cm in diameter, occurringfrom Cameroon east to the Central AfricanRepublic, and south to DR Congo and Cabinda(Angola). The lightweight wood, with a densityof 300-400 kg/ina at 12% moisture content, Isused for similar purposes as that of Cleis-topholis potens. The fibrous bark is used forcordage and hut walls. Bark decoctions or mac-erations are taken to treat bronchial coin-

plaints, tuberculosis, stomach-ache, diarrhoeaand scabies, and as emetic and vermifuge. Rootsap is used in mixtures to prepare dart poison.C!eistopho!is stowdtii Engl. & Diels is a sinaUtree up to 15(-25) in tall with hole up to 30 cmin diameter, occurring from southern Nigeriato Gabon. Its wood is probably used for similarpurposes as that of Cleistopholis potens. Thefibrous bark is used for cordage and hut walls.Lettowionthus stel!otus Diels differs from C!eis-

topholis spp. in its larger flowers and nearlysmooth seeds. It is a smalltree up to 15 in tallrestricted to Tanzania, where it occurs in low-land riverme and disturbed forests. Its light-weight wood is occasionally used for buildingpoles, spoons and tool handles. Several gera-nylbenzoquinonoids have been isolated fromthe fruits; lettowienolide and lettowiquinoneexhibited mild in-vitro activity against PIOs-inodium foieiporum

Ecology 01etstopholis potens is most coin-monly found in riverme and swamp forest, andin secondary forest. It prefers flat, disturbedand wet sites, but can also be found in ever-green forest on slopes, up to 1100 in altitude. Itdoes not tolerate fire

Management There are 600-700 seeds perkg. In Guinea seeds harvested in April started

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izontal branches drooping at tips; twigs oftenwith small ridges, glabrous. Leaves alternate,simple and entire; stipules absent; petiole up toI cm long; blade usually narrowly oblong-elliptical, (2.5-)10-25(-30) cm x (1.5-)3-6 cm,curieate to rounded and slightly asymmetricalat base, short-acuminate at apex, papery tothin-leathery, glabrous, shiny, pinnatelyveined with 8-24 pairs of lateral veins. Inflo-rescence a small axillary fascicle, up to 8-flowered, nearly glabrous. Flowers bisexual,regular, 3-merous, greenish yellow; pedicel I-2.5 cm long; sepals free, broadly ovate, c. 2 mmlong; petals free, in 2 whorls, outer ones obo-vate to obovate-oblong, 0.5-I cm long, Innerones shorter, short-hairy at margins; stamensnumerous, c. I min long; carpels c. 10, ovariesc. I mm long, styles minute. Fruit consisting ofup to 9 nearly globose follicles 1.5-2 cm in di-ameter, with stipe c. 7 min long and slightlyconstricted between the seeds, warty when dry,glabrous, indehiscent, I-2-seeded. Seeds glo-hose to ellipsoid, I-1.5 cm in diameter, stronglywarty. Seedling with epigeal germination; hy-pocoty1 8-12 cm long, epicotyl c. 1.5 cm long,

CLEISTOPHOLIS 207

208 TIMBERS2

germinating about 15 days after sowing. How-ever, dormancy may occur; seeds harvested inNovember took 5 months to germinate. Seed-lings are ready for planting 3-4 months aftergermination. They should be planted in fullsun. In plantations with a spacing of 3 in x 3in, weeding is necessary during 2-3 years. Thecanopy closes after 3-4 years and a thinning to50% of the trees is needed after 4-5 years. Asecond thinning can be done after 10 years. InGuinea mixed plantations, e. g. with rigyospp. , have been recommendedIn the forest, seedlings and saplings are onlyfound in canopy gaps. In Uganda the mortalityrate of seedlings was nearly 50% before theyreached the sapling stage. In waterlogged andriverIne areas 01eistopho!is potens can beabundant. In swamp forest in Uganda, up to 40individuals per ha can be found, with a propor-tion of trees with a hole diameter above 40 cm

of 8%. In forest in Cameroon an average densi-ty of 0.5 tree with a hole diameter of over 60 cmhas been reported per ha, with an averagewood volume of 1.1 in8tha. A hole with a lengthof 9 in and nearly 60 cm in diameter felled inDR Congo yielded 1.7 ina of wood. Logs shouldbe processed immediately after felling or treat-ed with fungicides, because the wood is verysusceptible to blue stain attack. In southernNigeria larvae of the Saturniid moth Bundeoo1cinoe have been reported to attack the foli-age; these caterpillars are edible

Genetic resources and breeding Cleis-topholis potens is widespread and locally coin-mon and there are no indications that it isthreatened. Immediate conservation measuresdo not seem to be necessary, as is the case for01eistopholis stowdtii which is classified asvulnerable on the IUCN Red List

Prospects 01eistopholis potens and relatedspecies will remain of local importance as pro-ducer of wood that is easy to work. Its invasivenature in disturbed forest and rapid growthseem to offer good possibilities for commercialtimber production on suitable sites in ever-green lowland forest, especially for veneer,plywood and particle board productionSeveral of the applications in traditional medi-cine have been confirmed by research, e. g. an-timicrobial, arithelmintic and antimalarial acti-vities. This may offer opportunities for drugdevelopment.

Major references Atuhe, 2010; Bolza &Keating, 1972; Burki11, 1985; Fouarge & G6r-ard, 1964; Vivien & Faure, 1985

Other references Akendengu6 at a1. , 2009;

Amadj, Odigie & SIminialayi, 2009; Boyom ata1. , 2011; de Koning, 1983; Ebi & Kamalu,2001; Ekundayo at a1. , 1988; Hufford at al. ,1987; Neuwinger, 2000; Takahashi, 1978;Verdcourt, 1971.

Sources of illustration Ako6gninou, vander Burg & van der Maesen (Editors), 2006

Authors R. H. M. J. Lemmens

COELOCARYONPREUSSllWarb.

Protologue Notizbl. K6nigl. Bot. Gart. BerhnI: 99 (1895)

Family MyristicaceaeSynonyms Coelocoryon MomeiPierre (1903).Origin and geographic distribution Coelo-

conyon prei, 88n occurs from southern Beninand Nigeria east to the Central African Repub-Iic, and south to Gabon and DR Congo

Uses The wood, known as 'ekoune' or 'ekun',Is used for interior Joinery, furniture, cabinetwork, veneer and plywood. It is suitable forlight construction, interior trim, ship building,toys, novelties, boxes, crates, matches, turnery,hardboard, particle board and pulpwood. Tra-ditionally, it is used for planks and paddles.The seed contains an edible fat. The bark iscommonly used in traditional medicine. Barksap is taken to treat blood in the urine, dys-menorrhoea, dysentery and haemorrhoids,whereas bark decoctions or macerations areadministered as expectorant, emetic and ano-dyne, and to treat cough, lung complaints,tachycardia, rheumatism and oedema. Barkdecoctions are used as an enema to treat colicand diarrhoea, and bark powder is applied towounds as haemostatic.

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Production and international trade Bet-

ween 1961 and 1968 Equatorial Guinea ex-ported on average 18,300 in3 of logs per year.In Gabon the export of 'ekoune' timber wassmall in 1991 with about 50 in3, but it in-creased to 7600 in3 in 1996, decreasing again to140 in3 in 1999 and 240 ina in 2005. Current

export volumes are smallProperties The heartwood is pale brown to

yellowish brown, with darker markings, andindistinctly demarcated from the slightly paler,wide sapwood. The grain is straight, texturemoderately fine. The wood is lustrous. It islightweight to medium-weight with a density of450-660 kg/ina at 12% moisture content. It airdries easily without serious defects. The ratesof shrinkage are moderate, from green to ovendry 3.0-5.1% radial and 6.2-8.5% tangentialOnce dry, the wood is unstable to moderatelystable in service.

At 12% moisture content, the modulus of rup-ture is 73-124 N/mm2, modulus of elasticity7550-12,500 N/mm2, compression parallel tograin 29-49 N/min2, shear 49 N/mm2, cleav-

11-19 N/min and Chalais-Meudon sideagehardness 1.4-4.0.

The wood is easy to saw and work with bothhand and machine tools. It nails and screws

well, with good holding capacity. It glues,paints and varnishes satisfactorily, and hasgood peeling and slicing properties. It Is notdurable, being susceptible to termite, Lyetusand marine borer attacks. The wood is quiteeasy to treat with preservativesThe wood contains 42-47% cellulose, 27-33%11gnin, 15-16% pentosan, 0.7-0.9% ash andvery little silica. The solubility is 2.5-3.4% inalcohol-benzene, 2.4-3.1% in cold water and17.3% in a I% NaOH solution. A test on the

kraft pulping characteristics of the woodshowed satisfactory results with a yield ofabout 45%

The fat from the seed has a smell of cacao but-

ter. Phytochemical screening of the barkshowed the presence of alkaloids, tannins, sap-onins and terpenoids

Adulterations and substitutes The wood

resembles that of Pycnonthi4s origolensisorelw. ) Warb. and is used for similar purposes

Description Evergreen, dioecious, medium-sized to fairly large tree up to 35 in tall; holebranchless for up to 20 in, generally straightand cylindrical, up to 80(-110) cm in diameter,often grooved at base or with small buttresses;bark surface scaly with elongate scales, reddishbrown, inner bark granular or fibrous, pinkish

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Coelocoryon preussii - I, 60se of 601e, . 2, led/: 3,mole inflorescence, . 4, port of mole inflorescence,5, fernole inflorescence, . 6, port of ferno!e milo-rescence, ' 7, dehisced fruit showing seed.Redrown und odopted by Ishoh Syomsudin

to yellowish brown with darker spots, with acreamy exudate; crown pyramidal, with hori-zorital branches; twigs glabrous. Leaves alter-nate, simple and entire; stipules absent; petioleI-2 cm long, grooved above; blade obovate-oblong, 15-25 cm x 6-8 cm, curieate at base,short-acuminate at apex, thin-leathery, gla-brous, pinnately veined with 6-10 pairs of Iat-eral veins. Inflorescence an axillary panicle upto IOC15) cm long, short-hairy, with flowers innumerous umbel-shaped clusters, each clusterinitially enclosed by an involucre and up to 30-flowered. Flowers unisexual, regular, up to 3mm long, yellowish, with short pedicel, perl-arith cup-shaped, 3-4-10bed, reddish brownshort-hairy; male flowers with 3-5 fused sta-mens, filaments merged into a column; femaleflowers with superior, ovoid, glabrous, I-celledovary, stigma 2-lobed. Fruit an ellipsoid to ob-long or globose drupe 3.5-4 cm x 2.5-3 cm, inbunches of up to 3, yellowish when ripe, split-ting longitudinalIy with 2 fleshy valves, I-seeded. Seed ellipsoid, 2-3 cm x I-1.5 cm,

2

210 TIMBERS2

glossy dark purplish brown to blackish, withpink to red anI, Iaciniate almost to the base.Seedling with hypogealgermination.

Other botanical information Coe!ocoryoncomprises 4 species and is restricted to Westand Central Africa.

The wood of Coelocoryon orycorpum Stapf, amedium-sized tree up to 30 in tall occurringfrom Senegal to Ghana, is used for similarpurposes as that of Coelocoryon preussit. Thebark sap is used as a purgativeCoelocoryon sphoerocorpum Fouilloy, a medi-urn-slzed tree up to 30 in tall occurring fromGuinea to C6te d'Ivoire, provides a wood simi-Iar to that of Coelocoryon preussii. The leavesand seeds are used in traditional medicine totreat drowsiness

Coe!ocoryon botryoides Vermoesen is also amedium-sized tree up to 30 in tall, and occursfrom south-eastern Nigeria and Cameroon tothe Central African Republic and DR CongoIts wood is also used for similar purposes

Anatomy Wood-anatointcal description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 14: scalariformperforation plates; 15: scalariform perforationplates with S 10 bars; 20: intervessel pits sca-Ianform; 21: intervessel pits opposite; 22: in-tervessel pits alternate; 26: intervessel pitsmedium (7-10 pin); 32: vessel-ray pits withmuch reduced borders to apparently simplepits 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; 46: S 5 vessels per square minimetre; (47:5-20 vessels per square minimetre). Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; (65: septate fibres present); 66:non-septate fibres present; 68: fibres very thin-walled; 69: fibres thin- to thick. walled. Axialparenchyma: 78: axial parenchyma scanty par-atrachea1; 93: eight (5-8) cells per parenchymastrand. Rays: 97: ray width I-3 cells; (102: rayheight > I mm); 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; (108: body ray cells procumbentwith over 4 rows of upright and/or square mar-ginalcells); 1/5: 4-12 rays per min.(R. Shanda, E. Uetimane, P. Baas & H. Beeck-man)

Growth and development The growth of

Coelocoryon preussiiis slow. Seedlings reachedonly 20-30 cm tall after 10 months and 140 cmafter 2 years. In Garnero0n trees flower mainlyin December-February and fruits ripen about 6months later. In Guinea trees fruit in July-September, in 06te d'TVoire usually in April-June. The seed anlis eaten by several animalspecies including hornbills, turacos, monkeysand chimpanzees, which disperse the seeds.

Ecology Coelocoryon preussiiis wldespreadin lowland rainforest up to 400 in altitude. Itcan be common in secondary forest, where it isoften found in small clusters. It usually occurson well-drained soils

Propagation and planting There are about375 seeds per kg. The seeds start germinating2-6 weeks after sowing, but the germinationrate Is often very low. In the nursery, seedlingsshould be kept in the shade; there was no sur-vivalafter planting in fullsun

Management In the forest of Garnero0n theaverage density of trees with a hole diameter ofmore than 60 cm is about 0.1 per ha, with anaverage wood volume of 0.3-0.6 matha. In Ga-bon the average wood volume is locally 2 in31ha,whereas an inventory in a large area in north-eastern Gabon resulted in an estimated aver-

age volume of 0.5-0.8 mama.Diseases and pests Studies in southern Ca-

meroon showed that several insect species at-tack the trees, particularly shoot and woodborers and insects feeding on the leaves orsucking sap, but damage is usually only slight

Handling after harvest Freshly harvestedlogs should be removed from the forest rapidlyor treated with preservatives to avoid bluestain and borer attacks. It is recommended to

leave the bark attached to logs. They float inwater and thus can be transported by river

Genetic resources Although Coelocoryonpreussii usually occurs scattered in the forest,it is quite widespread in different forest typesIncluding secondary forest. There are no indi-cations that it is threatened by genetic erosion.

Prospects Coe!ocoryon preussii and otherCoe!ocoryo, I spp. may be interesting for furthercommercialization, providing timber that, alt-bough not durable, can be used for variouspurposes including veneer and plywood produc-tion. Their common occurrence in secondaryforest formations may imply good prospects forsustainable production in natural forest, butresearch on growth rates, regeneration andproper managementsystems are needed.Phytochemical and pharmacologicalinvestiga-tions of the bark are recommended to assess

the value of medicinal applications. The seedmay have commercial importance for the pro-duction of edible oil.

Major references ATIBT, 1986; Bolza &Keating, 1972; Burki11, 1997; CTRAD ForestryDepartment, 2008; CTFT, 1977b; de Saint-Aubin, 1963; Foullloy, 1974; GIIbert & Troupin,1951; Takahashi, 1978; Vivien & Faure, 1985.

Other references Ako6gninou, van derBurg & van der Maesen (Editors), 2006; Chris-ty at a1. , 2003; Foahom, 2002; Fouilloy, 1965;Foullloy, 1972; Hawthorne & Jongkind, 2006;Hubert, undated; Keay, 1989; Maisonneuve &Manfredini(Editors), 1988b; Neuwinger, 2000;Onanga at a1. , 1999; Pauwels, 1993; Raponda-Walker & Sillans, 1961; Sallenave, 1955; Tail-for, 1989; Terashima & Ichikawa, 2003; White& Abernethy, 1997; Wilks & TSSemb6, 2000.

Sources of illustration FoulUoy, 1965;Wilks & TSSemb6, 2000.

Authors R. B. JiofackTafokou

ern Africa, where it occurs in Zambia, Malawi,southern Angola, northern Namibia, north-eastern Botswana, Zimbabwe, southern MD-zarubique and northern South Africa. It hasalso been planted in semi-and regions of India

Uses The wood of 0010phospermum moponeis traditionally used for posts and poles in theconstruction of houses and palisade fences, andfor carving. The wood accounts for more than90% of the wood used for living and storinghuts in large parts of southern Africa. It issuitable for heavy flooring, mine props, shipbuilding, vehicle bodies, railway sleepers, lad-ders, sporting goods, toys, novelties, agricul-turalimplements, tool handles, turnery, hard-board and particle board, and to a lesser ex-tent, due to its weight and hardness, for joineryand furniture. Colophospermum mopone pro-vides high quality firewood and is used forcharcoal production. The wood, that has beenclaimed to be the best firewood of Africa, burnswith great heat, slowly and evenly, and leaveslittle ash; it lights easily, even when green, butits hardness makes it difficult to fell, chop orsplit. IntricateIy shaped pieces of root are ex-ported from Namibia to eastern Asia, wherethey are used as decorations for fish tanks andhomes.

Seeds are consumed by humans asfaminefood0010phospermum moponeis host of caterpillarsof the mopane moth or emperor moth (Gonim-brosio balmo) which feed on its leaves. Thecaterpillars are widely dried, roasted and con-sumed in southern Africa, and are collectedand sold to generate income. The tree alsohosts the psyllid Retroactzzio moponi (syno-nym: Arytoino moponi), a sap-sucking insectthat produces 'mopane manna', a sweet wax-like cover on the leaves that is collected and

eaten by people but also monkeysGolophospermum mopone is used in the prepa-ration of traditional medicines. In Namibiaextracts from heated wood or the fibres fromchewed leaves are used to treat slow-heatingwounds or to stop bleeding of wounds. Topnaarpeople in Namibia drink a decoction of theleaves against colds. To treat pain of the eyesor headache, a leaf decoction is dropped in theeye or the head is washed with it. In SouthAfrica decoctions of the bark and wood are used

to treat eye inflammations, venereal diseases,syphilis, diarrhoea and stomach pains. In Zim-babwe bark infusions are sometimes given tocattle against diarrhoea. Infusions or decoc-tions of the leaves are drunk or given as enemato treat constipation and stomach-ache. Twigs

CoLOPHOSPERMUMMOPANE (Benth. )J. Leonard

Protologue Bull. Jard. Bot. Etat 19: 390(1949)

Family Caesalpiniaceae or. ,eguminosae - Cae-salpinioideae)

Chromosome number2n = 34.36Synonyms GOPoifero mopone Benth. (1865),

Horduiichio mopone (Benth. ) Breteler (1997)Vernacular names Mopane, turpentinetree,

balsam tree, butterfly tree, ironwood (En). Mo-pane (ET). Mupane, chanate (Po).

Origin and geographic distribution Colo-phospermum mopone Is widespread in south-

GOLOPHOSPERMUM 211

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212 TIMBERS2

are used as chew-sticks to clean the teeth. TheVenda people in South Africa use the roots tostop bleeding of the gums. In Mozambiqueroots have been used to killintestinalworms

Ash from the wood has high phosphate, calci-urn and lime contents and is used as fertilizer

and to make whitewash. The bark is used in

tanning hides. The inner bark provides fibrethat makes a very strong rope. The leaves andfruits are browsed by livestock. They retaintheir nutritional value after falling to theground; animals eat them off the ground dur-Ing the dry season. They are crucial to livestockfarmers for bridging the late dry season periodwhen many trees are leafless. Fresh leaves andseeds smell strongly of turpentine, but thisdoes nottaintthe meat or milk of animals feed-

Ing on them. The tree also acts as a food plantfor a wild silk moth (Gonometo rufobrunneo).Cocoons of the moth are harvested as wild silkand processed to make cloth.

Production and international trade Thereis little, if any, International trade in mopanetimber. Production of timber, firewood andcharcoal for local and regional markets is ex-tensive, but statistics are not available. Apartfrom mopane wood products, there is a Iucra-tive trade in mopane worms in formal and in-formal markets across southern Africa. The

total value of this trade is not known preciselybut has been estimated to be worth many inn-lions Us$.

Properties 0010phospermum mopone woodIs very attractLve in appearance. The heart-wood is medium to reddish brown, with blackstripes, darkening with age; it is clearly de-marcated from the thin, off. white to pale brownsapwood. The grain is interlocked, sometimesnearly straight, texture coarse to moderatelyfine and even. The wood is oily and shows littlelustre

The wood is very heavy, with a density of 990-1230 kg/ina at 12% moisture content, and hard.It air dries slowly but well, although it may beliable to surface checking. It does not easilyend-split during drying; the oily content of thewood might explain this feature. Kiln dryingmay cause moderate surface checking and dis-tortion. The rates of shrinkage are low to inod-erate, from green to oven dry about 4% radialand 5.2% tangential. At 12% moisture content,the modulus of rupture is 89-122 N/min2 inod-ulus of elasticity 10,880-14,000 N/mm2 coin-pression parallel to grain 69-75 N/min2 shear17 N/mm2, Janka side hardness 13,100-21,000N and Janka end hardness 19,400 N

Mopane wood is generally difficult to work withboth hand and machine tools, mainly on ac-count of its high density. Machining has a se-vere blunting effect on saw teeth and cuttingedges, and special tools are needed to obtain afine surface and a good finish. Planing is diffi-cult due to the presence of interlocked grainThe wood is difficult to nail and screw due to

its hardness, and requires pre-boring; the nail-holding power is good. Polishing gives fair togood results. The wood varnishes, glues andturns well. It is very durable as it containschemicals that resist attacks by termites andpowder-post beetles. The heartwood is ex-tremely resistant to treatment with preserva-tives, the sapwood is moderately permeableThe wood ash contains about 15% lime. Thebark contains 5.9% tannin. The nutritionalvalue of 0010phospermum mopone foliage wasassessed in South Africa. On average, the foli-age contained per 100 g dry matter: ash 5.8 g,crude protein 14.1 g, neutral detergent fibre380 g, acid detergent fibre 30.7 g, acid deter-gent insoluble N 1.0 g, acid detergent 11gnin14.6 g and ether extract 4.2 g. The mean con-tent of macro-nutrients in the foliage has beenestimated per 100 g dry matter at: Ca 1.0 g, P0.16 g, K 0.8 g, Mg 0.2 g, and Na 0,004 g. Themean content of micro-nutrients was assessed

per 100 g dry matter at: Cu 0.8 ing, Co 0.3 ing,Mn 5.6 ing and Se 001 ing. The foliage con-tains relatively high concentrations of con-densed tannins, which tend to diminish in old-er leaves, making them more acceptable tobrowsers during the dry Smallseason.

amounts of leaves added to the diet of youngpigs increased their growth rate and produc-tion.

The diterpene 8(S), 13(S)-dihydrogrindelic acidand its C-13 epimer were isolated from theseeds; both are intermediates in the biosynthe-sis of 9.13-epoxylabdanes. The aerial parts ofthe plant are rich in essential oil, consistingmainly of orpinene (68-72%) and 11monene(3.5-5%), which are presumably responsible forthe strong turpentine odour of the fruits andleaves. The leaves also contain significant con-centrations of p-sitosterol and stigmasterol,which are apparently the source of sterols invarious organs of the mopane moth. The poly-phenolic pool of the heartwood exhibits ex-treme diversity and complexity. It comprises avariety of monomeric navonoids, navan-3.4-diols Including mopanols and peltogynols, fla-vonols, dimeric proanthocyanidins, profisetini-dins, promopanidins, propeltogynidins, and a

variety of profisetinidin-type triflavanoidsDescription Deciduous smalltree, often re-

maining stunted as a shrub, up to 15(-24) intall, with one or more straight holes up to 100cm in diameter; bark rough, deeply verticallyfissured, dark grey to brown, Irregularly scal-ing off, often forming elongate-reticulatepatches, inner bark fibrous, laminated, pink tored; crown narrow, with few main branchesdiverging at a narrow angle; twigs glabrous,smooth, grey. Leaves alternate, compound witha single pair of leaflets; stipules ovate, up to 5mm long, early caducous; petiole (I-)1.5-4(-5)cm long, with a small, flat appendage 2-4(-5)min long at apex; pulvinus at base of leafletbroader than long, contiguous with the OPPo-site one; leaflets obliquely ovate or lanceolateto falcate-triangular, (2-)4-10(-13) cm x (1.5-)2.5-5(-65) cm, base asymmetrical, apex acuteto obtuse, margins convex, leathery, glabrous,with numerous penucid gland dots, with (7-)8-12(-14) veins from the base. Inflorescence anaxillary raceme up to 7 cm long, simple or withone or two branches at the base, 7-13-flowered.Flowers bisexual, regular, greenish white to

yellowish; pedice1 4-8 mm long; sepals 4, near-Iy round, c. 5 mm long, glabrous, reflexed; pet-als absent; stamens 20-25, free, c. 7 min long;ovary superior, flattened obovoid, c. 3 mm long,glabrous, style c. 2 min long, glabrous, stigmabroad. Fruit a flattened, obliquely obovoid tokidney-shaped pod (2.5-)3-4.5(-6) cm x 2-2.5(-3) cm, narrowed at base, rounded at apex, withstyle remnants half to two-thirds of the wayalong the upper side, glabrous, straw-colouredwith minute, sunken, glandular flecks of dark-er brown, usually with raised reticulate vena-tion, indehiscent, I'Seeded. Seeds almost fillingthe fruit, kidney-shaped, compressed, marginflattened, surface with deep folds, pitted withnumerous reddish, sticky glands. Seedling withepigealgermination.

Other botanical information 0010phosper-mum comprises a single species. It has beenincluded in Hordwichio (a genus of one speciesin India) and this is supported by pollen inor-phology, chromosome numbers and phylogenet-IC analyses; in 2005 the name 0010phospermumhas been conserved against Hordwichi0. 0010-phospermum seems also to be related to Prto-

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 pin); 29: ves-tured pits; 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 ILm);42: mean tangential diameter of vessellumina100-200 F1m; 47: 5-20 vessels per square inn-limetre; (48: 20-40 vessels per square minime-tre); 58: gums and other deposits in heartwoodvessels. Tracheids and fibres: 61: fibres withsimple to minutely bordered pits; 66: non-septate fibres present; 70: fibres very thick-walled. Axial parenchyma: (76: axial paren-chyma diffuse); 79: axial parenchyma vastcen-tric; 80: axial parenchyma allform; 81: axialparenchyma lozenge-anform; (83: axial paren-chyma confluent); 86: axial parenchyma innarrow bands or lines up to three cells wide;89: axial parenchyma in marginal or in seem-ingly marginal bands; (91: two cells per paren-chyma strand); 92: four (3-4) cells per paren-chyma strand. Rays: 97: ray width I-3 ceUs;104: all ray cells procumbent; 1/5: 4-12 rays

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per min. Mineral inclusions: 136: prismaticcrystals present; 142: prismatic crystals inchambered axial parenchyma cells; (143: prts-matic crystals in fibres)co. A. Obeng, P. Baas & H. Beeckman)

Growth and development The Tootsystemof Golophospermum mopone is remarkablyshallow (30-120 cm deep) but very extensiveand well adapted to and conditions. The rootscan take up water from drier soilthan coinpet-ing grasses. The leaves fold together and hangstraight down under Intense sunshine or waterstress, producing very little shade which addsto the harshness of mopane woodlands. Treesshed most of their leaves gradually during thedry season and can be leafless for up to 5months, but mostly shorter. New leaves appearjust before or soon after the first rains, alt-hough flushing seems independent of the rainsin Namibia and Botswana. New leaves are

yellowish and soft, and turn green and leatherywith age. Flowering begins when trees are 5years old and usually occurs soon after newleaves have developed, between October andMarch, but can be erratic with no flowers at allfor several years, Fruits usually appear be-tween March and June. They are dispersed byrain-wash and wind, and over short distancesonly.Root suckers are commonly found around thebase of established trees. They are producedwhen trees are damaged by fire, drought, hu-mans or elephants. GOPpice growth can be vig-orous and sometimes forms almost impenetra-ble thickets. Mycorrhizae are often present onthe roots, but N-fixing nodules have not beenfound. However, a symbiosis of the roots hasbeen observed with a bacterium, causing grad-ualdegeneration of the roots and the formationof new ones, thus forming clusters of new roots.It has been suggested that these clusters mightbe considered as a primitive form of root nod-ules

Golophospermum mopone is not a pioneer spe-cies and does not invade new, disturbed areasRegeneration under existing trees occurs, but itIs uncertain to what extent. Thousands of seed-

lings are commonly found during the rainyseason, but saplings of I-5 years old are rarelyencountered. Stands are often even sized and

possibly of even age, with few small or youngtrees. In a trial in 10 permanent sample plotsin a mopane woodland in northern South kiri-ca, the survival of seedlings was observed fornearly 3 years. The highest seedling mortalityrate of 39% was recorded during the estab-

lishment period. Only 3% of the original seed-lings, estimated at 294,000 seedlings per ha,survived tinthe end of the test period. Standsof seedlings under the canopies of mother treeswere the first to die out completely, probablydue to competition. Growth rates are generallyslow. In northern South Africa, the age of atree with a hole diameter of 10 cm was esti-mated at 42 years, which was believed to be inline with an average age of full-grown trees of100-200 years

Ecology Golophospermum mopone is foundin hot, low-lying areas, especially dry rivervalleys, up to 1000 (-1300) in altitude, with amean maximum temperature of about 30'Cand a mean annual rainfall of 400-700 min,although it occurs in some localities in Namibiareceiving only 100 mm rainfall per year. It oc-curs in areas with alluvial or colluvialsoils andis also widespread in Kalahari sands. Whilegrowth is best in fertile slightly acidic, friableand permeable deep clay soils, it tolerates alka-line and poorly drained and temporarily water-logged soils. The lack of clay-rich soils with afair nutrient level may explain the absence ofGolophospermum mopone at higher altitudesIn the drier savanna woodlands of the central

part of southern Africa, it is often gregariousand dominant, occurring in almost pure standsknown as 'mopane woodland' or 'Golophosper-mum woodland'. In some areas the plants re-main stunted, forming 'mopane scrub'. Mopanewoodlands cover an area of more than 500,000kin2. The bark is resistant to fire. Slight frostto -3'C is tolerated

Propagation and planting Colophosper-mum mopone is easily propagated by seed,wildlings and cuttings. The weight of 1000seeds is about 500 g. Germination is easy andno pre-treatment is needed. The seeds germi-nate inside the pod which does not break open,but germination is more rapid and even whenseeds are removed from the pod. Seeds can besoaked in cold water for 24 hours to speed upgermination. With proper care, fresh seeds canattain a germination rate of 100%. Storagebehaviour Is orthodox. ; seeds can be stored forseveral years and still attain good germination.In Botswana seeds stored for 8 years stillhad agermination rate of 78%, but in a trial in Indiagermination failed completely after storage forthe same periodIn the nursery, it is recommended to sow seedsin flat seedling trays in clean or sterilized riversand. Seeds should be partly covered whensown in trays and kept moist. Deeply buried

seeds will not germinate. Germination normal-Iy takes I-2 weeks. Seedlings should be care-fully transplanted from seedbeds or germina-tion trays into nursery bags filled with a loamysoil with some sand and compost added. Seed-lings are susceptible to damping-off and shouldnot be watered too excessively. Initially theygrow slowly, but growth speeds up once theyhave reached a height of 20 cm. Seedlings aresensitive to frost and should be given pro^C-tion in the nursery during the cold season.

Management There are no reports of man-made Golophospermum mopone plantations inAfrica, except for plantings of a few or singletrees for shade in homesteads. Planting onfarmland is not common, but during forestclearing farmers often leave a few isolatedtrees to provide some shade during the crop-ping season

Seedlings and wildlings should be transplantedat the start of the rainy season to enable themto establish and survive without watering. of-ter planting out, seedlings are light demandingand susceptible to competition, necessitatlrigseveral weedings. Only under very dry condi-tions can seedlings compete well with otherspecies. They should also be protected fromlivestock

Pollarding and lopping is quite well tolerated,with rapid re-growth of new shoots. Treesmanaged for producing poles can be left togrow for up to 10 years and can then be pal-larded to a height of2 in

Diseases and pests Mopane is often deroli-ated by the larvae of the mopane moth(Gonimbrosio beltno). Large population out-breaks of the caterpillars occur once or twice a

November/December and Febru-lnyear

aryMarch, and large stands of Colophosper-mum mopo, ,e trees are then completely defoli-ated. The larvae of the psyllid Betroocizziomoponi also feed on mopane leaves, causingthem to curl. Termite attack has been observedon young seedlings. No important diseaseshave been recorded either in southern Africa orin the and regions of India where the species Isplanted.

Harvesting Because of the hardness of thewood, stems of Colophospermum mopone aredifficult to fell, especially with hand tools, andheavy-duty chain-saws are normally used. Oldand large trees often show severe heart rot

Yield Forest inventories have reported den-sities for mature Golophospermum moponetrees ranging from a few trees/ha in semi-andnorth-western Namibia to 480 treesA1a in

south-eastern Zimbabwe. Typical mopane treedensities are approximately 200-400 tree/ha.In northern South Africa the total standingbiomass of a mopane woodland has been esti-mated at 24 tma

Genetic resources 0010phospermum ino-porte is not under threat of genetic erosion be-cause it is widespread and common over largeareas. However, some parts of southern Africaexperience a decline in natural stands due tothe multiple uses of the species, the growingpopulation, forest clearing for agriculture, ex-cessive burning, growing elephant populationsand possibly changes in climate. Systematicgermplasm collection and specific conservationprogrammes do not exist, but there are smallcollections in botanical gardens in southernAfrica

Prospects Golophospermum mopone is high-Iy valued for its wood used in construction ofhouses and fences because of its great durabil-ity and resistance to termites, and as firewoodbecause of its excellent burning propertiesThese uses are likely to increase in importanceand may locally become a threat to mopanewoodlands. 0010phospermum mopone is a truemultipurpose tree, not only Important for itswood but also as source of medicine, forage andedible caterpillars. Protection measures anddomestication should be explored to attain sus-tainable exploitation of this species, which isone of the most characteristic indigenous treesof southern Africa. As an important medicinalplant, research is needed into Its active coin-pounds

Major references BOLza & Keating, 1972;Brummitt at a1. , 2007a; Coates Palgrave, 2002;Mojeremane & Lumbile, 2005; Palmer & Pit-man, 1972-1974; Roodt, 1998; Ross, 1977;Timberlake, 1995; Timberlake, 1999; van Wyk& Gencke, 2000

Other references Baerts & Lehmann,2012; Parishawe, 1972; Goldsmith & Garter,1981; Gondo at a1. , 2010; Henning & White,1974; Kozanayi & Frost, 2002; Lukhele & vanRyssen, 2003; Makhado, Potgieter & Wessels,2009; Mojeremane & Kgati, 2005; Mulofwa,SImute & Tengnas, 1994; Mushove & Makoni,1993; Mushove at a1. , 1995; Sinit, 2001; Styles& Skinner, 1997; Tietema, Merkesdal &Schroten, 1992; Tietema at a1. , 1991; vanDarnme, van den Byriden & Vernemmen, 1992;van den Byriden, Vernemmen & van Darnme,1992; van Wyk & van Wyk, 1997; Wessels,undated

Sources of illustration Brummitt at al. ,

CoLOPHosPERMUM 215

216 TIMBERS2

2007a; Coates Palgrave, 2002.Authors R. MeIusi& W. Mojeremane

CoLUBRiNADECIPiENs (Baill. ) Capuron

Protologue Adansonia, s6r. 2, a 127 (1966).Family RhamnaceaeSynonyms Macrorhomnus decipiens Baill.

(1874)Origin and geographic distribution Golu-

67ino decipiens is endemic to western Mada-gascar

Uses The wood, known as 'tratramborondreo'or 'malamasafoy' in Madagascar, is used forconstruction, parquet flooring, joinery, Interiortrim, railway sleepers and furniture. The barkrubbed in water is used as a soap substitute.

Properties The heartwood is dark brownwith faint streaks, and distinctly demarcatedfrom the sapwood. The texture is medium. Thewood is medium-weight to fairly heavy, with adensity of 700-850 kg/ina at 12% moisture con-tent, and hard. It usually air dries satisfactori-Iy. The rates of shrinkage during drying aremoderately high. Once dry, the wood is moder-ately stable in service. The heartwood is fairlydurable and moderately resistant to impregna-tion with preservatives

Botany Deciduousshrub or smallto medium-sized tree up to 20(-25) in tall; hole up to 65(-90) cm in diameter; bark surface smooth butscaly, grey to blackish with distinct greyishpatches; crown open, Irregularly spreading;twigs reddish brown hairy. Leaves opposite,simple and entire; stipules up to 6 mm long,early caducous; petiole I-2 cm long; bladeovate to ovate-lanceolate, (2.5-)6-12 cm x (1.5-)

2.5-6 cm, base obtuse to rounded or shallowlycordate, apex obtuse to acuminate, nearly gla-brous, with translucent dots, 3-5-veined fromthe base and additionally with a few pairs oflateral veins. Inflorescence a short, axillarycyme, with short peduncle, densely reddishbrown hairy. Flowers bisexual, regular, 5-meTous; pedice1 2.5-3.5 mm long; calyx lobestriangular, c. 2 mm x 2 min; petals hood-shaped, nearly 2 mm long; stamens free, eachstamen partly enveloped by a petal; disk flat,glabrous, covering the ovary; ovary for thegreater part superior, 3-celled, style c. 0.5 mmlong, with 3 short lobes. Fruit a nearly globoseto ovoid capsule 8-15 mm long, with short cupat base, glabrous, yellow, later becomingbrownish black, splitting into three I-seededparts, each opening with a slit. Seeds c. 5 minlong, slightly compressed, dark brown to black-ish, with small anI at base. Seedling with hy-pogealgermination001ubrirto decipiens grows moderately rapidly,with an average annual heightincrement of 45cm and a mean annual hole diameter incre-merit of 5 min. Trees are usually leafless fromJuly to October and they flower around De-cember. Fruitstake 45 monthsto ripen.Colubrtno comprises about 30 species occurringthroughout the warmer parts of the world. InMadagascar 6 species occur, of which one (Co-Iubrino OSiotico (L. ) Brongn. ) also occurs alongthe coasts of East Africa.

Ecology 001ubrirro decipiens occurs in drydeciduous forest, from sea-levelup to 600(-800)in altitude. The mean annual rainfall in thedistribution area is 300-700(-1000) min, with5-7(-9) dry months, and the mean annualtemperature is 25'C. Colubrino decipiens isoften found on more humid localities and ittolerates temporary inundation. It seems toprefer limestone soils.

Management Ripe fruits can be collectedfrom the ground. They should be immersed inwater for 3-4 days and subsequently dried inthe sun untilthey open. The 1000-seed weightis 2-3 g. The germination rate of seeds is 25-40%. Seeds can be stored for up to one yearwithout losing much of their germination POW-er. They are sown in seed beds or in pots, usu-ally in January, and seedlings are ready fortransplanting Into the field when they are oneyear old and about 50 cm tall. They are usuallyplanted bare-rooted and in the fullsun.

Genetic resources and breeding Golubri-rid decipiens is widespread throughout westernMadagascar and locally common, and there are

.

Golubrino decipiens - wild

L,

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..

no indications that it is in danger of geneticerosion.

Prospects Colubrino decipiens seems tohave fair prospects as a plantation tree and forenrichment planting in the dry forests of west-ern Madagascar. However, long cutting cyclesare probably needed to obtain holes largeenough to be suitable for sawn timber produc-tion

Major references Blaser at a1. , 1993; CFPF,2008

Other references Johnston, 1971; Fernerde in Bathie, 1950a; Schatz, 2001.

Authors R. H. M. J. Lemmens

COLDBRINAFARALAOTRA (H. Perrier) Capuron

Protologue Adansonia, s6r. 2, a 130 (1966)Family RhamnaceaeSynonyms Mocrorhomnus forumotro H. Per-

tier (1943).Origin and geographic distribution Colu-

brimo fordlootro is endemic to northern andeastern Madagascar.

Uses The wood, known as 'faralaotra' or'malemisalaza' in Madagascar, is in demand forconstruction, high-quality parquetflooring, andfor indoor as well as outdoor joinery. It has alsobeen used for railway sleepers, often after jin-pregnation. It is suitable for interior trim, shipbuilding, hydraulic works in fresh water, rumi-ture, toys and novelties. The wood is also usedas firewood and for charcoal production

Properties The heartwood is greenish yel-low, turning reddish brown upon exposure. It isdistinctly demarcated from the whitish andwide sapwood. The texture is medium

The wood is medium-weight to fairly heavy,with a density of 710-830 kg/ina at 12% ino1s-ture content. It should be dried with care be-

cause it splits easily. The rates of shrinkageare moderately high, from green to oven dry3.3-4.0% radial and 7.6-8.4% tangentialQuarter-cutting of logs before drying is recoin-mended. Once dry, the wood is moderately sta-ble to quite unstable in service. At 12% ino1s-ture content, the modulus of rupture is 152-198 N/mm2 modulus of elasticity 11,950-15,690 N/min2, compression parallel to grain66-83 N/min2, shear 6-8 N/min2, cleavage 11-22 N/min and Chalais-Meudon side hardness

3.6-9.1

The wood saws and works fairly easily. It fin-ishes well. Nailing, painting and staining showsatisfactory results. The wood is moderatelydurable, being resistant to fungi and moderate-Iy resistant to termites, but susceptible to ina-rine borers. The sapwood is liable to Lyetttsattack. The heartwood is moderately resistantto Impregnation with preservativesChemical analyses of the leaves, bark andwood led to the identification of some apor-phine alkaloids, phenolic acids and flavonoidsThe navonoid aglycones kaempferol, quercetinand myricetin have also been demonstrated, aswell as the navonol-0-triosides faralatroside

and faratroside, which are derivatives fromkaempferoland quercetin, respectively

Botany More or less evergreen shrub orsmall to medium-sized tree up to 25(-30) intall; hole branchless for up to 18 in, usuallystraight and cylindrical, up to 90(-120) cm indiameter; twigs reddish brown hairy. Leavesalternate, sometimes nearly opposite, simple;stipules small, early caducous; petiole 2-6 mmlong; blade ovate to ovate-lanceolate or obo-vate, 3-11 cm x 2-7 cm, base rounded, some-times shallowly cordate, apex obtuse to short-acuminate, margins entire to slightly wavy,membranous, nearly glabrous, with translu-cent glands below, 3-veined from the base andadditionally with 2-3 pairs of lateral veinsInflorescence a short, axillary cyme, with veryshort to fairly long peduncle, densely reddishbrown hairy, 3-40-flowered. Flowers bisexual,regular, 5-merous, white, fragrant; pedice1 2-8min long; calyx lobes triangular, c. 1.5 mm x1.5 mm; petals hood-shaped, c. I min long;stamens free, each stamen partly enveloped bya petal; disk flat, glabrous, covering the ovary;ovary for the greater part superior, 3-celled,style c. I min long, with 3 short lobes. Fruit anearly globose capsule 8-10 mm in diameter,

CoLUBRiNA 217

.

Golubrirto foyer lootro - wild

..

o

,^

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218 TIMBERS2

with short cup at base, glabrous to slightlyhairy, becoming bluish black, splitting intothree I-seeded parts, each opening with a slitSeeds c. 2.5 mm long, slightly compressed,black, with smallarilat baseGolubrino comprises about 30 species occurringthroughout the warmer parts of the world. InMadagascar 6 species occur, of which one (Co-Iubrino OSiotico (L. ) BTOngn. ) also occurs alongthe coasts of East Africa.

001ubrinu orborescens (Mill. ) Sarg. is a smalltree up to 8(-30) in tall, native of Middle Amer-ICa and the Caribbean and occasionally plantedas ornamental shade tree in Africa. The dura-

ble wood is mainly used for posts and piles. Intropical America, bark decoctions are used intraditional medicine to treat diabetes, hyper-tension and stomach disorders. Leaf decoctions

are applied in a bath against fever, skin coin-plaints, haemorrhoids and joint pains, and aretaken to treat gonorrhoea. Leaves and barkrubbed in water are used as a substitute of

soap.

Golubrinoforolootru is variable, and some sub-species and varieties have been distinguished,based on leafshape and size, number offlowersin the inflorescence, and hairiness of the fruit

Ecology Golubrino fordlootro occurs in dense,humid evergreen forest, from sea-level up to1000(-1700) in altitude

Genetic resources and breeding Colubri-rid fordlootro is fairly widespread and locallycommon, and there are no indications that it isin danger of genetic erosion

Prospects Too little information is availableto assess the prospects of Golubrino foro100troas an economicalIy important tlinber tree withsustainable production

Major references Blaser at a1. , 1993; Bolza& Keating, 1972; CFPF, 2008; Gu6neau, Bedel& Thie1, 1970-1975; Takahashi, 1978

Other references Bedolla, 1997; Gu6neau,1971a; Gumaudeau at a1. , 1976; Gumaudeau ata1. , 1981; Johnston, 1971; Parant, Chichignoud& Rakotova0, 1985; Ferner de in Bathie, 1950a;Schatz, 2001

Authors L. FA. Oyen

tree (En). Flamboyant d'avril, gloire de Golville(Fr).

Origin and geographic distribution Goluil-Ieo rocemoso Is endemic to western Madagas-car, where it occurs in the north near kiltsir-ariana, and from the Bemaraha Plateau to thesouth

Uses The wood, known as 'sarongaza', isused for posts, carpentry, shuttering, fencesand veneer. It is suitable for light joinery, inte-nor trim, furniture, boxes and crates. The holesare used to make dug-out canoes. Goluil!eorocemoso is planted as an ornamental treebecause of its bright orange flowers and dell-cate foliage, also in several tropical countriesoutside Madagascar.

Properties The wood is yellowish white togreyish yellow. The grain is straight, texturecoarse. The wood is moderately lightweight,with a density of 410-580 kg/ms at 12% inois-ture content, and soft. It air dries rapidly andwithout much defects when it is properlystacked. The rates of shrinkage are moderate,from green to oven dry about 3.8% radial and7.3% tangential. At 12% moisture content, themodulus of rupture is about 124 N/min2 inodu-Ius of elasticity 18,300 N/min2 compressionparallel to grain 40 N/min2 and Chalais-Meudon side hardness 2.8. The VeneerIng

properties are satisfactory. The wood is notdurable, but fairly easy to impregnate withpreservatives

Botany Deciduous, small to medium-sizedtree up to 20(-30) in tall; bole usually cylindri-cal and straight, sometimes slightly sinuous,up to 90(-100) cm in diameter; bark surfacefinely grooved, peeling off in thin strips, palegrey, with many lenticels, inner bark green;

CoLVILLEARACEMOSABojer

Protologue Bot. Mag. 61:t. 3325-3326 (1834)Family Caesalpiniaceae or, eguminosae - Cae-

salpinioideae)Chromosome number2n = 26.28Vernacular names ColviUe's glory tree, whip

.

Goluilleo racemoso - wild

..

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..

crown elongate, with Irregularly spreading orpendulous branches. Leaves alternate, hipin-nately compound with (6-)9-16 pairs of pinnae;stipules small, caducous; petiole 3-8.5 cm long,swollen at base, rachis up to 30 cm long,grooved and greyish hairy; leaflets opposite,numerous, nearly sessile, oblong, (0.5-)I-2 cmlong, slightly asymmetrical, short-hairy. Inno-rescence a large, terminal panicle c. 50 cmlong, with pendulous branches up to 40 cmlong, short-hairy, many-flowered. Flowers bi-sexual, zygomorphic, 5-merous, bright orangewith orange-red petals; pedice1 0.5-I cm long;calyx 2-2.5 cm long, leathery, with 4 shortlobes in lower part of flower and I long lobe inupper part; petals free, the lower one up to 1.5cm long with strongly mrolled margins, lateraltwo rhombic, up to 1.5 cm long, upper two nar-rowly rhombic, up to 3 cm long; stamens 10,free, 2.5-4.5 cm long, curved outwards from thetop of the flower; ovary superior, elongate, flat-tened, c. 1.5 cm long, I-celled, style slender,4.5-5 cm long. Fruit a narrowly oblong, flat-tened, pendulous pod 25-35 cm x 4-7 cm, gla-brous, dehiscing along one side, 12-20'seededSeeds oblong-obovate, flattened, c. 1.5 cm x Icm, pale brown inottled with darker brownSeedling with hypogealgerminationYoung trees grow fairly rapidly. A mean annu-al hole diameter increment of 7-14 mm has

been reached during the first 10 years afterplanting. 0010illeo rocemoso flowers from De-cember to March(-April). The flowers, whichproduce abundantly nectar, are pollinated bysunbirds. They are often eaten by lemursFruits ripen about 6 months after flowering001ui!led comprises a single species. It is relat-ed to Delonix and Lemuropisum

Ecology Goluilleo racemoso occurs in decid-uous woodland and seasonalIy dry forest onsandy soils, up to 300 in altitude. The meanannual rainfall in its area of distribution is upto 700(-1200) mm, with 7-9 dry months, andthe mean annual temperature 24.5-27'C.

Management The fruits are collected fromthe tree or the ground when they have becomedry and blackish, from May to July. The seedsare extracted manually and non-viable seedsare separated by floating in water. Viable seedsare dried in the sun. The weight of 1000 seedsis 275-400 g. The seeds can be stored for up to4 years, when still a germination rate of 50-70% can be reached. It is recommended to

submerge them in boiling water before sowing.When seedlings are raised in the nursery, theycan be planted into the field after 6-12 months

when they are 50-100 cm tall. Clipping thelateral roots is advised. Direct sowing into thefield is also practised. Propagation by cuttingshas been successful

Genetic resources and breeding Coluilleorocemoso is included in the IUCN Red List,where it is considered to be at lower risk, alt-hough near threatened. It is locally common,but its natural habitat of deciduous woodland

is strongly fragmented.Prospects Tests with Colonleo rocemoso in

Madagascar showed positive results. The woodfulfils the requirements for technical pro-cessing and has characteristics rendering itsuitable for economic exploitation, e. g. for ve-neer production. Colu, !Ieo rocemosu has goodcolonizing potential, and reforestation can beachieved by direct sowing. As such, it seems tohave good potential for reforestation in regionswith seasonalIy dry forest in Madagascar. Itdeserves wider application as an ornamentaltree.

Major references Blaser at a1. , 1993;CFPF, 2008; du Puy at a1. , 2002; Rakotovao atal. , en preparation; Randrianas010, 1992a

Other references du Puy, Phillipson &Rabevohitra, 1995; Randrianasolo at a1. , 1996;World Conservation Monitoring Centre, 1998

Authors R. H. M. J. Lemmens

COMBRETmi 219

COMBRETUMIMBERBEWawra

Protologue KaiserI. Akad. Wiss. Wien, Math. -Naturwiss. in. , Anz. 38: 556 (1860)

Family CombretaceaeVernacular names Leadwood (En).Origin and geographic distribution Coin-

I.

Combretum tinberbe - wild

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220 TIMBERS2

67etum jinberbe occurs from southern Tanzania

south to northern Namibia, Botswana, Zimba-bwe, Mozambique and northern SouthAfrica

Uses The wood is used for fence poles, mineprops, railway sleepers, mortars, walking sticks,inlay work, toys and turnery. Because of itsdurability, it is commonly used for the mainsupporting poles of huts. It Ts popular for sculp-ture and lathe work, whereas it is also used tomake heavy, extremely durable furniture. Thewood is favoured for use as firewood and for

charcoal production; it burns slowly and withgreat heat. The ash from the wood has a highlime content, and is sometimes used as tooth-paste and as a substitute of whitewash to deco-rate walls of houses.

Several plant parts are used in traditional me-dicine. Powdered roots or leaves and decoctions

from roots and leaves are taken to treat diar-rhoea. On the other hand, a leaf decoction isapplied as an enema to treat constipation. Aroot Infusion is drunk to treat schistosomiasis.

Bark powder is applied externalIy against Iep-rosy. Tea made from the roots or leaves isdrunk to treat coughs and colds, and the smokeof burnt leaves is inhaled for the same PUTpos-es. A leaf decoction is drunk to relieve chest

pain. A root maceration is taken to treat stom-ach-ache. The roots are used to treat infertilityin women. The tree yields an edible gum. Thebark has been used for tanning leather. Thefoliage is eaten by livestock, although it report-edly contains only 4% protein. The tree isclaimed to have protective power, and severalplant parts are used in ritual ceremoniesCombretum jinberbe is useful as an ornamentalshade tree

Production and international trade In

southern Africa Combretum tinberbe is in highdemand in the carving industry, together withBereftemto spp. , 01eo europoeo L. subsp. cuspi-doto Ovall. ex G. Don) Gif. and Spirostochysof neono Sond. In Mozambique the wood isclassified as first class for carving. In Tanzaniait has replaced the wood of Bobgunnio mudo-gascortensis (Desv. ) J. H. Kirkbr. & Wiersema.In 2003 the official harvest was 3900 in3 oflogs, butthen a nation-wide ban on the harvestwas declared

Properties The heartwood is dark brownand distinctly demarcated from the thin, yel-low-brown sapwood. The grain is straight, tex-ture fine. The wood is very heavy, with a densi-ty of about 1200 kg/ms at 12% moisture con-tent. It is difficult to work, rapidly bluntingcutting edges, but it takes a very nice lustrous

polish. The turning properties are excellentThe wood is very durable, even in contact withthe ground.Acetone and ethylacetate extracts of the leavesof Combretum jinberbe showed pronouncedanti-inflammatory activity in the cyclooxygen-ase-I bioassay, whereas an aqueous extractshowed activity against Schistosomo hoemoto-bium. The pentacyclic triterpene jinberbic acidhas been isolated from the leaves; this coin-pound showed potent antibacterial activityagainst Mycobacterium fortuitum and Stophy-IOCoccus oureus. Another pentacyclic triterpeneisolated from the leaves, 10.23-dihydroxy-12-o1eanen-29-o1c acid-3^-0-2.4-di-acetyl-L-rhamno-pyranoside, showed strong antibacterial activi-ty agalnst Staphylococcus oureus and ESChe-richio coli, as well as strong inhibition of 30-hydroxysteroid dehydrogenase and moderatecytotoxic activity against some human cancercelllines. In tests the wood ash showed goodresults in managing cowpea weevils in storedcowpea seeds

Adulterations and substitutes In South

Africa the wood of Combretum tinberbe is

commonly used to supplement the wood of0010phospermum mopone (Benth. ) J. Leonardas the main constituent of palisade fences.

Description Deciduous shrub or small tomedium-sized tree up to 20(-30) in tall; bolecommonly crooked, up to 100 cm in diameter;bark surface whitish to pale grey or dark grey,with deep longitudinal furrows and irregulartransverse cracks; crown spreading, roundedand rather open; twigs glabrous, sometimesspiny at tip. Leaves opposite, simple and en-tlre; stipules absent; petiole up to I cm long;blade elliptical to oblong-elliptical or elliptical-obovate, 2.5-8.5 cm x I-3 cm, curieate torounded at base, rounded to obtuse at apex,often with short point, papery to slightly Ieath-ery, densely silvery scaly, pinnately veinedwith 3-7 pairs of lateral veins. Inflorescence anaxillary or terminal spike up. to 5(-10) cm long;main axis scaly. Flowers bisexual, regular, 4-merous, yellowish, densely scaly, scented, ses-SIIe; receptacle consisting of 2 parts, lower partc. 2.5 min long, upper part indistinct, nearlyflat; sepals ovate-triangular, c. 1.5 mm long;petals free, obovate to spatulate, c. I mm long,glabrous; stamens 8, free, c. 2 min long; ovaryinferior, I-celled, style c. 2 min long, denselyglandular at base. Fruit a 4-winged nut, broad-Iy ovoid to nearly orbicular in outline, 1.5-2 cmlong, with 2-3 min long stipe and c. 7 mmbroad wings, pale yellowish green, silvery

3

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wood not ring-porous); 46: s 5 vessels persquare minimetre; 58: gums and other depositsin heartwood vessels. Tracheids and fibres: 60:

vascular/vasicentric tracheids present; 61: fi-bres with simple to minutely bordered pits; 66:non-septate fibres present; 69: fibres thin- tothick-walled. Axial parenchyma: (76: axial pa-renchyma diffuse); (79: axial parenchyma va-siCentric); 80: axial parenchyma allform; 81axial parenchyma lozenge-anform; (89: axialparenchyma in marginal or in seemingly mar-ginal bands); (92: four (3-4) cells per paren-chyma strand); 93: eight (5-8) cells per paren-chyma strand. Rays: 97: ray width I-3 ceUs;109: rays with procumbent, square and uprightcells mixed throughout the ray; 1/5: F12 raysper mm; (1/6: z 12 rays per min). Mineral in-clusions: 136: prismatic crystals present; 137:prismatic crystals in upright and/or square raycells; (138: prismatic crystals in procumbentray cells); (141: prismatic crystals in non-chambered axial parenchyma cells).re. 06tienne & P. Baas)

Growth and development Combretum jin-be76e trees grow very slowly. Under naturalconditions, mean annual diameter Incrementrates of only 0.3 min to 2 min have been rec-orded, but under optimal conditions in cultiva-tion young trees may reach 6 in tallin 15

Combretum jinberbe flowers from No.years

veinber to March, and fruits ripen from Febru-ary to August. The fruits are dispersed bywind, up to a distance of 50 in from the parenttree.

Combretum Imberbe trees can become very oldRadiocarbon dating showed that some speci-mens were over 1000 years old. Dead trees canremain uprightfor as much as 80 years'

Ecology Combretum tinberbe occurs in openwoodland and wooded savanna, on a wide van-ety of soils from sandy soils to limestone out-crops, also on alluvial soils and black cottonsoils, but only occasionally on heavy clay soils,from sea-level up to 1000 in altitude. It is mostcommon along rivers. Combretum Jinberbe hasan important ecological value for animal spe-cies, e. g. as fodder for desert elephants in Na-inIbia and as nesting site for Ruppell's parrotand hornbills

Propagation and planting One kg containsabout 11,500 fruits, with the seed accountingfor about 45% of the fruit weight. Seeds shouldbe sown while still fresh. They should besoaked in water for a few hours before beingpressed into seedling trays filled with riversand. After they have been covered with a thin

4

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Combretum tinberbe - I, tree herbit, . 2, port ofledb, twig, . 3, flower in longitudinol section, . 4,frui'Redrown grid odopted bylshoh Syomsudin

COMBRETUM 221

I

,

scaly, indehiscent, I-seeded. Seedling withhypogeal germination, with stalked, nearlyorbicular cotyledons.

Other botanical information Combretumis a very large genus, comprising about 250species and distributed worldwide in the trop-ics and subtropics. About 140 species occur intropical Africa

Anatomy Wood-anatointcal description CAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; (7:vessels in diagonal and/or radial pattern); (9:vessels exclusively solitary (90% or more)); 13simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-nab; 25: intervessel pits small (4-7 prn); 26intervessel pits medium (7-10 prn); 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 prn;(45: vessels of two distinct diameter classes,

222 TIMBERS2

layer of sand, the seeds should be kept moist.They germinate in I-2 weeks, but very lowgermination rates have been recorded (3-5%),as well as high mortality among seedlings. Theseedlings can be transplanted into nurserybags after development of the second leaf.

Management The trees can be managed bycoppicing. The number of shoots produced fromcut stems is negatively correlated with theheight at which the trees are cut, but shortershoots are produced when the tree is cut closeto the ground. A cutting height of I in appearsmost advantageous. In South Africa the major-ity (nearly 80%) of the Combretum jinberbetrees has hole diameters of less than 20 cm. InTanzania the minimum diameter limit for har-vesting has been fixed at 24 cm.

Diseases and pests Seeds seem to be liableto attacks by insects and rodents. Seedlings areapparently often destroyed by large herbivoressuch as antelopes and elephants

Harvesting Good equipment with hardenedand sharp cutting edges is needed to fell thetrees and to saw and work the wood.

Genetic resources Combretum tinberbe iswidespread and at least locally common. It istherefore not easily liable to genetic erosionHowever, studies in Malawi have shown thatthe wood is in high demand for charcoal pro-duction, and that many trees are felled for thispurpose. In South Africa Combretum tinberbetrees of over 20 cm in hole diameter are selec-

tively felled on a large scale for their timber,whereas smaller-sized trees are commonly cutfor firewood. The extent of exploitation appearsto be unsustainable in many regions with highhuman population pressure within the distri-bution area of the species.

Prospects There is a lack of knowledge onnatural regeneration of Combretum tinberbeand on threats to young trees, and the responseof Combretum tinberbe to harvesting pressureunder different environmental and arithropo-genic Influences is poorly understood. Researchon these aspects Is needed to develop modelsfor sustainable exploitation. Protection of thisslow-growing tree is becoming important, butIs certainly worthwhile considering its manyservices to humans and animals.

Major references ingeh at a1. , 2007; Exell,1978; Herrinann, Milton & Seymour, 2003;Mtsweni, 2006; Palmer & Pitman, 1972-1974;van den Eynden, Vernemmen & van Damme,1992; van Wyk & Gencke, 2000; Vogel & EUls,2005.

Other references Coates Palgrave, 1983;

Gelfand at a1. , 1985; Javaid & Ramatlakapela,1995; Katerere at a1. , 2003; MCGaw at a1. , 2001;Milledge, Gelvas & mirends, 2007; Neuwinger,2000; Steenkamp, 2003; Wickens, 1973; Willi-amson, 1955

Sources of illustration Coates Palgrave,1983; Exe11, 1978; van Wyk, 1972-1974.

Authors R. H. M. J. Lemmens

COMBRETUMSCHUMANNllEngl.

Protologue PITanzenw. OSt-Mrikas 0 289(1895)

Family CombretaceaeChromosome number2n=26

Synonyms Combretttm errg!eriSchinz (1901).Vernacular names Forest tree combretum,

sand bushwillow (En). Mgurure, inpera-inwitu,ingongolo (Sw).

Origin and geographic distribution Coin-bretum schumonnii occurs from southern So-mana and Kenya south to Namibia, Botswana,Zimbabwe, Mozambique and northern SouthAfrica.

Uses The wood is used for heavy construc-tion, flooring, pestles, combs and carvings. It issuitable for interior trim, ship building, vehiclebodies, mine props, handles, ladders, sportinggoods, musical instruments, pulley blocks, toys,novelties, precision equipment and turnery. InNamibia the branches of Combretum schu-

morinii shrubs are used for making bows forchildren to practice hunting; they are not suit-ed for bows for adults because they break tooeasily. The wood is also used as firewood andfor charcoal productionBark and leaves are used in traditional medi-

.

Combretum schumonnii- wild

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cine. Pulp made from the root bark is appliedto swellings. Heated leaves are applied to thechest to treat pneumonia, and the leaves arealso used to treat epilepsy and headache. Theflowers are a source of nectar and pollen forhoney bees. It has been reported that the treegives mulch of good quality

Production and international trade In

Kenya Combretum schumonnii belongs to thefour most commonly used species in the carv-ing industry, together with Do16ergio melono-xylon Gulll. & Perr. , Brochyloeno hutllensisO. Hoffm. and 01eo europoeo L. subsp. cuspido-to (Wall ex G. Don) Gif. However, there is noinformation on volume and price of wood intrade

Properties The heartwood is dark purplishbrown when freshly sawn, slowly darkening tonearly black after years, and distinctly demar-cated from the narrow, whitish sapwood. Thegrain is straight, texture moderately fine tofine, but not always evenThe wood is heavy, with a density of 1040-1/20 kg/ina at 12% moisture content. The woodair dries well but slowly, with little degradeThe rates of shrinkage are rather low, fromgreen to 12% moisture content about I% radialand 2-3% tangential. The wood is easy to workas long as sharp tools are used, and it takes avery nice polish. The turning propertles areexcellent. Pre-boring is necessary for nailingThe wood joints well. It is fairly durable, evenin contact with the ground, and resists termiteand marine borer attacks quite well. It cannotbe impregnated with preservatives

Adulterations and substitutes The woodof Combretum schumonnii has some resem-

blance to that of Do16ergio melonoxylon, whichis also in high demand for carving. Ebony, thewood of Diospyros spp. , is also used for similarpurposes as that of Combretum schumonrrii.

Description Deciduous shrub or small tomedium-sized tree up to 20(-30) in tall; holeoften crooked or fluted, up to 60(-80) cm indiameter; bark surface pale yellowish brown tobrown, scaling off in large, elongate flakes,inner bark pinkish with paler streaks, exudinga thick whitish sap; crown open; twigs slender,nearly glabrous. Leaves opposite, simple andentire; stipules absent; petiole up to I cm long,slender; blade elliptical to oblong-elliptical,ovate or obovate, (2-)6-12(-15) cm x (I-)2-4.5(-7.5) cm, curieate to rounded at base, roundedto obtuse or short-acuminate at apex, papery,slightly warty and scaly, pinnately veined with3-8 pairs of lateral veins. Inflorescence a short

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COMBRETUM 223

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Combretttm schumonrtii - I, tree hobit, . 2, legfi,twig, ' 3, flower in longitudinol section, . 4, fruit.Redrown und adopted by AChmod Sotiri Nur-hornon

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axillary spike up to 1.5 cm long, sticky. Flowersbisexual, regular, 4-meTous, pale yellow, scent-ed, sessile; receptacle consisting of 2 parts,lower part I-1.5 mm long, upper part 1.5-2.5min long, with 4 pouch-like swellings; sepalsshallowly triangular, indistinct; petals free,nearly orbicular, 1.5-2 mm long, glabrous;stamens 8, free, c. 5 mm long; ovary Inferior, I-celled, style c. 5 mm long. Fruit a 4-winged nut,ovoid to nearly orbicular in outline, 3-4 cmlong, with I-2 cm long stipe and c. I cm broadwings, green turning pale brown, lridehiscent,I-seeded. Seedling with epigeal germination,with sessile kidney-shaped cotyledons

Other botanical information Combretttm

is a very large genus, comprising about 250species and distributed worldwide in the trop-ICs and subtropics. About 140 species occur intropical Africa. In several African floras, Coin-bretum errgleri Schinz from the western part ofsouthern Africa has been included in the syn-onyiny of Combretum schumonnii, althoughthe former is only a shrub up to 4 in tall withsmaller leaves and fruits. More research is

needed to elucidate the status of Combretum

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224 TIMBERS2

errg!eri; it might be a separate taxon (either onspecific or infraspecific level), or the differencesmight be due to climatic conditionsCombretum kroussii Hochst. occurs in eastern

South Africa and Swaziland, but a few speci-mens have been collected in southern Mozam-

bique. It is a shrub or smalltree up to 12 intall. Its yellow wood with usually straightgrain and fine texture is suitable for flooring,joinery, mine props, ship building, vehicle bod-ies, furniture, cabinet work, handles, ladders,sporting goods, toys, novelties, agriculturalimplements and turnery; it is also used asfirewood. At 12% moisture content, the densityof the wood is about 770 kg/ina modulus ofrupture 110 N/min2, modulus of elasticity13,230 N/mm2 and compression parallel tograin 63 N/min2. Although the wood is hardand tough, it is moderately easy to work; itpolishes well. It is only moderately durable,and resistant to Impregnation by preservativesThe roots of Combretum kroussii are used intraditional medicine to treat wounds, and as ananodyne, tonic and appetite stimulant. Thepliable young stems are used in basket making

Anatomy Wood-anatoiincal description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(7: vessels in diagonal and/or radial pattern); 9:vessels exclusively solitary (90% or more); 13:simple perforation plates; 22: intervessel pitsalternate; 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; 41: mean tan-gential diameter of vessellumina 50-100 pin;(45: vessels of two distinct diameter classes,wood not ring-porous); 47: 5-20 vessels persquare minimetre; 58: gums and other depositsin heartwood vessels. Tracheids and fibres: 60:vascular/vasicentric tracheids present; 61: fi-bres with simple to minutely bordered pits; 66:non-septate fibres present; 70: fibres verythick-walled. Axial parenchyma: 83: axial pa-renchyma confluent; 92: four (3-4) cells perparenchyma strand. Rays: 97: ray width I-3cells; (104: all ray cells procumbent); (109: rayswith procumbent, square and upright cellsmixed throughout the ray); 1/5: 4-12 rays permin; 1/6: Z 12 rays per mm. Mineral inclu-SIons: 156: crystals in enlarged cells.(P. Derienne & P. Baas)

Growth and development In Namibia theflowers only last for about 3 weeks, from inid

September until the beginning of October,when the shrub is nearly leafless. The flowersare pollinated by insects such as honey bees.Fruits ripen about 7 months after flowering.Monkeys have been reported to feed on thefruits and may serve as seed dispersers.

Ecology Combretum schumonniioccurs in awide range of habitats, from lowland rainforestto riverme forest, dry deciduous forest andwooded savanna, from sea-level up to 1150(-1600) in altitude. In southern Africa it is coin-monly found on Kalaharisand and rocky out-crops

Propagation and planting Seeds shouldbe sown while still fresh. Germination can be

fast and the germination rate high. Wholefruits can be stored for a few weeks only with-out affecting the germination capacity of theseed. It has been recommended to strip off thewings of the fruit before sowing. Trees mayproduce root suckers that can be used for prop-agation.

Management The trees can be managed bycoppicing and lopping.

Handling after harvest The holes are usu-ally short and commonly fluted, which reducesthe possibilities of usage for sawn wood.

Genetic resources Combretum schumonniiis widespread and at least locally common.This makes that it is not easily liable to genetic

However, in some regions, e. g. inerosion

coastal regions in Kenya, Combretum schu-monn, i trees are selectively felled for theirwood that is highly valued for carving, therebyaffecting populations of larger-sized treesStudies on the genetic vanability are warrant-ed, covering populations all over the distribu-tion area; this may also elucidate the status ofCombretum errgleri

Prospects There is no information on growthrates and natural regeneration of Combretumschumonnii. Research is needed to developmodels for sustainable harvesting of its woodIt does not seem to be a serious candidate for

planting with the objective of economicalIyviable timber production because the hole isoften of too small size and poor shape, and be-cause growth is probably slow

Major references Bolza & Keating, 1972;Bryce, 1967; Exe11, 1978; Lovett at a1. , 2007;Maundu & Tengnas (Editors), 2005; Mbuya ata1. , 1994; Wimbush, 1957.

Other references Beentje, 1994; CoatesPalgrave, 1983; Dale & Greenway, 1961; Ko-kwar0, 1993; Leger, 1997; Msangi, 1991; Neu-winger, 2000; Takahashi, 1978; Tee1, 2004;

Thulin, 1993a; Wickens, 1973.Sources of illustration Exe11, 1978; Maun-

du & Tengnas (Editors), 2005Authors D. Dongock Nguemo

CoPAIFERAMiLDBRAEDllHarms

Protologue Notizbl. Bot. Gart. Berlin-Dahlema 147 (1922).

Family Caesalpiniaceae or, eguminosae - Cae-salpinioideae)

Origin and geographic distribution GOPoi-Iero mildbroediiis distributed from south-eastern Nigeria eastward to the Central Mri-can Republic and southward to Gabon and DRCongo.

Uses The wood of GOPoi/;?ro mildbroedii, trad-ed as 'andem-evin6' and 'anzem noir' and as

'etimoe' together with that of GOPoifero soli-houndo Heckel, is mainly used for veneer. It issuitable for heavy construction, joinery, shipbuilding, vehicle bodies, furniture, musicalinstruments, boxes, crates, carving and drain-ing boards. It is also used as firewoodPygmy people in the Central African Republicuse the bark in perfume. The fruit contains aresin that is inflammable. In the Central Mri-

can Republic, fruits dried in the sun are usedfor lighting fires and they are burned in lampsand as candles. In traditional medicine in Ni-

gena, the fruits and seeds are used as purga-tive, stimulant and diuretic. In the CentralAfrican Republic, the bark is eaten to increasevirility, bark decoctions are drunk or used inenemas or fumigations for the treatment ofstomach problems, and a decoction of the barkof GOPoifero mildbroedii and that of Anonidi-

urn monnii(011v. ) Engl. & Diels is taken tocure female sterility.

Production and international trade Accord-

ing to ATIBT statistics, Gabon exported 530 inaof logs in 2001, 685 ms in 2002, 235 main 2003,225 ing in 2004 and 265 ms in 2005. Congo ex-ported 150 ina of logs in 2004, and 80 ms of logsand 100 ing of sawn wood in 2006

Properties The heartwood is reddish brown,often with decorative streaks, and distinctlydemarcated from the white or greyish white,up to 10 cm wide sapwood. The grain isstraight, texture medium to fine and even. Thewood contains resin

The wood is moderately heavy, with a densityof (600-)750-850 kg/ms at 12% moisture con-tent. Drying should be done slowly and withmuch care to prevent severe checks and distor-tion. It is advisable to quarter-saw the woodbefore drying. The shrinkage rates are moder-ately high, from green to oven dry 6.0% radialand 8.5% tangential. Once dry, the wood ismoderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 127-155 N/mm2, modulus of elasticity10,800-15,300 N/mm2 compression parallel tograin 64-70 N/mm2, cleavage 21.5-24.5 N/minand Chalais-Meudon side hardness 40-60.

The wood works well with both machine and

hand tools. It planes very well, and gives anattractive finish. It takes nails well. The wood

is only moderately durable, being susceptible tostain and termite attack. The sapwood is sus-ceptible to Lyetus attack. The heartwood isextremely resistant to impregnation with pre-servatives

The wood contains 40% cellulose, 30.5% 11gnin,18% pentosan and 0.5% ash. The solubility isabout 5.5% in alcohol-benzene, 2.7% in hotwater and 27.1% in a I% NaOH solution. Wood

fibre cells in samples from DR Congo were onaverage 1.4 min long and 20 prn wide, with acell wanthickness of 4 I'm and a lumen widthof 12.5 Urn

Botany Evergreen, large to very large treeup to 60(-65) in tall; hole branchless for up to30 in, straight and cylindrical, up to 200 cm indiameter, sometimes thickened and grooved atbase, without buttresses; bark surface smoothto slightly fissured, dark grey, inner bark fi-brous, hard, brittle, pinkish yellow, turningbrown on exposure, aromatic; crown umbrella-shaped; twigs hairy but soon becoming gla-brous. Leaves alternate, pinnately compoundwith 20-40 leaflets; stipules caducous; petioleand rachis 13-25 cm long, grooved above; petio-

CoPAiFERA 225

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GOPotfero mildbroedii- wild

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Iules I-2 mm long, hairy; leaflets alternate ornearly opposite, oblong, 1.5-4(-6) cm x I-2(-2.5) cm, base rounded and asymmetrical, apexrounded to notched, glabrous except for theinIdvein below, with translucent dots, pinnate-Iy veined with 14-20 pairs of lateral veinsInflorescence a terminal or axillary raceme 8-30 cm long, with lateral spike-like branches 2-5 cm long, reddish hairy. Flowers bisexual,zygomorphic, white; pedicelc. I min long, with2 caducous bracteoles up to 2.5 mm long; sepals4, ovate-lanceolate, 3-4 min x I-2.5 min, Ibroader than other 3, acute, reddish hairy;petals absent; stamens 9-10, free, 5-6 mmlong; ovary superior, sessile, hairy at one side,I-celled, style c. 2.5 mm long. Fruit an oblique-Iy oblong to elliptical or nearly globose pod 3-5cm x 2.5-3.5 cm, leathery, glabrous, brownishblack, often shiny and covered with resin drop-lets, dehiscent with 2 valves, I'Seeded. Seedsoblong, slightly flattened, I-3 cm x I-1.5 cm,black, with thin, orange-red, waxy anI. Seed-ling with epigeal germination; hypocoty1 3-7.5cm long, epicoty1 4-9 cm long; first 2 leavesopposite.

In Nigeria GOPoifero mildbroedii fruits ripenin FebruaryGOPoifero comprises about 40 species, of whichabout 35 occur in tropical America, 4 in Africaand I in tropical Asia. GOPoifero seems closelyrelated to Bothioeo, Deturium, Stridoro andTessmonnio

Ecology GOPoifero mildbroedit occurs inrainforest and gallery forest, also in secondaryforest, rarely in periodically inundated localI-ties.

Management In the Central African Repub-11c and Gabon, the minimum hole diameterallowed for harvesting is 70 cm. Logs do notfloat well in water and cannot be transportedby river. They should be extracted rapidly fromthe forest or treated with a preservative toavoid attacks by fungiand insects

Genetic resources and breeding GOPot-Iero mildbroediiis widespread and locally fair-Iy common within its distribution area. Thereare no indications of overexploitation and Co-potfero mildbroedtt does not seem to be threat-ened by genetic erosion.

Prospects In view of the limited durabilityof the wood, it is more suitable for interior thanfor exterior use. Small quantities of GOPoiferumildbroedii timber are traded on the interna-

tional market, mainly for veneer production.Although it is locally fairly common, infor-mation on local use of its timber in tropical

Africa is lacking.Major references Aubr6ville, 1970; Bolza &

Keating, 1972; Sallenave, 1955; Takahashi,1978; Vivien & Faure, 1985.

Other references are Assi at a1. , 1985;Burki11, 1995; CIRAD Forestry Department,2008; Danforth & Noren, 1997; de Saint-Aubin,1963; 1stas, Raekelboom & Heremans, 1959;Lewis at a1. , 2005; Neuwinger, 2000; White &Abernethy, 1997; Wilczek at a1. , 1952.

Authors E. A. Obeng

CoPAIFERARELIGIOSAJ. Leonard

Protologue Bull. Jard. Bot. Etat 19: 398(1949)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Origin and geographic distribution Copui-Iero religioso is distributed from Cameroon toDR Congo

Uses The wood, traded as 'n'tone', sometimesalso as 'anzem', is suitable for light construc-tion, light flooring, joinery, interior trim, vehi-cle bodies, toys, novelties, boxes, crates, veneer,plywood, hardboard and particle board. It isalso suitable as pulp for paper production.The bark and theresin from the bark and wood

are locally used as perfume. The resin is alsoused in torches and for sealing canoes. In Ga-bon the bark of young trees is made into boxesused for ritual purposes. In traditional medi-cine in Gabon bark macerations are taken for

the treatment of cough and stomach-ache, barkdecoctions for the treatment of cardiovascular

diseases, whereas the bark is used in fumiga-tions against headache and kidney pain. Bark

.

GOPoiferu religioso - wild

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preparations are also used to treat malariaThe bark is added to a bath together with otheringredients to treat leprosy. In Cameroon andGabon the tree is credited with magical proper-ties and used for ritual purposes

Production and international trade In the

1960s Gabon and Congo exported small quanti-ties of logs. According to ATIBT statistics,Congo exported 165 ina of'n'ten6'logs in 2006.

Properties The heartwood is pale brown tored-brown, inottled, turning darker brown up-on exposure, and distinctly demarcated fromthe wide, yellow-white to pale pink sapwoodThe grain is straight to slightly interlocked,texture medium to fine, rarely uneven. Thewood contains a sticky, transparent resin, andblackish resin ducts are visible. Sawn wood has

a bitter, almond-like scent.The wood is lighter and softer than that of oth-er GOPoifero species. It has a density of about550-750 kg/ms at 12% moisture content. Thewood dries rapidly, with only a slight risk ofdistortion and checking. During drying, theresin in the wood solidifies and produces whit-ish clots if the drying temperature is above70'C. The shrinkage rates are quite low, fromgreen to oven dry 3.8% radial and 4.5% tangen-tial. Once dry, the wood is stable in service. At12% moisture content, the modulus of ruptureis about 1/7 N/min2, modulus of elasticity10,000 N/min2, compression parallelto grain 46N/min2 cleavage 21 N/mm and Chalais-Meu-don side hardness 2.3.

The wood saws well, with only slight bluntingeffects on saw teeth and cutting edges. It workswell with most tools and usually planes well,although irregular grain may sometimes causeproblems when machine tools are used. It peelswell. The gluing properties are good. Varnish-ing and painting may be difficult because of thepresence of resin. The wood is moderately du-rable, being resistant to Lyetus beetles butliable to attacks by fungi, termites and marineborers. The heartwood is extremely resistant toimpregnation with preservatives, the sapwoodmoderately resistant.The wood contains 43% cellulose, 29% 11gnin,17% pentosan and 0.8% ash. The solubility isabout 4.0% in alcohol-benzene, 3.0% in hotwater and 18.6% in a I% NaOH solution.

Bark and roots contain tannin. A dichloro-

methane bark extract has shown in-vitro an-

tiplasmodial activity against Plusmodium 101-ciporum, while the methanol extract was mac-tive. The dichloromethane bark extract also

showed considerable in-vitro cytotoxicity

Botany Large tree up to 45 in tall; holebranchless for up to 30 in, straight and cylin-drical, up to 200 cm in diameter, thickened atthe base, without buttresses; bark surfacesmooth or finely scaly, orange to reddishbrown, inner bark fibrous, brittle, pale yellow,aromatic; crown umbrella-shaped; twigs short-hairy but soon becoming glabrous. Leaves al-ternate, pinnately compound with 10-16 leaf-lets; stipules minute, early caducous; petioleand rachis 10-20 cm long, grooved above; petio-Iules 2-5 min long, twisted; leaflets alternate,oblong to elliptical or obovate, 2.5-6.5 cm x1.5-3.5 cm, base slightly asymmetrical androunded or curieate, apex rounded to notched,leathery, nearly glabrous, with translucentdots, pinnately veined with c. 20 pairs of Iat-eral veins. Inflorescence an axillary raceme upto 19 cm long, with spike-like branches 2(-6)cm long, densely hairy. Flowers bisexual, zy-gomorphic, white, nearly sessile, with 2 small,caducous bracteoles; sepals 4, ovate-lanceolate,c. 4 min x 2-3 mm, I broader than other 3,densely hairy; petals absent or sometimes 4present and then I-2 min long; stamens 8-10,free, c. 5 mm long; ovary superior, sessile,hairy at base and sides, I-celled, style c. 3 minlong. Fruit an ellipsoid to nearly globose pod 3-5 cm x 2-3.5 cm, slightly flattened, leathery,rough, glabrous to hairy, blackish, somewhatshiny, covered with small resin droplets, dehis-cent with 2 valves, I-seeded. Seeds oblong,with a red anI.

GOPoifero comprises about 40 species, of whichabout 35 occur in tropical America, 4 in Africaand I in tropical Asia. GOPoifero seems closelyrelated to Bothioeo, Detorittm, Stridoro andressmunniu

Ecology GOPoijero religiosa occurs very scat-tered in dense primary lowland rainforest. It isfound on well-drained localities and avoids

swampy or temporarily inundated sitesManagement In Gabon the minimum hole

diameter allowed for harvesting is 70 cmGenetic resources and breeding It is un-

known to what extent Copoifero religioso isthreatened with genetic erosion. In Cameroonit is considered a vulnerable species, because ithas a limited ecological range and occurs inregions where forest land is being converted inland to be used for other purposes

Prospects The wood of Copoifero religioso islighter and softer than that of other GOPoiferospecies, and only moderately durable. There-fore its range of potential uses is limited. Smallamounts of its timber are traded on the inter-

GOPAIFERA 227

228 TIMBERS2

national market, but it is not known for whichend-use. Information on the use of the timber

in tropical Africa is also lacking, but the tree islocally used as a source of resin (used for per-fume, sealing and as fuel) and various medi-cinesfrom the bark

Major references ATIBT, 1986; Aubr6ville,1970; Bolza & Keating, 1972; Lekana-Douki ata1. , 2011; Vivien & Faure, 1985.

Other references ATIBT, 2007; Costa,2009; de Saint-Aubin, 1963; Koumba Madingouat a1. , 2012; Neuwinger, 2000; Normand, 1950b;Raponda-Walker & Sinans, 1961; Sallenave,1955; Volavka, 1998; Witczek at a1. , 1952

Authors M. Brink

CoPAIFERASALIKOUNDAHeckel

Protologue min. Fac. Sci. Marseille 3: 4, t16 (1891).

Family Caesalpiniaceae (Leguminosae - Fa-baceae)

Chromosome number2n=24

Origin and geographic distribution GOPoi-Iero sankoundo is distributed from GuineaBISsau eastward to Ghana

Uses The wood, traded as 'etimoe' is usedfor producing veneer. It is locally used for inak-ing furniture. It is suitable for heavy construc-tion, heavy flooring, joinery, interior trim, mineprops, ship and boat building, vehicle bodies,sporting goods, toys, novelties, boxes, crates,agricultural Implements and turnery.The wood and bark contain an aromatic resin,which is locally used for making a scented un-gent for cosmetic use. The resin outflow is toolimited for commercial production. In Liberia

the pounded bark is rubbed on the body as aperfume. In Sierra Leone the fragrant seed ismade into a pomade, whereas in Liberia it isdried and powdered to be added to snuff forscenting it. In Guinea fragrant necklaces aremadefrombroken seeds

In traditional West African medicine, pulpedleaves are applied directly onto sores or used inhot poultices, whereas powdered dried leavesand bark are mixed with baked and powderedclay and applied to ulcers. A maceration of thefruit valves is drunk to purify the blood. A coldinfusion of the seed is used for the treatment of

vertigoProduction and international trade Ghana

exported small quantities of logs to the UnitedKingdom around 1960. The wood was probablytraded in mixed consignments and thus statis-tics are not available. In 2000 and 2001 Ghana

exported about 55 ina and 35 in3 respectively,of GOPoifero salthoundo wood

Properties The heartwood is grey with apinkish tinge when freshly cut, turning cop-pery reddish brown on exposure;it is distinctlydemarcated from the paler, up to LOG15) cmwide sapwood. The grain is interlocked orwavy, sometimes straight, texture fine to me-dium and even. When cut, the sapwood exudesa sweetly scented resin; the blackish resinducts give sawn wood a striped appearance,The wood is moderately heavy, with a densityof about 700-850 kg/ina at 12% moisture con-tent. It should be dried slowly to avoid distor-tion. The shrinkage rates are quite high, fromgreen to oven dry 5.1-5.4% radial and 8.4-9.2%tangential. Once dry, the wood is moderatelystable to unstable in service. At 12% moisture

content, the modulus of rupture is 144-190N/min2, modulus of elasticity 13,400 N/mm2compression parallel to grain 71-75 N/min2,cleavage 20 N/mm and Chalais-Meudon sidehardness 4.9

The wood works fairly easily with hand andmachine tools, but has a tendency to blunttools and to chip off. It polishes well. Pre-boring before nailing and screwing is advisableThe wood paints satisfactorily, and varnishesand glues well. The wood is durable, being re-sistant to attacks by termites and xylophagousbeetles. The heartwood is resistant to impreg-nation with preservatives, the sapwood moder-ately resistantThe wood contains 36.5% cellulose, 31% 11gnin,17% pentosan, 1.0% ash and Do silica. The sol-ubility is about 6.6% in alcohol-benzene, 3.9%in hot water and 23.9% in a I% NaOH solution

.

GOPoifero sonhoundo- wild

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The bark, wood and seeds contain coumarin.Description Deciduous, medium-sized to

large tree up to 50 in tan; hole branchless forup to 32 in, straight and cylindrical, up to 200cm in diameter, base thickened or with narrowroot ridges developing into buttresses; barksurface rough, irregularly scaly, greyish brownto dark brown, inner bark hard, fibrous, pink-ish to reddish brown, sweetly scented, exudinga clear, brown to purplish, sticky, fragrant res-in when cut; crown spreading and denselybranched; twigs pale brown hairy. Leaves al-ternate, panpinnately compound with 3-8(-9)pairs of leaflets; stipules c. 2 cm long, cadu-cous; petiole I-1.5 cm long, with joint at base,rachis 6-13 cm long, grooved above, greenishbrown hairy; petiolules very short, twisted;leaflets opposite to slightly alternate, ellipticalto oblong, 1.5-6 cm x I-3.5 cm, base asymmet-Tical, apex obtuse to notched, margins thick-ened, leathery, glabrous except at base andmargins, with translucent dots, pinnatelyveined with 10-20 pairs of lateral veins. Inno-rescence a terminal or axillary panicle 3-12 cmlong, with spike-like branches up to 3 cm long,hairy. Flowers bisexual, zygomorphic, greenish

white, fragrant, sessile, with 2 triangular, ca-ducous bracteoles c. I min long; sepals 4, ellip-tical-oblong, c. 3 mm long, I broader than other3, acute, hairy; petals usually absent; stamens10, free, unequal in length, up to 7 mm long;ovary superior, sessile, hairy at sides, I-celled,style c. 3 mm long. Fruit an ellipsoid to oblongpod 3-5 cm x 1.5-3.5 cm, slightly flattened,thick-leathery, green-red when ripe, sticky,dehiscent with 2 valves, I-seeded. Seeds c. 2.5cm x 1.5 cm, black, sweetly scented, with thin,red, waxy anI. Seedling with epigeal germina-tion; hypocoty1 (3.5-)5-7.5 cm long, epicotyl4.5-7.5 cm long; cotyledons fleshy, c. 2 cm x Icm; first 2 leaves opposite, with 4-5 pairs ofleaflets

Other botanical information GOPoiferocomprises about 40 species, of which about 35occur in tropical America, 4 in Africa and I intropical Asia. GOPoifero seems closely relatedto Bothioeo, Detorittm, Stridoro and Tessmon-

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;26: intervessel pits medium (7-10 pin); (27:intervessel pits large (z 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 I'm; (43:mean tangential diameter of vessellumina ^200 pin); 46: s 5 vessels per square minimetre;47: 5-20 vessels per square minimetre; 58:gums and other deposits in heartwood vessels.Tracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled; 70fibres very thick-walled. Axial parenchyma: 79:axial parenchyma vasicentric; 80: axial paren-chyma alitorm; 81: axial parenchyma lozenge-anform; 83: axial parenchyma confluent; 89:axial parenchyma in marginal or in seeminglymarginal bands; 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; 104: allray cells procumbent; 1/5: 4-12 rays per minSecretory elements and cambial variants: 127:axial canals in long tangentiallines. Mineralinclusions: 136: prismatic crystals present; 142:prismatic crystals in chambered axial paren-

CoPAiFERA 229

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230 TIMBERS2

chyma cells.(P. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Seedlings andsaplings tolerate shade. The growth of seed-lings is slow; they reached about 20 cm tall 4months after sowing and only 50 cm after 2years when planted in the field in fullsunIn Sierra Leone the leaves are gradually shedfrom November to April, when new leaves areproduced; fruits have been observed in Mayand more frequently in November. In Liberiaflowering trees have been observed in May andAugust, and ripe fruits have been collected inOctober and November. The tree is leafless for

a short period at the beginning of the rainyseason, before flowering. In C6te d'TVoire fruit-ing is in December-March. In Ghana floweringhas been recorded from March to May and fromSeptember to October, and ripe fruits are pre-sent in October-February. The seeds are prob-ably dispersed by birds

Ecology GOPoifero sonhoundo is most abun-dant in moist evergreen forest, but also occursin semi-deciduous forest. It prefers well-drained locations, but also occurs on periodical-Iy inundated land near rivers. It is rare inmany regions within its distribution area

Propagation and planting Natural regen-eration usually occurs abundantly near themother tree, especially in more or less exposedlocations. GOPoifero salthoundo can be propa-gated with seeds. The 1000-seed weight is c. 2kg. The seeds are collected from the ground assoon as possible after falling because they aremuch appreciated by rodents. They are sownfresh in nursery pots and shaded. Germinationtakes 5-21 days. The germination rate is about60%. The seedlings can be planted into thefield when they are 4 months old. When plant-ed in full sun, mortality is high, about 60%,suggesting that some shade is needed in earlystages of growth. Keeping the seedlings for oneyear in the nursery may give better results

Management Larger GOPuifero sonkoundotrees usually occur scattered and in low densi-ties in the forest. In Ghana it may be locallycommon; about 230 trees with a diameter lessthan 30 cm and 80 trees with a diameter over

30 cm were found per ha, indicating good natu-ral regeneration but a high mortality rate dur-Ing growth

Harvesting The prescribed minimum bolediameter for harvesting in Ghana is 70 cm.

Handling after harvest Logs of GOPoiferosankoundo do not float in water and cannot betransported by river

Genetic resources GOPotfero sonhoundo isincluded in the TUGN Red list of threatened

species as vulnerable because of over-exploita-tion and habitatloss

Prospects The prospects of GOPoifero suit-houndo as a timber tree of commercial jin-

portance are limited because of its restricteddistribution and scattered occurrence. Re-

search may be warranted on appropriate man-agement methods to reduce the mortality rateof young trees under natural regeneration.

Major references Ayensu & Bentum, 1974;Bolza & Keating, 1972; Burki11, 1995; Haw-thorne & Jongkind, 2006; Hubert, undated;Irvine, 1961; 0teng-amoako (Editor), 2006;Savill & Fox, 1967; Taylor, 1960; Voorhoeve,1979

Other references African Regional Work-shop, 1998; Catarino, Martins & Morelra, 2001;CIRAD Forestry Department, 2008; Costa,2009; de in Mensbruge, 1966; Hawthorne,1990; Hawthorne & Gyakari, 2006; Kryn &Fobes, 1959; Kunke1, 1965; Mangenot &Mangenot, 1957; Neuwinger, 2000; N'guessan,Soro & funori, 2011; Sallenave, 1955; Savard,Besson & Morize, 1954; Takahashi, 1978; Tay-10r, 1960

Sources of illustration Voorhoeve, 1979.Authors A. A. Oteng-kirioako & E. A. Obeng

CORDIAAFRICANA Lain.

Protologue Tab. encyc1. I: 420 (1792)Family BoraginaceaeSynonyms Cordio abyssinico R. Br. (1814)Vernacular names Large-leaved cordia,

East African cordia, Sudan teak (En). S6bestierd'Mrique, faux teck, teck d'Arable (Fr). Ma-kobokobo, innngamringa (Sw).

Origin and geographic distribution CordioQincono is widespread, from Guinea east toEritrea, Ethiopia and Kenya, and south to An-gola, Zimbabwe, Mozambique and northernSouth Africa. It is also found in Saudi Arabia

and Yemen, and has been planted in manytropical countries, including Reunion and Mau-ritius.

Uses The wood, in East Africa often knownas 'inukumari', is commonly used for joinery,Interior trim, panelling, furniture, cabinetwork, drums, beehives, boxes, mortars andcanoes. It is suitable for light construction, shipbuilding, vehicle bodies, toys, novelties, vats,draining boards, food containers, matches, ve-neer, plywood, hardboard, particle board and

Cordio ofricuno - wild

pulp for paper making. It is also used as fire-wood. The fruit pulp is edible and is added as asweetener to food. The leaves serve as fodderfor livestock. Several parts of the plant areused in traditional medicine. Leaf decoctionsare administered to treat headache, nose bleed-ing, dizziness and vomiting during pregnancy,wounds and worms. Fresh bark is applied tofractures and bark extracts and taken againstfatigue. Root decoctions are drunk to treatjaundice and schistosomiasis. Wood ash is ap-plied to skin diseases. Cordio of neono is plant-ed as a shade tree in coffee plantations, and asroadside tree, ornamental shade tree andboundary marker. The leaves make a goodmulch. The flowers provide nectar for honeybees

Properties The heartwood is pinkish brownto reddish brown and fairly distinctly demar-cated from the greyish, 2.5-4 cm wide sapwood.The grain is usually interlocked, texture medi-urn to coarse but even. The wood ISIustrous

The wood is moderately lightweight, with adensity of 440-580 kg/in3 at 12% moisture con-tent. It air dries well without serious degradeOnce dry, it is exceptionally stable in serviceThe rates of shrinkage are quite low, fromgreen to oven dry 3.4% radial and 5.7% tangen-tial. At 12% moisture content the modulus of

rupture is 91 N/mm2, modulus of elasticity8040 N/mm2, compression parallel to grain 47N/mm2 shear 9 N/min2, cleavage 9 N/min, Bri-nellside hardness 23.5 N/mm2 and Brinell endhardness 40 N/mm2.

The wood works well with both machine andhand tools, and it planes and moulds to a nlcesurface. It polishes wellwhen a filler is used. It

holds nails and screws well and has good glu-ing properties. The peeling and slicing charac-tenstics are satisfactory. The wood is moder-ately durable, being moderately susceptible totermite and pinhole borer attacksThe polysaccharide from the fruit pulp of Cor-di0 @1ncono consists mainly of galactose, man-nose, xylose, arabinose, glucose, rhamnose,galacturonic acid and about 2.5% protein. Itmay have applications in food, textile andpharmaceutical industries, and could act as anemulsifying agent also in systems with rela-tively high salt concentrations.

Botany Deciduous shrub or small to medi-urn-sized tree up to 25(-30) in tall; hole branch-less for up to 8(-12) in, often curved or crooked,up to 90 cm in diameter; bark surface smoothin young trees, becoming cracked or longitudi-nany fissured, pale brown to dark brown, innerbark fibrous, whitish, turning greyish to nearlyblackish upon exposure; crown rounded, dense,much-branched; twigs velvety hairy, becomingglabrous. Leaves alternate, simple; stipulesabsent; petiole I-13 cm long; blade ovate toelliptical or nearly orbicular, 7.5-20(-30) cm x3.5-18(-22.5) cm, rounded to cordate at base,rounded to acuminate at apex, margins entireto slightly toothed, leathery, rough above,short-hairy below, pinnately veined with 5-7pairs of lateral veins. Inflorescence a terminalcompact panicle up to 15 cm long, composed ofcymes, minutely hairy, many-flowered. Flowersbisexual, regular, 5-merous, nearly sessile,sweet-scented; calyx tubular, (5-)7-9 mm long,10-ribbed, slightly irregularly toothed; corollafunnel-shaped, (1.5-)2-2.5 cm long, stronglyfolded, white, with short lobes; stamens insert-ed near base of corolla tube, c. I cm long, in-cluded; ovary superior, ovoid, 2-3 mm long,glabrous, style I-2 cm long, twice bifid with 4stigmas. Fruit an ovoid to nearly globose drupe10-12 mm x 6-10 mm, glabrous, yellow, en-closed at base by the persistent calyx, withsweet pulp and slightly 4-angled stone contain-ing I-2(-4) seeds. Seeds ovoid, flattened, up to6 min x 3 mm, creamy whiteTrees have been reported to reach 7-8 in tallafter 7 years' In trial plots at 2100 in altitudein Kenya, mean annual increments of 30-years-old trees were 0.5 in in height and 0.9 cm indiameter. Trees may startflowering when theyare 3-5 years old. They usually flower duringthe dry season, but in Ethiopia flowering treeshave been recorded throughout the year. Theflowers are poUinated by insects such as bees.Fruits take 3-5 months to ripen after flower-

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CORDiA 231

232 TIMBERS2

ing. They are probably mainly dispersed bybirds.

Cordio is a large pantropical genus of about250 species, with the majority of the speciesoccurring in the New World and about 35 spe-cies indigenous in tropical Africa. It is a vana-ble genus and it has been suggested that itshould be split up in several genera.Cordio senegolensis Juss. is a shrub to smallormedium-sized tree with hole up to 40 cm indiameter occurring in savanna and drier forestfrom Senegal east to Rwanda. The yellowishwood is used in West Africa for drums and ca-noes. The bark fibre is used for wickerwork and

tying material. Leaf decoctions and macera-tions are applied as a wash or bath to treatoedema, fatigue and stiffness, and as anodyne,and they are taken to treat colic and alsoagainst pain. The sweet fruit pulp is edible.

Ecology Cordio of neono occurs in open for-est, riverme forest, edges and clearings in mon-tane forest, and wooded grassland at 500-2200(-2700) in altitude. The annual rainfall inits area of distribution is 900-2000 min, with adry period of 3-4 months and an estimatedmean annual temperature of 16-22"C. Cordioof neono prefers deep, moist but well-drainedsoils, but can also be found on rocky slopes

Management Natural regeneration of Cor-dig of neono in more closed forest is restrictedto gaps. In the nursery seedlings showed goodgrowth rates at an ITradiance level of 19% offull sun, but growth was enhanced by increas-Ing Irradiance, as well as the supply of nutri-ents at high level. The results of tests in Ethio-pia indicated that Cordio of ticono has evolveda light-quality sensing mechanism that pre-vents seed germination beneath the forest can-opy. Deep sowing as well as covering and shad-ing of seed beds in nurseries with leaf littershould be avoided

One kg contains about 1700 seeds. Germina-tion of seeds usually starts after 40-60 daysand is promoted by soaking in water for 12-48hours, and the germination rate of healthyseed is 50-80%. Seeds can be stored at 3'0 forat least I year when they have been dried inthe sun to 6-8% moisture content. Seedlingscan be transplanted from the nursery into thefield after 4-6 months. Seeds are commonlyinfested by insects; tests showed that near-infrared spectroscopy can be used in the seedcleaning process to select sound seeds.Cordio of neono is in many areas quite coin-mon although trees usually occur scattered. Itis locally retained after forest clearing and

commonly planted in agricultural fields. In aninventory in southern Ethiopia it occurred inmore than 88% of the farms, being the mostpreferred indigenous tree species, often plantedtogether with enset (Errsete Dentricosum orelw. )Cheesman). The tree can be managed by cop-picing, pollarding and pruning, Regular pol-larding is recommended when Cordio of neonois used as shade tree for crops to reduce shad-Ing

As a timber tree, Cordio girtcono can be grownin rotations of 40-50 years. Close spacing andregular pruning are recommended because theboles have a tendency to develop a poor formand numerous branches

Young trees can be attacked by nematodes.Genetic resources and breeding Locally,

especially in Ethiopia and Kenya, the exploita-tion of Cordio of neono has been severe and ledto depletion of natural stands. However, itseems unlikely that this species is under sen-ous threat because it is widespread in differenttypes of habitat, appears to have fair rates ofregeneration and is commonly planted.The genetic vanability is considerable. Severalprovenance tests have been performed. InEthiopia the differential growth characteristicswere tested using 19 provenances. The annualheight growth of 3-year-old trees varied be-tween 2 cm and 62 cm, but all provenancesproduced multi-leader stems and numerousbranches. Other tests in Ethiopia showed con-siderable genetic variation in seed inorphomet-

traits, seed germination and seedlinggrowth. The results of exposure of seedlings todrought in southern Sudan showed considera-ble intraspecific variation, with provenancesfrom Kordofan having the best response andthus most suitable as a seed source for affor-estation in drought-prone environments.

Prospects Cordt0 o17icono is an importantmultipurpose tree used mainly for timber pro-duction, as an auxiliary plant in agricultureand as medicinal plant, but also yielding ediblefruits, forage and firewood. In southern Ethio-pia it is considered one of the most importantindigenous tree species, and the demand forseed is stillincreasing. Research in selectivelylogged rainforest in south-western Ethiopiademonstrated that Cordto of neono has ade-quate regeneration and seems to have goodprospects for sustainable management of tim-her production forest. More provenance trialsare needed to select elite trees that can be used

for establishing seed orchards or establishingplantations. The main drawback as a timber

ric

tree is the often short and poorly shaped hole,which could be overcome by SIIvicultural prac-tices, I. e. close planting and regular pruning,and by selection of superior tree types

Major references Bekele-Tesemma, 2007;Janick & Paull(Editors), 2006; Jansen, 1981;Verdcourt, 1991; World Agroforestry Centre,undated.

Other references Arbonnier, 2004; Benhura& Chidewe, 2004; Bolza & Keating, 1972; Bur-kill, 1985; CAB International, 2005; Chikamaiat al. , undated; Feuillet & Bosser, 2005; Mar-tins & Brummitt, 1990; Neuwinger, 2000;Takahashi, 1978

Authors E. A. Obeng

CORDLAMiLLENii Baker

Protologue Bull. Misc. Inform. Kew 1894: 27(1894)

Family BoraginaceaeSynonyms Cordioplotythyrso Baker(1894)Vernacular names African cordia, drum tree

(En). Cordia d'Mrique, ebais (Fr).Origin and geographic distribution Cordio

milleniiis widespread, occurring from SierraLeone east to western Kenya and western Tan-zania, and south to DR Congo and northernhagola. It is commonly planted, especially inWest Africa.

Uses The wood, often traded as 'cordia wood'or 'pooli', Is commonly used for construction,joinery, interior trim, panelllrig, furniture, inu-siCalinstruments including drums, boxes, toys,utensils, toolhandles, shingles, canoes and car-ving. It is suitable for boat building, cabinetwork, veneer, plywood and hardboard. It is also

used asfirewood

In Gamero0n large pieces of bark are strippedfrom the hole to make hut-walls and partitionsIn West Africa seed powder mixed with palmoilis taken as a vermifuge and applied exter-nany to ringworm and itching skin, whereas aleaf decoction is also taken to dispel worms,and to treat asthma, cough and colds. Cordiominenitis commonly planted as a village shadetree. It has also been planted in reafforestationprogrammes. In Uganda Cordio milleniitreesare often left for shade in Eucalyptus and cof-fee plantations. The flowers provide nectar andpollen for honey bees.

Production and international trade The

wood of Cordio minenit Is only occasionallyexported and has no importance in the Interna-tionaltrade. It is mainly used locally

Properties The heartwood is pale brown toyellowish brown or medium brown, occasional-Iy pale purphsh or pinkish brown, and not dis-tinctly demarcated from the whitish or yellow-ish, 4-6 cm wide sapwood. The grain is straightor interlocked, often producing a stripe figure,texture medium to coarse

The wood is moderately lightweight, with adensity of 410-500(-600) kg/in3 at 12% ino1s-ture content. It air dries rapidly and well, butin kiln drying a high temperature schedule isrecommended. The rates of shrinkage are inod-erate, from green to oven dry 2.6-3.6% radialand 4.6-4.8% tangential. Once dry, the wood isstable in service.

At 12% moisture content, the modulus of rup-ture is 67-79 N/mm2, modulus of elasticity6860-8600 N/min2, compression parallel tograin 28-43 N/mm2 shear 6.5-9 N/min2 cleav-age 10 N/mm, Janka side hardness 2615 N andChalais-Meudon side hardness 1.3

The wood saws and works well with ordinarysaw teeth and cutting tools and with onlyslight blunting effect. Planing is satisfactory ata standard angle of 30' for back-sawn surfaces,but for quarter-cut material reduction to anangle of 20' is advisable to obtain best results,especially when the grain is interlocked. Thewood polishes well when a filler is used. Itholds nails and screws well, but is somewhatlissile. The wood has good gluing, peeling andslicing characteristics. It is moderately durableto durable with an expected service life of 8-25years. It is resistant to termite and dry-woodborer attacks and moderately resistant to fun-gal attacks. Wood dust may cause allergic reac-tions of skin and mucous membranes in wood

workers

CORDiA 233

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234 TIMBERS2

The wood contains 42% cellulose, 33% 11gnin,13% pentosan, 1.7% ash and 0.02% silica; thesolubility in alcohol-benzene is 8.2%, in hotwater 1.0% and in a I% NaOH solution 10.7%.

The terpenoid benzoquinone derivatives cordi-achromes A-F have been isolated from the

heartwood. Allantoin, ^-sitosterol, balanopho-rim, precesterol and 3-0-^-D-glucopyranosyl-^-sitosterol have been isolated from the bark,together with sphingolipids and cerebrosides

Adulterations and substitutes The wood

of Cordio minenit resembles some grades ofAfrican mahogany (Khoyo and Errtondrophrog-ino) and it has been exported as a substitute

Description Deciduous, dioecious, small tomedium-sized tree up to 30(-35) in tall; holecylindrical, often irregularly bent, branchlessfor up to LOCI5) in, up to 100(-120) cm in di-ameter, old trees often with stout but low but-tresses; bark surface smooth in young treesbecoming scaly and fissured in older trees, palebrown to greyish brown, inner bark soft, fi-brous, yellowish, rapidly becoming greenish todark brown upon exposure, with an earthysmell; crown fairly thin, flattened, with widelyspreading branches; twigs short-hairy. Leaves

alternate to nearly opposite, simple; stipulesabsent; petiole slender, 2.5-16 cm long; bladebroadly ovate-elliptical to nearly round, (3.5-)10-28(-33) cm x (2-)5-17(-26) cm, rounded tocordate at base, rounded to short-acuminate atapex, margins nearly entire to slightly andirregularly toothed, deep green above and olivegreen and hairy below, more or less 3-veinedfrom the base and with 4-8 pairs of lateralveins. Inflorescence a large, terminal, very lax,spreading panicle up to 45 cm long, consistingof condensed cymes. Flowers unisexual, regu-Iar, whitish, nearly sessile, sweet-scented; ca-Iyx tubular, 6-12 min long, finely hairy out-side, with 3-4 lobes; corolla with funnel-shapedtube 5-12 min long and 4-7 lobes 4.5-7 minlong; stamens alternating with corolla lobes,exserted; ovary superior, ovoid, glabrous, 4-celled, style twice bifid with 4 linear stigmas;male flowers with rudimentary ovary, femaleflowers with sterile stamens. Fruit an ovoid to

ellipsoid drupe up to 4.5 cm x 3 cm, yellowishto brown when ripe, enclosed at base by thepersistent and enlarged calyx, stone containingI-2(-4) seeds. Seedling with epigeal germina-tion; hypocoty1 2-4 cm long, finely hairy, epi-cotyl c. 0.5 cm long, hairy; cotyledons leafy,fan-shaped and folded, c. 2 cm x 2 cm, toothedat the apical edge, with slender petiole up to 8min long; first leaves alternate, toothed, hairy

Other botanical information Cordio is a

large pantropical genus of about 250 species,with the majority of the species occurring inthe New World and about 35 species indige-nous in tropical Africa. Cordio plotythyrsoBaker is usually kept separate from Cordiominenit differing in its often smaller, ovate andshort-hairy leaves and smaller fruits. However,there is much overlap in characters betweenthe two species and they are combined here, asproposed already by Warfa (1988). In the liter-ature, they have been much confused, and thewood has alwaysbeen mixed.

Anatomy Wood-anatomical description co\WAhardwood codes)Growth rings: I: growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; 13: SIm-PIe perforation plates; 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;25: intervessel pits small (4-7 pin); 26: inter-vessel pits medium (7-10 I'm); 30: vessel-raypits with distinct borders; similar to intervesselpits in size and shape throughout the ray cell;43: mean tangential diameter of vessellumina^ 200 pin; 46: s 5 vessels per square minimetre;56: tyloses common. Tracheids and fibres: 61

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fibres with simple to minutely bordered pits;66: non-septate fibres present; 69: fibres thin-to thick-walled. Axial parenchyma: 80: axialparenchyma allform; 82: axial parenchymawinged-anform; 83: axial parenchyma conflu-ent; 85: axial parenchyma bands more thanthree cells wide; 89: axial parenchyma in mar-ginal or in seemingly marginal bands; 90: fusi-form parenchyma cells; 91: two cells per paren-chyma strand. Rays: 98: larger rays commonly4- to 10'seriate; 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: 4-12rays per mm. Stoned structure: 122: raysand/or axial elements irregularly storied. Se-cretory elements and cambial variants: 131:intercellular canals of traumatic origin. Miner-alinclusions: 136: prismatic crystals present;137: prismatic crystals in upright and/orsquare ray cells; 138: prismatic crystals in pro-cumbent ray cells; 141: prismatic crystals innon-chambered axial parenchyma cells; 152:crystals of other shapes (mostly small); 154:more than one crystal of about the same sizeper cellor chamber; 157: crystals in tyloses(E. Ebanyenle, P. E. Gasson & E. A. Wheeler)

Growth and development Young treesusually produce a nat crown of horizontalbranches, one of which bends upwards anddevelops into a leading shoot. This results in acurved hole and because this process is repeat-ed the hole becomes irregularly bent in largertrees. Cordio minenit trees grow rapidly. InSierra Leone mean annual increments in hole

diameter of 3.3-6.3 cm have been recorded for

trees up to 18 years old. The tree is leaness fora period of up to 2 months, but seem to be ev-ergreen at some localities. In West Africa Cor-did millenii usually flowers from March toMay, after new leaves have developed. Fruitsripen towards the end of the rainy season, fromJune to August. The fruits are eaten by anI-inals including elephants, chimpanzees andbombills, which probably serve as seed dis-persers.

Ecology In West and Central Africa Cordiomilleniiis most common in semi-deciduous

forest, where it is most abundant in secondaryforest. In East Africa it is usually found inrainforest, but also in riverme forest and thick-ets in grassland, up to 1650 in altitude. It pre-fors well-drained localities.

Propagation and planting Cordio milleniiis a light-demanding pioneer species. Seedlings

are mainly found in gaps in the forest canopy,and can be very common along roadsidesThere are about 4000 seeds per kg. Germina-tion of seeds takes 10-32 days after sowing. Agermination rate of 77% has been reportedfrom Ghana, but 20-25% only from C6ted'TVoire. Hand scarification of seeds significant-Iy accelerates germination and may increasethe germination rate to nearly 100%. The seedscan be stored when they are spread on a dryconcrete floor and protected against rodentsWildlings are sometimes collected for planting

Management In general the density of Iarg-er Cordio minenit trees in the forest is quitelow, but in some areas they are common. Insouthern Cameroon the density is locally up to0.5 hole of more than 60 cm in diameter per ha,with an average total wood volume of up to 4.8mama, and in some forests in Sierra Leone upto 0.3 hole of more than 60 cm in diameter perha. In western Uganda Cordio milleniiis local-Iy abundant.The trees can be managed by coppicingGrowth of coppice from cut stumps is usuallyvigorous under favourable light conditions.Although Cordto minenit is planted in manyareas, detailed management practices are lack-Ing.

Harvesting The minimum hole diameter forharvesting Cordio minenit is 60 cm in Came-roon and 70 cm in the Central African Republicand Ghana

Yield A hole branchless for 15 in and 80 cm

in diameter may yield 4 ms of timber.Handling after harvest Freshly harvested

holes float in water and thus can be transport-ed by river. Brittle or pulpy heart may be pre-sent in large logs.

Genetic resources Although Cordio mine-nit Is locally common and its exploitation Ismoderate, it is included in the IUCN Red Listof threatened species. However, it is classifiedas a lower-risk species of least concern. Thestatus of the species should be kept under re-

CORDIA 235

view.

Prospects More research is needed on ap-propriate management systems in natural for-est to ensure sustainable exploitation of Cordiomillenti. Currently Cordio milleniiis not ex-ported on a large scale, butthe decorative woodqualities present a great potential as a coin-mercial timber, as well as its ability to regen-erate wellin disturbed forest and its fairly rap-Id growth. The often irregular shape of the holeis a drawback, which can possibly be overcomeby selective breeding and proper management

236 TIMBERS2

Cordio minenit may have prospects for use inagroforestry systemsTaxonomic studies are needed to confirm that

Cordio minenit and Cordio plotythyrso areconspecific. These should take into account thewhole distribution area

Major references Bolza & Keating, 1972;Burki11, 1985; CTRAD Forestry Department,2008; Hawthorne, 1995; Hindle, 1994; Katende,Birnie & Tengnas, 1995; Savill & Fox, 1967;Verdcourt, 1991; Vivien & Faure, 1985; Warfa,1988.

Other references African Regional Work-shop, 1998; Ako6gninou, van der Burg & vander Maesen (Editors), 2006; ATIBT, 1986; Au-br6ville, 1959; Beentje, 1994; Dalzie1, 1937; dein Mensbruge, 1966; Hausen, 1981; Keay, 1989;Mapongmetsem, Nkongmeneck & Duguma,2002; Mapongmetsem at a1. , 1999b; Neuwinger,2000; 0teng-Amoako (Editor), 2006; Sommer-Iatte & Sommerlatte, 1990; Tailfer, 1989; Ta-kahashi, 1978; Tapondjou at a1. , 2005; Taton,1971; Taylor, 1960; Wilks & Issemb6, 2000

Sources of illustration Taton, 1971; Wilks& TSSemb6, 2000

Authors R. B. Jiofack Tafokou

Ocean, tropical Asia and Australia, and islandsof the Pacific Ocean. In tropical Africa, it oc-CUTS from southern Somalia along the coasts ofKenya and Tanzania to Mozambique, and it isalso found in Comoros, Madagascar and Sey-chelles. It has been introduced in Mauritius

Uses Although no uses have been document-ed from Africa, the wood is used in tropicalAsia for light construction, beams and posts,wharves, furniture, cabinet making, musicalinstruments, tools, utensils, boxes, carvings,fancy articles, canoes, paddles and sliced ve-neer. It is suitable for food vessels and utensils

because it does not impart taste to food. It isalso used as firewood.

The seeds are edible. Leaves are sometimes

chewed together with betel nut, and they arefed to pigs. They are used to dye clothes andfishing nets and lines yellowish brown, andthey are also used in traditional medicine, e. gto treat stings of poisonous fish. In Polynesiabark, leaves, flowers and fruits are used inphytomedicine, whereas the inner bark is usedto make baskets, dance skirts, hats and fans.Cordio subcordoto is planted as a windbreakand life fence, for coastal protection and asornamental tree. The flowers are commonlyused in collars and garlands

Production and international trade The

wood is not traded in Africa, but in the Solo.mon Islands, Papua New Guinea and otherisland in the Pacific Ocean it is highly prizedand locally a significant source of Income

Properties The heartwood is pale brown todark brown, often with a purplish tinge anddark brown to nearly black streaks, and dis-tinctly demarcated from the pale yellowishbrown sapwood. The grain is usually inter-locked, texture moderately fine. The wood israther glossy. It is moderately lightweight,with a density of 470-650 kg/ms at 15% inois-ture content. However, wood from French Pol-ynesia is recorded to be heavier, with a densityof 740-850 kg/in3 at 12% moisture content. Itair dries slowly, and drying should be donecarefully to avoid distortion. The rates ofshrinkage are moderate, from green to ovendry 3.1-4.5% radial and 4.9-8.9% tangentialOnce dry, the wood is stable in serviceAt 12% moisture content, the modulus of rup.ture is 88-128 N/min2, modulus of elasticity12,300-18,100 N/min2, compression parallel tograin 45-81 N/mm2, shear 3.5 N/min2 cleavage18 N/mm and Chalais-Meudon side hardness3.1-5.9.

The wood is easy to saw and can be worked

CORDIASUBCORDATALam.

Protologue Tab. encyc1. I: 421 (1792)Family BoraginaceaeVernacular names Sea trumpet, beach

cordial, island walnut (En). Noyer d'Oceanie,faux-thene (Fr).

Origin and geographic distribution Cordiosubcordoto is extremely widespread along thecoasts of eastern Africa, islands of the Indian

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well with both hand and machine tools. It

takes a good finish with nice lustre when afiller is used. It is not durable in exposed situa-tions and attacked by powder-post and long-horn beetles, termites, as well as decay fungi,but it is durable for interior work. The heart-

wood is resistant to impregnation with pre-servatives, even when using a pressure treat-merit. The wood is reported to burn fast andcan easily be ignited by rubbing 2 pieces ofwood together

Botany Evergreen shrub or smalltree up to15(-20) in tall; hole branchless for up to 8 in,often crooked, up to 60(-100) cm in diameter,without buttresses; bark surface smooth inyoung trees, becoming fissured and scaly inolder trees, brown to dark brown, inner barkpale brown, turning greenish brown upon ex-POSure; crown rounded to irregular, withspreading branches; twigs slightly ridged,nearly glabrous, pale grey. Leaves alternate,crowded near the ends of twigs, simple andentire; stipules absent; petiole 1.5-11 cm long;blade broadly ovate to broadly elliptical ornearly round, 4-34 cm x 3-17 cm, usuallyrounded at base, rounded to acute or short-

acuminate at apex, papery but slightly fleshy,nearly glabrous, pinnately veined with c. 6pairs of lateral veins. Inflorescence an axillaryor terminalcyme up to 14 cm long, short-hairywhen young, up to 20nowered. Flowers bisex-ual, regular, heterostylous; pedicel up to 0.5(-I) cm long; calyx cylindrical to funnel-shaped,I-2 cm long, with 3-5 lobes 2-5 min long; co-rolla orange to red, sometimes white, with fun-nel-shaped tube 2-3 cm long and 5-7 spreadinglobes 1.5-2.5 cm long; stamens Inserted justbelow the middle of corolla tube, 3-8 min long;ovary superior, conical, c. 3 min long, glabrous,style 2-3 cm long, twice bind with 4 stigmasFruit an ovoid to nearly globose drupe 1.5-4 cmx 2-3 cm, glabrous, yellow to red, becomingblackish, completely enclosed by the persistentcalyx, with corky pulp and angular and ridgedstone containing I-2 seeds. Seeds elongate, 10-13 min long, whitish.Trees grow fast on moderately fertile and well-drained sites. In Indonesia seedlings reached aheight of I-1.5 in 10 months after planting, 4-5 in after 2 years and on average 7 in after 4years with an average diameter of 6.5 cm.Heavy branches often develop low on the stemand form a wide spreading crown. Cordto sub-cordoto has an extensive, shallow root system.The flowers are probably poUinated by insectssuch as bees. Trees of 3 years old may alreadyproduce fruits. The corky fruits float in waterand are dispersed by sea currents.Cordiu is a large pantropical genus of about250 species, with the majority of the speciesoccurring in the New World and about 35 spe-cies indigenous in tropical Africa. It is a varia-ble genus and it has been suggested that itshould be split up in several generaCordio cuffro Sond. occurs in southern Mozam-bique and eastern South Africa, often in coastalforest and scrub vegetation on dunes, some-times inland in woodland and forest marginsIt is also found in southern Madagascar. It is ashrub or smalltree up to 7(-20) in tall. Thepale brown and moderately heavy wood is 10-cally valued for construction, furniture andfences. It is suitable for joinery, interior trim,boat building, toys, novelties, boxes, crates,vats, veneer and plywood. It is used as fire-wood and for charcoal production. The orangefruits are edible. Cordio cuffro is an attractiveornamental tree

Ecology In many areas Cordio subcordoto isa fairly common constituent of vegetation alongthe beach and of secondary forest and thicketclose to the seashore. It is adapted to strong,

CORDiA 237

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salt-laden winds. It mainly occurs in regionswith more than 1500 min/year of rainfall andwith no real dry season, but can also be foundin regions with 1000-1200 mm of annual rain-fall and a pronounced dry season. The meanannual temperature is usually in the range of24-28'C. Cordio subcordoto is usually found ondeep sandy soils, also on sand dunes and oftenon coralline sands over limestone. It prefersneutral to slightly alkaline soils. The soil maycontain moderate levels of salt. In East Africa

Cordio subcordoto is most commonly found onsea shores above the high tide mark in sheI-tered localities

Management There are 560-700 fruits inone kg. Usually the seeds are not removed fromthe fruits because it is difficult to do so withoutdamaging them. In Malaysia seeds have beenreported to take 19-62 days to germinate, withabout 25% of the seeds being viable. Data fromIndonesia show that viahinty offresh seed maybe as high as nearly 100%, but seed collectedfrom the ground showed up to 50% germina-tion. Soaking fruits in water for one or twonights promotes germination, as well as clip-ping the ends of the fruits. Storage under am-bient conditions reduces the viahinty from 60%initially to 40% after 7 months, but dried fruitscan be stored in cool and dry conditions for upto a year with slight decrease in viabinty.It is recommended to cover the fruits by 1.5-2cm of soil in seed beds. Seedlings should bewatered sparingIy. In eastern Polynesia seedsare sown in germination boxes filled with sand,sometimes mixed with humus. The seedlingsshould be grown under well-ventilated condi-tions to avoid fungal attacks. They can beplanted into the field when 6-8 months oldwith a height of 40-50 cm. In Indonesia a spac-ing of 2.5 in x I in and 2.5 in x 3 in has beenused.

Cordio subcordoto has been classified as a pio-neer species and can grow in full sunlightHowever, trees grown under these circum-stances are hardly suited for timber produc-tion, and it is recommended to plant youngtrees in the partial shade of other trees. Theyare not very shade tolerant but can developreasonably well under light conditions above30% relative light intensity. Frequent weedingmay be necessary, as well as thinning andpruning. For timber production rotations of 60-70 years are recommended. The hole of oldertrees often develops heart rot. The larvae of themoth Ethmio wigroopicello may cause seriousdamage to trees by defoliation

Genetic resources and breeding Cordiosubcordoto Is extremely widespread and notlikely to be under threat of genetic erosionHowever, in some areas larger trees have be-come rare due to extensive exploitation. Cordiosubcordotu has been identified as a priorityspecies on several island in the Pacific Oceanfor genetic research

Prospects Cordio subcorduto is very poorlyknown in tropical Africa, but it deserves moreresearch attention because it has interestinguses in tropical Asia and the Pacific. It haspotential for the production of timber of smalldimension suitable for local handicraft manu-

facture. In many areas the wood is consideredvaluable for carving because of its moderatelyfine texture and nice figure. Cordi0 81,600rdotocan have great value for coastal protectionTests in Hawaiishowed that it is promising foruse in phytoremediation of soils contaminatedby petroleum hydrocarbon. It is a nice orna-mental tree for coastal areas with its beautiful

flowers, although it may fruit prolifically pro-ducing rounded, hard fruits which may be ahazard for pedestrians.

Major references Butaud, G6rard & GUIbal,2008; CAB International, 2005; Friday & Oka-n0, 2006; Verdcourt, 1991; Warfa, 1988.

Other references Feuillet & Bosser, 2005;mapwijk, 2003; Martins & Brummitt, 1990;Sun at a1. , 2004; Thulin, 2006; Wong & Sudo,1995.

Sources of illustration Wong & Sudo,1995.

Authors E. A. Obeng

CORDYIAAFRICANA Lour.

Protologue F1. cochinch. 2: 4/2 (1790)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Chromosome number2n=20

Vernacular names Cordyla, wild mango,bush mango, sunbird tree (En). Tondo, rondocoo). Mumbwe, ingwata, mroma, invoo, intigonzi(Sw).

Origin and geographic distribution Cordy-to of neono occurs from the coast of Kenyasouth to Zimbabwe, Mozambique, north-eastern South Africa and Swaziland

Uses The wood is used for heavy construc-tion, bridge decking, furniture, tool handles,household utensils, beehives and carvings. It issuitable for medium-heavy flooring, joinery,interior trim, mine props, ship building, vehicle

.

Cordylo of neono- wild

bodies, railway sleepers, toys and novelties. Itis not recommended for uses where it may besubject to shocks. In Malawi it is one of thefavourite woods for making canoes. The hole isoften hollowed out to make drums, which aresaid to be sonorous and can be heard at greatdistance. The wood is also used as firewoodThe fruit pulp is edible, but mainly eaten inperiods of food shortage. It has an unpleasantturpentine or bean-like smell and is said tocause vertigo. The fruits are sometimes eatenafter removal of the fruit wall and cooking.Seeds are occasionally eaten after drying over afire

Production and international trade Thewood is not traded internationally, but is coin-monly used locally.

Properties The heartwood is yellowishbrown to brown, with darker bands, and fairlydistinctly demarcated from the paler, up to 7.5cm wide sapwood. The grain is wavy or inter-locked, texture coarse. It has a nice figure dueto wavy stripes.The wood is medium-weight to heavy, with adensity of 750-910 kg/ms at 12% moisture con-tent. It air dries fairly rapidly with some ten-dency to splitting and checking but with littledistortion; backsawn boards may be subject to

Boards of 2.5 cm thick takesome cupping

about 6 weeks to air dry from green to 12%moisture content. The rates of shrinkage arefairly high, from green to 12% moisture content3.8% radial and 5.3% tangential. After drying,the wood is moderately stable in service, butsometimes quite unstable. At 12% moisturecontent, the modulus of rupture Is 67-85N/mm2 modulus of elasticity 8625-12,000

N/min2, compression parallel to grain 38-60N/mm2 shear 12 N/mm2 and Janka side hard-ness 6225-7025 N.

The green wood is easy to saw. Dried wood iseasy to saw and work with machine tools, butmore difficult with hand tools. It may give arough finish in planing operations due to thepresence of Interlocked graln. It is recommend-ed to reduce the cutting angle to 15' to avoidrough surfaces. The wood moulds, drills andmortises cleanly, but turning often gives acoarse finish. Pre-boring is needed to preventsplitting upon nailing. The wood is durable,being resistant to termite attack and moderate-Iy resistant to Lyetus, but it is liable to marineborer attack. It is resistant to impregnationwith preservatives.The fruits are astringent. The pulp representsabout 48% of the fruit weight. It has a highcontent of ascorbic acid (vitamin C), about 75ing per 100 g fresh pulp.

Description Deciduous small to medium-sized tree up to 25(-40) in tall; hole branchlessfor up to 15 in, straight or curved, up to 100(-120) cm in diameter, without buttresses; barksurface fissured, greyish brown, Inner bark

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fibrous, yellowish with reddish bands; crownrounded, with spreading branches; twigs gla-brous or minutely hairy. Leaves arranged spi-rally, imparipinnately compound with 11-30leaflets; stipules small, caducous; petiole andrachis together up to 25 cm long; petiolules upto 4 min long; leaflets usually alternate, oblongto oblong-elliptical or oblong-ovate, (I-)2-5 cmx (0.5-)I-2.5 cm, rounded to slightly notchedat apex, minutely hairy beneath, with minutepenucid dots. Inflorescence an axillary racemeup to 11 cm long, short-hairy. Flowers bisexualor male, regular; pedice1 0.5-I cm long; bypan-thium bell-shaped, c. 0.5 cm long, greenish;calyx initially entire but splitting into (3-)5reflexed lobes, greenish with yellowish hairs atapex; petals absent; stamens numerous, insert-ed at rim of hypanthium, 1.5-2 cm long, or-ange-yellow; ovary superior, ellipsoid, I-celled,on a long stipe, style short. Fruit an ellipsoid tonearly globose, indehiscent, berry-like pod 4.5-8 cm x 3-6 cm, yellow when ripe, with stipe ofc. 2 cm long, with I-3(-4) seeds embedded inyellowish, sticky pulp. Seeds oblong, c. 3 cmlong, without seed coat and endosperm.

Other botanical information Cordylocomprises 5 species and is restricted to main-land tropical Africa. The two species describedfrom Madagascar have recently been trans-forred to a separate genus Dupuyo, based onthe presence of staininodes and on differencesin seed morphology. Cordylo is also closelyrelated to Mildbroediodendron. Traditionally,Cordylo is placed in Cueso!pintoceoe (Legumi-nosoe - Goeso!pintotdeoe), but chemistry, cytol-ogy, palynology and wood anatomy support theinclusion in Pupilionoceoe (Legumtnosoe - Po-pinonoideoe), and this is also supported bymolecular studies

The name Cordy10 o17icuno has commonly beenused incorrectly for Cordy!o pinnoto (Lepr. exA. Rich. ) Milne-Redh. , which occurs in the Sa-bel region from Senegal to Garnero0n and dif-fors in its inflorescences that usually appearbefore the leaves on older shoots, more hairyflowers and more numerous, slender, whitishstamens. Literature should therefore be inter-preted with care.Cordylo densiftoro Milne-Redh. is a small de-ciduous tree up to 10 in tall, endemic to Tanza-nia, where it occurs in deciduous woodland andbushland at 800-1200 in altitude. The wood isused for poles, stools, drums, beehives and jin-PIements such as mortars and pestles. Thefruit pulp is eaten fresh or after cooking. Leafdecoctions are administered as an enema to

treat constipation, and charcoal made from thewood is applied to burns. The tree is planted asa life fence

Cordylo richordii Milne-Redh. is a small tomedium-SLzed tree up to 20 in tall, occurring inSudan and northern Uganda in deciduouswoodland on rocky hillsides. The wood is usedfor similar purposes as that of Cordylo densi-11070, whereas the fruits are eaten fresh.Cordylo somoliensis J. B. Gillett is a shrub orsmall tree up to 5 in tall, occurring in easternEthiopia and Somalia. Its wood is probablyused for similar purposes as that of Cordylodensi/1070, and the fruits are eaten.

Anatomy Wood-anatointcal 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-Iygona1; 26: intervesselpits medium (7-10 pin);(27: intervessel pits large (z 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;46: S 5 vessels per square minimetre; 58: gumsand other deposits in heartwood vessels. Tra-cheids and fibres: 61: fibres with simple to inI-nutely bordered pits; 66: non-septate fibrespresent; 70: fibres very thick-walled. Axial pa-Tenchyma: (76: axial parenchyma diffuse); 80:axial parenchyma allform; (81: axial paren-chyma lozenge-allform); (82: axial parenchymawinged-allform); 83: axial parenchyma conflu-ent; 85: axial parenchyma bands more thanthree cells wide; 91: two cells per parenchymastrand; 92: four (3-4) cells per parenchymastrand. Rays: 97: ray width I-3 cells; 104: allray cells procumbent; (106: body ray cells pro-cumbent with one row of upright and/or squaremarginal cells); 1/5: 4-12 rays per mm. Stonedstructure: (118: all rays stoned); (120: axialparenchyma and/or vessel elements stoned);122: rays and/or axial elements irregularlystoned. Mineral inclusions: 136: prismaticcrystals present; 142: prismatic crystals inchambered axial parenchyma cells; (154: morethan one crystal of about the same size per cellor chamber)re. Mugabi, P. Baas & H. Beeckman)

Growth and development The rate ofgrowth of young seedlings is high, but growthslows down later. In Tanzania trees of 14 yearsold showed a mean annual hole diameter

growth of 8.5 mm and had an average heightof

,

88 in. In southern Africa trees flower in July-October, when also new leaves develop. Theflowers are rich in nectar and pollinated bysunbirds. Fruits are ripe in November-January. They are eaten by animals such aselephants and monkeys, which serve as seeddispersers

Ecology Cordylo of neono occurs in ever-green forest and woodland, often along rivers,also in swamp forest, up to 1000 in altitude. Itis mainly found in hot areas.

Propagation and planting Fresh seedgerminates easily, but the viahinty of seed isshort. Seeds often start germinating while stiUin the fruit. Pre-treatment is unnecessary. Thetree produces suckers that can be used forpropagation.

Management Trees can be managed by pol-larding, lopping and coppicing

Genetic resources In general Cordylo Qin-curio is considered not liable to genetic erosionbecause it is widespread and locally common.In Malawiit is regarded as endangered and itis prohibited from fuelwood collection

Prospects The wood of Cordylo ofricono willremain important for local applications. Thefruits are considered valuable for human con-

sumption, but research is needed on the phyto-chemistry and nutritional value. Cordylo Girt-curio is worth of being planted as ornamentaltree; flowering trees are spectacular. Studieson growth rates and propagation are recoin-mended to give directives for sustainable ex-PIOitation and domestication of this multipur-pose tree, which has potential for use in agro-forestry systems.

Major references Bolza & Keating, 1972;Brummitt at a1. , 2007b; Chikamai at al. , undat-ed; Huxham at a1. , 1998; Maundu & Tengnas(Editors), 2005; Meyer, 2006; Palmer & Pit-man, 1972-1974; Ruffo, BITnie & Tengnas,2002; SEPASAL, 2011; Takahashi, 1978.

Other references Beentje, 1994; Brenan,1967; Bryce, 1967; Chudnoff, 1980; Coates Pal-grave, 1983; mrkbride, 2005; Mbuya at al. ,1994; Thulin, 1989; van Wyk & Gencke, 2000;Watt & Breyer-Brandwijk, 1962; Westphal,1975; Wimbush, 1957

Sources of illustration Brenan, 1967; Coa-tes Palgrave, 1983

Authors R. H. M. J. Lemmens & N. Nyunai

CORDYIAPiNNATA (Lepr. ex A. Rich. ) Milne-Redh.

Protologue Repert. Spec. Nov. Regni Veg.41: 232 (1937).

Family Caesalpiniaceae (I. eguminosae - Cae-salpinioideae)

Chromosome number2n=20

Vernacular names Gayor pear tree, bushmango (En). Poirier du Cayor (Fr).

Origin and geographic distribution Cordy-to pmnoto occurs through the dry forest andwooded savannah zones from Senegal east toNiger, northern Nigeria and northern Came-

CORDYLA 241

roon

Uses The wood is used for poles in houseconstruction, Joinery, furniture and utensilssuch as mortars. It is suitable for heavy floor-ing, interior trim, mine props, ship building,vehicle bodies, railway sleepers, sporting goodsand agricultural implements. The bole is oftenhollowed out to make drums, which are said tobe sonorous and can be heard at great distanceThe fresh fruit pulp is edible, although it isslightly astringent and said to cause vertigo.The fruits are important in times offo0d short-age near the beginning of the rainy season andare often consumed after cooking. Cooked un-ripe fruits are sometimes added to couscous ormixed with groundnut. Pounded seeds are oc-casionally added to foodDecoctions of roots, bark and leaves are addedto a bath to treat stomach-ache and orchitis

Root and bark decoctions are taken as diuretic,cholagogue, oxytocic, tonic and aphrodisiac,and to treat gastro-intestinal complaints, diar-rhoea, worms, schistosomiasis, lumbago andsyphilis. Leaf decoctions or macerations are

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Cordylopinnoto - wild

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242 TIMBERS2

taken to treat colic and anorexia, and are ap-plied as a vapour bath to treat fever. Leafpow-der is applied to abscesses and bark powder issniffed against headache and colds. Leaf decoc-tions are given to cattle as a tonic. Powderedbark and leaves are given to cattle against in-testinal worms. Bark and leaves are added to

water as purifiers. The foliage is browsed bylivestock. Cordylo pinnoto is valued as nursetree for crops

Production and international trade The

wood has been traded as 'metondo', but cur-rently it is only used locally. In Senegal in1960, a hole of Cordylo pinnoto yielded on av-erage 2 ms of sawn wood, in 1975 only I ina andto date most of the well-formed trees have dis-

appeared. The legal annual yield between 1964and 1972 comprised about 2800 trees

Properties The heartwood is yellowish brownto reddish brown and rather indistinctly de-marcated from the about 2 cm wide, paler sap-wood. The grain is usually interlocked, texture

bark surface fissured and scaly, pale brown todark brown, inner bark yellow-orange; crownrounded, dense; twigs short hairy, becomingglabrous. Leaves arranged spiralIy, imparipin-nately compound with 10-20 leaflets; stipulessmall, caducous; petiole and rachis together upto 25 cm long; petiolules up to 5 mm long;leaf-lets alternate to nearly opposite, oblong to ob-long-erriptical or oblong-ovate, 47 cm x 2-3cm, rounded to slightly notched at apex, gla-brous or minutely hairy beneath, often withminute penucid dots. Inflorescence an axillaryraceme up to 8 cm long, densely hairy. Flowersbisexual or male, regular; pedicel up to I cmlong; hypanthium bell-shaped, c. 0.5 cm long,greenish; calyx initially entire but splittinginto (3-)5 reflexed lobes, greenish, short-hairy;petals absent; stamens numerous, inserted atrim of hypanthium, 2-3 cm long, whitish; ovarysuperior, ellipsoid, I-celled, on a long stipe,style short. Fruit an ellipsoid to nearly globose,indehiscent, berry-like pod 4-8 cm x 3-6 cm,green to yellow when ripe, with stipe ofc. 2 cmlong, with 2-3 seeds embedded in yellowish,sticky pulp. Seeds oblong, c. 3 cm long, withoutseed coat and endospermTrees usually flower from the middle of the dryseason to the beginning of the rainy seasonbefore the development of new leaves. In Sene-gal it fruits in June-July, at the beginning ofthe rainy season. The growth of seedlings isvery slow; they are about 20 cm tall at the endof the first rainy season and 40 cm tall after 2years

Cordy!o comprises 5 species and is restricted tomainland tropical Africa. The two species de-scribed from Madagascar have recently beentransferred to a separate genus Dupuyo, basedon the presence of staininodes and on differ-ences in seed morphology. Cordy!o is also close-Iy related to Mildbroediodendron. Traditional-Iy, Cordylo is placed in Cuesolpinioceoe (Le-guminosoe - Coesolpinioideoe), but chemistry,cytology, palynology and wood anatomy sup-port the inclusion in Pupilionoceoe (Legumi-nosoe - Pupilionoideoe), and this is also sup-ported by molecular studies.The name Cordy!o of ticono has commonlybeen used incorrectly for Cordylo pmnoto. Thetrue Cordylo of neono Lour. is restricted toEast and southern Africa. It differs in its milo-

rescences that usually appear together withthe leaves on young shoots, less hairy flowersand less numerous, stouter, orange-yellowstamens

Ecology Cordylo pinnoto occurs in dry for-

coarse.

The wood is heavy, with a density of 840-880kg/in3 at 12% moisture content, and hard. It airdries fairly rapidly with some tendency to split-ting and checking. The rates of shrinkage aremoderately high, from green to oven dry 4.7-5.2% radial and 6.6-7.8% tangential. Afterdrying, the wood is moderately stable to unsta-ble in service. At 12% moisture content, themodulus of rupture is 110-151 N/min2 inodu-Ius of elasticity 12,740-15,600 N/min2 coin-pression parallelto grain 48-71 N/min2 shear10-12.5 N/mm2, cleavage 15-19 N/min andChalais-Meudon side hardness 10.2.

The wood is fairly easy to saw and work withmachine tools, but difficult with hand tools. Itmay give a rough finish due to the presence ofinterlocked grain; a 15' cutting angle is rec-ommended in planing. Pre-boring is needed toprevent splitting upon nailing. The wood iseasy to sand, varnish and paint. It is durable,being resistantto termite attack and moderate-Iy resistant to Lyetus, but it is moderately 11a-ble to marine borer attack. It is resistant toimpregnation with preservatives.The approximate composition of ripe fruits per100 g edible portion is: water 80 g, energy 294kJ (70 kcal), protein 1.4 g, fat 1.0 g, carbohy-drate 18 g, fibre 1.1 g, Ca 29 ing, P 142 ing, Fe1.8 ing, ^-carotene 310 pg, thiamin 002 ing,niacin 8 ing and ascorbic acid 74 ing.

Botany Deciduous smalltree up to 15(-20)in tall; hole branchless for up to 10 in, but usu-ally shorter, often straight and cylindrical;

est, woodland and savanna, in regions with anannual rainfall of 600-1500 mm. In parklandsavanna in Senegal it Is the dominant woodyspecies, with as other major tree species Loft-neo acido ARIch. , Anogeissus leiocorpo (DC. )Guill. & Perr. and Tamarindus indica L. In

this parkland, the soils are characterized bymuch sand (about 84%), little loam (8%), littleorganic matter (0.8%) and poor water holdingcapacity. Cordylo pinnoto grows better on al-Iuvial soils

Management The 1000-seed weightis about10.4 kg. The germination rate of fresh seed isclose to 100%. The seeds have a moisture con-

tent of about 50%, and the loss of viabinty israpid when the moisture content drops below30%; seeds do not germinate at all when It isbelow 24%. In storage at -5'0, seed longevitydoes not exceed 5 days. Survival is best at15'C, with still a germination rate of nearly50% after 2 months of storage, but O% after 3months. Sowing is done in plastic bags or innursery beds in the shade, which is progres-SIvely reduced, and removed 3 weeks aftergermination. Some weeks before transplantingthe roots should be trimmed. Because seedlingsdo not coppice, they are very sensltlve to fireand browsing. Although seedlings may be veryabundant at the beginning of the rainy season,regeneration is often very poor. In tests in Sen-egal, inoculation with the arbuscular mycorrhi-zalfungus Glomus erggregotum showed positiveeffects on seedling growth.In Senegal Cordylo pm, ,oto is locally the doin-inarit tree species in a parkland vegetation,comprising more than 60% of the woody plants.This is a traditional agroforestry system, whichexists for many generations. Millet andgroundnut are the major crops, and the litter ofCordylo pinnoto increases soil fertility andmodifies crop yield. In this parkland the meanheight of the Cordy!GPinnotu trees is 12 in andthe average hole diameter 48 cmFruits should be picked before they are fullyripe because they rot rapidly. Fresh or cookedfruit pulp is often dried and kept in storageDry fruits are put in water for some time andthen cooked. Dried pulp is used for cakes.Fresh fruits can be made into jam

Genetic resources and breeding In gen-eral Cordylopinnuto is considered not liable togenetic erosion because it is widespread andlocally common, but locally It is over-exploitedIn 1990 the Centre National de RecherchesForestfores (Institut Senegalais de RecherchesAgricoles)initiated a domestication programme

for the principal fruit-producing and multipur-pose forest trees of Senegal, including Cordylopinnoto.

Prospects The wood of Cordylopinnoto willremain important for local applications. It is ahighly valued tree in traditional agroforestrysystems in West Africa, which deserve pro^C-tion. It is a true multipurpose tree, not onlyyielding wood but also edible fruits and tradi-tional medicines. Cordylo pinnoto has beensuggested for planting in the Saheland Sahelo-Sondanian regions for reafforestation. It de-serves more research on SIIviculture, particu-Iarly concerning natural regeneration andgrowth, and on selection of productive fruittrees.

Major references Arbonnier, 2004; Bolza &Keating, 1972; Burki11, 1995; Huxham at al. ,1998; Takahashi, 1978

Other references B6. at a1. , 2000; Busson,1965; Danthu at a1. , 2000; mrkbride, 2005;Leung, Busson & Jardin, 1968; Neuwinger,2000; Samba, 2001; Samba, Camir6 & Margo-lis, 2001; SEPASAL, 2011; Vivien & Faure,1996

Authors N. Nyunai

CORNUS 243

CORNUSVOLKENSllHarms

Protologue Engl. , Pflanzenw. OSt-Mrikas C:301 (1895)

Family CornaceaeSynonyms Afrocronio uolhensii(Harms)

Hutch. (1942)Vernacular names Afrocrania (En). Mnyan-

dege (Sw).Origin and geographic distribution Cornus

,.

Cornus o01kensii- wild

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244 TIMBERS2

uolhensii occurs in mountain regions from DRCongo and southern Sudan east to Kenya andsouth to Zimbabwe and Mozambique.

Uses The wood, known as 'inukorombosi' inKenya, is used for utensils and handles of jin-PIements. It is suitable for light construction,light flooring, joinery, interior trim, ship build-ing, furniture, cabinet work, toys, novelties,boxes, crates, turnery, veneer and plywood. Itis also used as firewood. In Tanzania the leaves

applied as a poultice to boils. Cornuso01hensit is planted in reforestation pro-grammes and as roadside tree.

Properties The heartwood is yellowish whiteto pale reddish brown, occasionally with grey-Ish streaks, indistinctly demarcated from thenarrow sapwood. The grain is straight, texturemedium, even. The wood is medium-weight,with a density of about 610 kg/ms at 12% ino1s-ture content. It air dries without much de-

grade, but slowly. The rates of shrinkage aremoderate. It saws and works well with both

hand and machine tools. The nailing, screwingand gluing properties are satisfactory. Thewood is not durable and susceptible to attacksby blue stain fungi, termites, Lyetus borers andmarine borers

Botany Evergreen, dioecious, small to medi-urn-sized tree up to 20(-30) in tall; hole usuallystraight and cylindrical, up to 100 cm in diam-eter; bark surface rough, grey, with large lenti-cels, inner bark granular; crown with spread-ing branches; twigs initially short-hairy, be-coining glabrous. Leaves opposite, simple andentire; stipules absent; petiole up to 2 cm long,grooved above; blade elliptical to ovate or Ian-ceolate, 3.5-17.5 cm x 1.5-7 cm, curieate atbase, acuminate at apex, finely appressed-hairy when young but becoming glabrous, palegreen, pinnately veined with 3-4 pairs of prom-merit, curving and ascending lateral veinsInflorescence a terminal false umbel, up to 100-flowered, very young inflorescence enclosed byan involucre of 4 ovate to orbicular bracts, 7-8min x 6-7 min, soon falling; peduncle 2.5-5 cmlong, thickened at apex, silvery hairy. Flowersunisexual, regular, 4-merous; pedicel up to 12min long; sepals minute; petals free, 2-3 mmlong, sparsely hairy, creamy white to greenishyellow; stamens free, alternating with petals, c1.5 mm long; ovary Inferior, cylindrical, 2-celled, style short and thick, lower part sur-rounded by disk, stigma club-shaped or slightly2-lobed; male flowers with rudimentary style,female flowers with rudimentary stamensFruit an ellipsoid drupe I-1.5 cm long, minute-

are

Iy hairy, red turning purplish black when fullyripe, crowned by the remains of calyx, disk andstyle, 2-seededCornus comprises about 60 species, most ofthem occurring in temperate regions of theNorthern Hemisphere. Cornus DoIhensiiis theonly Cornus species of tropical Africa and theonly species which is dioecious and has spinypollen; it has often been considered to repre-sent a distinct genus, i. e. Afrocronio. However,phylogenetic and biogeographic reconstructionof the evolutionary history based on DNA dataas well as extant and fossil morphology showedthat Cornus uolhensiirepresents an old lineageof a group of Cornus species from Eurasia andSouth America. It was postulated that thegroup arose in Europe and distributed fromthere to Africa by the middle of the Eocene,about 45 million years ago

Ecology Cornus DoIkensii occurs in moun-tain forest at (1200-)1800-3000 in altitude,often along streams. It may dominate the can-opy layer of the forest. In Kenya it often occurstogether with East African camphor wood(Ocoteo usumborensis Engl. ) and in Ugandawith real yellowwood (Podocorpus lotifolius(Thunb. ) R. Br. ex Mirb. ). In mountains of Tan-zania, large areas of forest composed almostentirely of Cornus o01hensii and Picolhoo IOUri-folio Hiern are found, and in Zimbabwe CornusDothensit locally co-dominates with Ilex mitts(L. ) Radlk. and 01eo copensis L. It is often as-SOCiated with bamboo species

Management Cornus uolhensii may regen-erate in intensiveIy grazed areas

Genetic resources and breeding Cornus001hensiiis widespread and in several regionsabundant, and does riotseem to be threatened

Prospects Cornus DoIkensitis an interestingspecies for planting in agroforestry pro-grammes in high-altitude regions of tropicalAfrica. Research is needed on growth rates andmethods of propagation

Major references Bamps, 1971; Bolza &Keating, 1972; Cannon, 1978c; Chikamai at al. ,undated; Maundu & Tengnds (Editors), 2005.

Other references Barnps, 1973; Beentje,1994; Lovett at a1. , 2007; Sommerlatte & Som-merlatte, 1990; Verdcourt, 1958b; Xiang at al. ,2005

Authors R. H. M. J. Lemmens

CROTONMEGALOCARPUS Hutch.

Protologue 011v. , F1. trop. Mr. 6(I): 760(1912).

Family EuphorbiaceaeVernacular names Croton (En). Msenefu

(Sw).Origin and geographic distribution Croton

megolocorpus occurs from eastern DR Congoeast to Kenya and south to Malawi, Zambiaand Mozambique.

Uses The wood, most commonly known as'musine', is used for construction, flooring,stools, mortars, beehives, veneer and plywood.It is suitable for joinery, Interior trim, shipbuilding, vehicle bodies, furniture, cabinetwork, railway sleepers and agricultural imple-merits. It is also used as firewood and for char-

coalproduction.A maceration or decoction of the bark Is taken

as a vermifuge, and to treat whooping cough,pneumonia, stomach-ache, fever including ina-Iana, and abdominal complaints associatedwith gall bladder and spleen problems. Sapfrom leaves and young twigs are applied towounds. An infusion of the powdered bark withpotash is given to goats as a conditioner. Alt-bough trees are usually not browsed by live-stock, leafy twigs may serve as forage for goatsSeeds are used as poultry feed. Seed oilis test-ed as bio-fuel. The seeds can be used to dyewool yellowish. The flowers provide nectar forhoney bees; the honey produced is dark andhas a strong flavour. Oroton megolocorpus isplanted in hedges, live fences, shelterbelts andwindbreaks, and as an ornamental shade treeIt may serve as shade tree for coffee planta-tions. The foliage provides good mulch. The

fruit shells are used as mulch in vegetable gar-dens and as a component of potting mixtures.

Production and international trade Thereare no trade statistics for Croton megalocorpustimber, which is only used locally. However, Ithas been used for flooring in the United King-doin. In recent years, Croton megolocorpus hasgained interest for large-scale planting pro-grammes as a commercial poultry feed and bio-fuel crop with low agro-ecological demands,mainly in Kenya and Tanzania. Most of theplantations are still in the pilot stage of devel-opment.

Properties The heartwood is yellowishwhite to brownish grey, often with irregulardark brown streaks, and not distinctly demar-cated from the 2.5-5 cm wide sapwood. Thegrain is usually straight, texture medium.Freshly sawn wood has an unpleasant smellThe wood is medium-weight, with a density of700-750 kg/ina at 12% moisture content. It isliable to splitting and moderate distortion dur-ing air drying. Once dry, it is often not stablein service, especially when used in larger sizesAt 12% moisture content, the modulus of rup-ture is about 97 N/min2, compression parallelto grain 52 N/mm2, Janka side hardness 6000N and Janka end hardness 7560 N

The wood is easy to saw and work with handtools, but moderately difficult to machine. Itusually planes to a smooth and lustrous sur-face. It is resistant to abrasion. The nailing,screwing, gluing, varnishing, painting andjointing properties are all satisfactory. Thewood is only suitable for sliced veneer. It ismoderately durable, being slightly susceptibleto termite, dry-wood borer and marine borerattacks. It is liable to attacks by blue stainfungi. It is easy to impregnate with preserva-tives under pressure. Dry sawdust may irritatenose and throat in wood workers. It has been

reported that smoke from the wood Irritatesthe eyes.Seeds have an oil content of about 30% and a

protein content of about 50%. The oil has pur-gative activity and also showed Bpstein-Barrvirus activating potency. Bark extracts showedweak antibacterial activity in in-vitro tests.From the bark the clerodane diterpene chiro-inodine has been isolated as a major constitu-ent, together with Iupeol, betulin, ^-sitosteroland long-chain fatty esters. Ground seedsshowed good results in preliminary tests aschicken feed with no adverse effects on produc-tion and hatchability of eggs.

Adulterations and substitutes As an oil

CROTON 245

.

Croton megalocurpus - wild

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246 TIMBERS2

producer for bio-fuel, Oroton megolocorpus iscoinparable to jatropha (Jotropho cureos L. ),which has become the focus of large-scaleplanting programmes in several tropical coun-tries on account of its potential as a bio-fuelcrop with low agro-ecological demands

Description Medium-sized to fairly largetree up to 35 in tall; hole cylindrical, branchlessfor up to 20 in, up to 100(-120) cm in diameter;bark surface densely longitudinalIy fissured,rough, cracking, pale grey-brown, inner barkyellowish or pale brown; crown spreading, flat,with branches in layers; twigs densely scalyLeaves alternate, simple and entire; stipuleslinear-finform, 0.5-I cm long; petiole 2-8(-11)cm long, densely scaly; blade ovate to elliptical-ovate or oblong-lanceolate, (4-)7-14(-19) cm x(1.5-)3-7(-11) cm, curieate to rounded or shal-lowly cordate at base, with 2(-4) basal glands,shortly acuminate at apex, thickly papery tothinly leathery, scaly at both surfaces, silverybelow, pinnately veined with 15-25 pairs oflateral veins. Inflorescence a terminal pendu-IOUs raceme up to 30 cm long, densely scaly,completely male or with a few female flowers

at base. Flowers unisexual, regular, 5-meTous,yellowish; pedice1 0.5-1.5 cm long; calyx withtriangular-ovate lobes up to 4.5 mm x 3.5 mm;petals free, c. 5 mm long; male flowers withobovate petals, free disk glands and 25-30(-40)free stamens up to 8 mm long; female flowerswith linear petals, annular to shallowly 5-lobeddisk, superior, globose to shghtly 3-lobed ovaryc. 4 min in diameter, and 3 shortly 2-fid stylesFruit an ellipsoid-ovoid to globose capsule 2.5-4.5 cm long, scaly, greyish brown, with thick,woody endocarp, dehiscent from the apex, 3-seeded. Seeds ellipsoid-ovoid to oblong-ellip-soid, flattened, 2-2.5 cm long, slightly ridgedand rough, whitish to grey-brown.

Other botanical information Croton coin-

prises about 1200 species and occurs through-out the warmer parts of the world. It is bestrepresented in the Americas; about 65 speciesoccur in continental Africa and about 125 in

Madagascar.The wood of several other Croton species isused in tropical Africa, but none of them isimportant. In Ghana the wood of Croton pen-dun/107us Hutch. , a smallto medium-sized treeup to 20 in tall, is used for rafters.In DR Congo the wood of 070ton longiroce-inOSus Hutch. , a medium-sized tree up to 30 intall with hole up to 40 cm in diameter, is usedfor construction, whereas in Congo the leavesare eaten as a tonic and applied to maturatefuruncles. In Gabon the whitish wood of Oroton

wellensii De Wild. , a medium-sized tree up to25(-30) in tall with hole up to 45(-80) cm indiameter, is used for stools.In Kenya the wood of Croton diehogomus Pax,a shrub or smalltree up to 7.5 in tall, is used inbuilding huts. The roots and stems are used toflavour food and drinks. Roots and sometimesleaves are used in traditional medicine to treatcolds, fever, tuberculosis and syphilis, and as atonlc

In southern Africa the whitish wood of CFOton

megolobotrys Minl. Arg. , a small tree up to 15in tall with hole up to 60 cm in diameter, isconsidered useful for unspecified purposes. Thebark and seeds have some reputation as atreatment for malaria, whereas roots and barkare used as purgative and to treat ascites andfemale sterility. Leaves are used to treat bodypain and seeds as vermifuge

Anatomy Wood-anatointcal description CAWAhardwood codes):Growth rings: I: growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; (10: ves-sels in radial multiples of 4 or more common);

, . - ~,,.. . -.,. ~... ,. .*..... .... ..~

, ."4, -",.*, .".*

,..... J. ., . ,. " ....,.,". .,,... . , .<,.

,

*.

,

\

2

*

I.

,. ..

.

Croton megalocorpus - I, tree herbit, . 2, flower-ing twig, ' 3, seeds.Redruu)it grid odopted by AChmod Satin Nur-hornon

~,."~

."

.,.. .

3

13: simple perforation plates; 22: intervesselpits alternate; (23: shape of alternate pits po-Iygona1); 27: intervessel pits large (Z 10 pin);30: vessel-ray pits with distinct borders; simi-Iar to intervessel pits in size and shapethroughout the ray cell; 42: mean tangentialdiameter of vessellumina 100-200 prn; 47: 5-20 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;69: fibres thin- to thick-walled. Axial paren-chyma: 76: axial parenchyma diffuse; 77: axialparenchyma diffuse-in-aggregates; 78: axialparenchyma scanty paratrachea1; 86: axialparenchyma in narrow bands or lines up tothree cells wide; 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;104: all ray cells procumbent; (1/3: disjunctiveray parenchyma cell walls present); 115: 4-12rays per mm. Mineral inclusions: 136: pris-matic crystals present; 138: prismatic crystalsin procumbent ray cells; (139: prismatic crys-tals in radial alignment in procumbent raycells).re. Mugabi, P. E. Gasson & E. A. Wheeler)

Growth and development 070ton megalo-corpus grows rapidly when conditions are fa-vourable. In Kenya seedlings reached 1.7 in tallin one year, and in Rwanda 3 in in 2 years and11.5 in in 5 years. In Burundi planted treesreached on average only 3.6 in tall 7 years afterplanting, and 15 in tallwith a hole diameter of24 cm after 32 years. Trees may already startflowering when they are 4 years old. The flow-ers are short-lived. They are pollinated by in-sects such as bees. Fruits take about 5 months

to ripen after flowering. The seeds are eaten bybirds including poultry

Ecology 070ton megolocorpz, s occurs in ev-ergreen and semi-deciduous forest at (700-)900-2100(-2400) in altitude, sometimes also inriverme woodland and wooded grassland. It ismost commonly found in regions with a meanannual rainfall of 900-1900 min, with a dryseason of 3-4 months and a mean annual tern-

perature of 11-26'C. It prefers light, deep andwell-drained soils.

Propagation and planting Natural regen-eration is often prolific, Croto, I me8010corpusbeing a pioneer of large forest gaps and forestmargins. It has been reported to become inva-SIve under favourable climatic conditions. It isusually propagated by sowing, either directlyinto the field or in pots. There are 1000-1700

seeds per kg. The germination rate of freshseed is up to 95% after 45 days. Mature fruitscan be collected from the ground. Seed can beextracted after cracking the fruit wall andshould be sun dried to 5-9% moisture content.

The high oil content of the seed (about 30%)makes storage difficult, but seed can be storedin plastic containers up to I year at 3'C; agermination rate of 80% can then still bereached. Traditionally farmers prepare 8 cmdeep furrows where they drop ripe fruits 15-20cm apart. When seedlings are raised in seedbeds, they should be lightly shaded and initial-Iy watered twice a day. Croton megalocorpus issometimes propagated by wildlings and stumpplanting is also practised

Management Croton me8010corpus is local-Iy dominant in secondary natural forest and itis commonly planted. On slopes of Mount Ken-ya, 070ton megalocorpus has been found on40% of the farms at an average density of 15trees per farm. It is also retained after forestclearing as a shade tree for crops such as coffeeand sugar cane, but it may provide too muchshade for maize cultivation.

The tree can be managed by lopping, pollardingand coppicing. When planted in hedges, plantsshould be pruned for the first time after 2years' Oroton megalocorpus trees develop adeep taproot. This makes them quite droughttolerant and allows food crops to be grown un-derneath

Harvesting Fruits are most commonly col-Iected from the ground underneath the treesand seeds are separated manually. Simpletools in the form of a tin with a notch have

been developed. However, many fruits collectedfrom the ground do not contain viable seeds.Plant parts used for medicinal purposes areharvested throughout the year when the needarises. They can be used immediately or aredried and stored for later use.

Yield Preliminary observations indicate thata yield of 25-30 kg of seed per tree per year lsrealistic.

Handling after harvest The presscakeremaining after oil extraction can be incorpo-rated in high-calorie and high-protein supple-merits in chicken feed, and also as bio-fuel andas organic fertilizer. Oil extraction from theseed is done by hand-operated or mechanizedscrew presses. The o11 may be refined in a con-tinuous transesterification reactor to producebio-fuel of diesel oil quality and glycerol as avaluable by-product.

Genetic resources Croton megolocorpus is

CROTON 247

248 TIMBERS2

quite widespread, both in the wild and planted,and is not in danger of genetic erosion. Sys-ternatic germplasm collection or preservationprogrammes do riotseem to exist.

Breeding No breeding work has been un-dertaken yet. The most important selectioncriteria would be long and straight holes fortimber production and high oil yield of theseeds. Most plant material used so far is de-rived from simple selection within semi-wildpopulations or landraces. Great variance hasbeen observed in seed production, related tofrequency of flowering, number of innores-cences, number of female flowers per inno-rescence, number of seeds per fruit and seedweight. This indicates that through selectionand breeding oil yields could be considerablyincreased.

Prospects Oroton megolocorpus is a multi-purpose tree that is important for people as asource of timber, firewood and medicine, and asauxiliary plant. It is already an importantcomponent of agroforestry systems in EastAfrica. Types with large and straight bolesmight be interesting for timber plantations,but research on SIIvicultural and propagationaspects is still needed. Seeds seem to havegreat potential as a source of protein for poul-try feed and perhaps also feeds for other live-stock, but more research is still needed on thephytochemistry and possible adverse effectsbefore it can be developed into an importantexport product. However, oilproduction for bio-fuel seems to be most promising, and alreadylarge investments are being made to establishlarge-scale plantations in East Africa. Breedingwork is needed to select types with high oilproduction and optimal management methodsof plantations need to be established. Possibili-ties of vegetative propagation including tissueculture techniques have to be explored to ena-ble rapid and large-scale multiplication ofgood-quality genetic material. At the end oftheir productive life in bio-fuel plantations,approximately 50 years after planting, treescan be felled for their timber

Major references Bolza & Keating, 1972;CAB International, 2005; Chikamai at al. , un-dated; Katende, Birnie & Tengnas, 1995;Maundu & Tengnas (Editors), 2005; Mbuya ata1. , 1994; Radcliffe-Smith, 1987a; Radcliffe-Smith, 1996; Thijssen, 1996; World Agroforest-ry Centre, undated

Other references Addae Mensah at al. ,1989; Bryce, 1967; Chudnoff, 1980; GIIbert &Bellefontaine, 1973; Govaerts, Frodin & Rad-

cliffe-Smith, 2000; Hindmarsh, 1982; Hines &ECkman, 1993; Ichikawa, 1987; Johns at al. ,1994; Kokwar0, 1993; Leonard, 1962; Matu &van Staden, 2003; Neuwinger, 2000; Nioroge &Bussman, 2007; Noad & Birnie, 1989;Takahashi, 1978; Tee1, 1984; Wimbush, 1957;Yanase & It0, 1984.

Sources of illustration Noad & Birnie,'989; Tee}, 1984

AuthorsA. Maroyi

CROTONMONGUE Baill.

Protologue Adansonia I: 158 (1861).Family EuphorbiaceaeOrigin and geographic distribution Oroton

mongue is endemic to northern and easternMadagascar

Uses The wood, locally known as 'ino1anga'is occasionally used, mainly for laths of tiledroofs, support poles for crops, fence posts, boxesand matches, and as firewood. It is suitable forjoinery, moulding, plywood and particle boardminfusion of the bark is administered as a

tonic in cases of anorexia, fatigue and depres-SIon. Bark and leaf decoctions are applied inbathes and lotions to treat blennorrhoea

Properties The heartwood is whitish andnot distinctly demarcated from the sapwood.The grain is straight, texture medium. Woodsurfaces are slightly lustrous. The wood islightweight, with a density of 400-560 kg/ing at12% moisture content. It air dries rapidly, withmoderate risks of distortion, checking and col-lapse. Boards of 4 cm thick take 5-6 months toair dry. Once dry, the wood is only moderatelystable in service. The rates of shrinkage during

I.

Oroton mongue- wild

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drying are low.At 12% moisture content, the modulus of rup-ture is 98-139 N/min2, modulus of elasticity9000-11,000 N/mm2, compression parallel tograin 38-48 N/mm2 and Chalais-Meudon sidehardness 1.5-2.8

The wood is easy to saw and work with handand machine tools. It usually planes and sandsto a smooth and lustrous surface. The screw-

ing, gluing, varnishing, painting and jointingproperties are all satisfactory. However, thenail-holding capacity is only moderate. Thewood is suitable for rotary-cut veneer. It is notdurable, being liable to attacks by fungi andinsects. It is easy to impregnate with preserva-tives

A toxic protein, monguine, was extracted fromthe seeds; it inhibits protein synthesis

Botany Evergreen shrub to small or medi-urn-sized tree up to 20(-30) in tall; hole up to50(-60) cm in diameter; bark surface smooth toslightly rough, whitish, with lenticels, innerbark brown to greenish brown; crown rounded;twigs densely short-hairy. Leaves alternate,simple; stipules caducous; petiole up to 15 cmlong, hairy; blade ovate, up to 18 cm x 16 cm,cordate at base, with 2 basal glands, shortlyacuminate at apex, margins finely toothed,papery, stellate-hairy above, woolly silveryhairy below, (3-)5-7-veined from the base andwith few pairs of lateral veins. Inflorescence aterminal or axillary raceme up to 35 cm long,laxly flowered, completely male or with a fewfemale flowers at base. Flowers unisexual, reg-ular, 5-merous, yellowish to creamy white, withvalvate sepals, jinbricate petals and free diskglands; male flowers with c. 16 free stamens;female flowers with superior, globose to slight-Iy 3-10bed ovary, and 3 shortly 2-fid styles.Fruit a globose capsule I-1.5 cm in diameter,pale green, greyish to brownish hairy, dehis-cent, 3-seeded. Seeds ovoid, c. I cm long, slight-Iy ridged and rough, whitish to grey-brownSeedling with epigealgerminationSeedlings reached an average height of 28 cmafter I year. Croton mongue is said to be fastgrowing. It can be found flowering from July toSeptember and fruits are ripe from January.Croton comprises about 1200 species and oc-curs throughoutthe warmer parts of the worldIt is best represented in the Americas; about 65species occur in continental Africa and about125 in MadagascarOccasionally the wood of some other Crotonspecies is used in Madagascar. The wood of070ton cotoneoster Milll. Arg. (synonym: Croton

inghofoliensis Leandri) is used for parts of tra-ditional canoes, and that of Croton inuricotusVahl for cross-lathes in light constructionwork

Ecology Croton mongue occurs in forest,often in secondary or degraded forest alongrivers, up to 1300 in altitude. It is found inregions with 2000-3000 min annual rainfallwith up to 3 dry months, and an average annu-altemperature of about 17'C. It occurs on clay-ey and latentic soils

Management Oroton mongue is a pioneerspecies. Seeds may germinate abundantlywhen they are exposed to the sun. Immersionin water for 48 hours improves germination.The germination rate is about 65%, and germi-nation starts after 3-7 weeks.

Genetic resources and breeding Crotonmongue is widespread in Madagascar and oc-curs in disturbed habitats. Therefore, there isno reason to consider it threatened by genetic

CRUDiA 249

erosion.

Prospects Croton mongue has been proposedas suitable for large-scale production of match-sticks, but the usually smallbole diameter is aserious drawback. It has good prospects forreforestation of degraded land

Major references Blaser at a1. , 1993; Bot-teau, Bolteau & Allorge-Bolteau, 1999; Gu6-neau, Bedel & Thie1, 1970-1975; Parant, Chi-chignoud & Rakotova0, 1985; Rakotovao at al. ,en preparation

Other references Ranson at a1. , 1986; Ra-zakanirina, 1980; Schatz, 2001

Authors R. H. M. J. Lemmens & D. Louppe

CRUDIASENEGALENSIS Planch. ex Benth.

Protologue Trans. Linn. SOC. 25: 314 (1865)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Origin and geographic distribution Crudio

senegalerrsis is widespread in West Africa, fromGuinea-Bissau eastto south-western Nigeria.

Uses The very heavy and hard wood is rare-Iy used, but it has been used for boards in SIer-ra Leone. The tree is valued as an ornamentalshade tree.

Properties The wood is greyish brown, oftendark brown in the centre of the log. It has amoderately fine texture and it is very hard,making it very difficult to saw and work

Botany Smallto medium-sized tree up to 30in tall; hole often low-branching and crooked,up to 90(-100) cm in diameter, with narrow

250 TIMBERS2

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Crudio senegolensis - wild

buttresses; bark surface irregularly scaly, grey-ish brown, inner bark fibrous, reddish brown;crown spreading; twigs glabrous. Leaves alter-nate, imparipinnately compound with 6-12leaflets; stipules narrowly ovate, united atbase, I-3(-4) cm long, persistent; petiole I-2(-2.5) cm long, rachis up to 15(-19) cm long; peti-o1ules I-3 mm long, twisted; leaflets alternate,ovate-elliptical to obovate, (1.5-)5-IOC15) cmx (I-)2-5(-7) cm, slightly asymmet"ical, gla-brous. Inflorescence an axillary or terminalraceme up to 21 cm long, glabrous, looselyflowered. Flowers bisexual, regular, greenish;pedice1 1.5-2.5 cm long; sepals usually 4, ellip-tica1, 5-7 min long, glabrous outside, short-hairy inside; petals absent; stamens 10, free,I-1.5 cm long; ovary superior, c. 0.5 cm long,hairy, style c. I cm long. Fruit an elliptical tonearly round, flattened pod 5-12 cm x 4-7 cm,hairy, dehiscing by 2 woody valves, I-2-seededSeeds elliptical to nearly round, flattened, 4-6cm long, glabrous, brown. Seedling with epige-al germination, but seemingly hypogeal; hypo-cotyl c. 0.5 cm long, epicoty1 14-20 cm long;cotyledons remaining within the seed coat; firstleaves alternate, with 3(-4) pairs of oppositeleaflets.

There is no information on growth rates ofCrudio senegolensis, but probably trees growslowly. In Gabon young trees of Orudio gobo-itensis Pierre ex De Wild. reached a mean an-

nualbole diameter growth of 8 min, older treesI-3 mm. In C6te d'TVoire trees of Crudio serve-golensis flower in October to February andfruits mature in January to March(-Maxi.Crudio comprises about 55 species, of whichabout 30 in tropical Asia, 10 in tropical Amen-

ca and 10 in West and Central Africa. Its affin-

ity is still uncertain. Several other Orudio spp.become medium-sized and sometimes even

large trees, and their wood may be used occa-sionally, although it is extremely hard anddifficult to cut. One of these is CrL, din gobonen-sis, which may become 50 in tall with a hole upto 125 cm in diameter; it is found from C6ted'TVoire to Gabon and Congo.

Ecology Crudio senegalerrsis occurs in low-land rainforest, often in swamp forest andalong rivers and lagoons, up to 50 in altitude

Management There are about 100 seeds perkg. Germination starts 2-3 months after sow-ing and the germination rate is low. Pre-treatment of the seeds is needed to obtain fair

germination results.Genetic resources and breeding Although

CFudio senegolensis is widespread in WestAfrica, it occurs usually scattered, most coin-monly in riverme and swamp forest. Therefore,it may be liable to genetic erosion when theseforest types decline.

Prospects The timber of Crudio senego-lensis and other Crudio spp. will not becomecommercially interesting because of scatteredoccurrence and low growth rates of the trees,and the difficulties in sawing and working thewood

Major references Berhaut, 1975b; Breteler& Nguema Miyon0, 2008; Burki11, 1995; deKoning, 1983; Savill& Fox, 1967

Other references Aubr6ville, 1959b; de inMensbruge, 1966; Hawthorne & Jongkind,2006; Keay, 1989; Lewis at a1. , 2005; Normand,1950a; Normand & Paquis, 1976; Vivien &Faure, 1985.

Authors R. H. M. J. Lemmens

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CRYPTOSEPALUMSTAUDTllHarms

Protologue Bot. Jabrb. Syst. 26: 267 (1899).Family Caesalpiniaceae (I, eguminosae - Cae-

salpinioideae)Origin and geographic distribution Crypto-

sepo!urn stoudtii occurs from Cameroon southto eastern Gabon

Uses The wood, known as 'tani' or 'ekop tani'is suitable for heavy construction, heavy floor-Ing, Joinery, interior trim, ship building, vehi-cle bodies, furniture, railway sleepers, ladders,sporting goods and toolhandles.

Production and international trade The

wood of Cryptosepolum stot, dtiiis mainly usedlocally and rarely traded on the international

I.

Cryptosepo!urn stoudtz, - wild

market

Properties The heartwood is pale brown toorange-brown and usually not distinctly de-marcated from the up to 15 cm wide, greyish toyellowish white sapwood. The grain is wavy tointerlocked, texture moderately coarse. Thewood is sometimes slightly glossy.The wood is heavy, with a density of 890-950kg/ing at 12% moisture content, hard andtough. Drying often causes distortion andshould be done with care. The rates of shrink-

age are high, from green to oven dry 5.5-5.8%radial and 10.4-11.5% tangential. It is recoin-mended to quarter-saw logs before drying. At12% moisture content, the modulus of ruptureis 189-210 N/min2 modulus of elasticity15,090-16,950 N/min2, compression parallel tograin 73-83 N/mm2, shear 10-13.5 N/mm2,cleavage 22-24 N/min and Chalais-Meudonside hardness 6.5-10.2.

The wood is difficult to saw and work with bothhand and machine tools; considerable power isrequired and the wood has rather seriousblunting effect on saw teeth and cutting edges.The interlocked grain may cause problems inobtaining a smooth finish. Pre-boring is neededfor nailing. The wood spitts easily. It takesvarnish and paint well. It is moderately dura-ble, being susceptible to marine borers andoccasionally to termites. The sapwood is sus-ceptible to Lyetus attack. The heartwood isvery resistant to impregnation with preserva-tives and the sapwood moderately resistant

Description Deciduous, small to medium-sized tree up to 25(-35) in tall; hole branchlessfor up to 15(-20) in, usually straight and cylin-drical, up to 100(-120) cm in diameter, usually

without buttresses; bark surface smooth, pink-ish grey, inner bark fibrous, reddish pink;twigs glabrous. Leaves alternate, panpinnatelycompound with 10-13 pairs of leaflets; stipuleslinear, fused at base, caducous; rachis finelyhairy; leaflets opposite, sessile, oblong, I-1.5cm x 3-4 mm, usually truncate at base, round-ed or slightly notched at apex, glabrous, oftenwith a translucent glandular dot near base,pinnately veined. Inflorescence a short termi-nal raceme, usually on lateral twig. Flowersbisexual, zygomorphic, whitish, with 2 concavebracteoles 5-6 mm long, enclosing bud, persis-tent; pedicel up to 8 mm long; calyx with c. Imm long tube and usually 4 smallteeth; petalI, elliptical, up to 6 mm long; stamens 3, c. 6mm long; ovary superior, ellipsoid, with stipe,nearly glabrous. Fruit an obovate to oblong,flattened pod 6-8 cm x 3 cm, pointed at apex,smooth with thickened edges, greyish brown,dehiscing with 2 woody valves, I-2-seededSeeds rounded, flattened, brown

Other botanical information Cryptosepo-Ium comprises about 11 species and is restrict-ed to tropical Africa. It seems most closely re-lated to Poromocrolobium.

Cryptosepolum tetrophyllum (Hook. f. ) Benth. isa small to medium-sized tree, occasionally up

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CRYPTOSEPAl, un 251

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Cryptosepolum stoudtii - I, flowering trotg, 2,flower, . 3, fruit.Redrown and udopted bylshoh Syomsudin

3

252 TIMBERS2

to 30 in tallwith straight, cylindrical hole up to80 cm in diameter. It is characterized by 4 leaf-lets per leaf, the basal ones smaller. It occursin West Africa, from Guinea to Ghana. Itswood, described as whitish to pinkish and ofrather poor quality, Is sometimes used in con-struction although difficult to nail.

Anatomy Wood-anatomical description (IAWAhardwood codes):Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: slinPIe perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 26: intervessel pits medium (7-10 F1m);29: vestured pits; 30: vessel-ray pits with dis-tinct borders; similar to intervessel pits in sizeand shape throughout the ray cell; 42: meantangential diameter of vessellumina 100-200pin; 46: S 5 vessels per square minimetre; 47:5-20 vessels per square minimetre; 58: gumsand other deposits in heartwood vessels. Tra-cheids and fibres: 61: fibres with simple to inI-nutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled; 70:fibres very thick-walled. Axial parenchyma: 80:axial parenchyma allform; (81: axial paren-chyma lozenge-anform); (82: axial parenchymawinged-allform); 83: axial parenchyma conflu-ent; 89: axial parenchyma in marginal or inseemingly marginal bands; 91: two cells perparenchyma strand; 92: four (3-4) cells perparenchyma strand; 93: eight (5-8) cells perparenchyma strand. Rays: 97: ray width I-3cells; 104: all ray cells procumbent; (106: bodyray cells procumbent with one row of uprightand/or square marginal cells); 1/5: 4-12 raysper mm. Secretory elements and cainbial van-ants: 131: intercellular canals of traumatic

origin. Mineral inclusions: 136: prismatic crys-tals present; 142: prismatic crystals in chain-hered axial parenchyma ceUs; (143: prismaticcrystals in fibres).(P. Mugabi, P. Baas & H. Beeckman)

Growth and development In Gabon Cryp-tosepolum stoudtii usually flowers in Septem-her-October when also young leaves develop;the trees with their white flowers mixed with

pinkish young leaves are then very attractive.The young leaves are eaten by monkeys, chim-panzees, gorillas and elephants. Fruits ripen inJanuary-February; they open explosiveIy witha sharp crack, dispersing the seeds over shortdistances

Ecology Cryptosepolum stoudtii occurs inlowland evergreen rainforest, often along riv-

Genetic resources Cryptosepo!urn stoudtiihas a limited distribution area and it seems

fairly uncommon in most parts of its range. Atpresent, there are no indications of threats bygenetic erosion, but it might easily becomethreatened when logging becomes more inten-SIve.

Prospects Despite its good timber proper-ties, there is too little information on manyaspects of Cryptosepolum stoudtiito assess itsprospects as a commercial timber source. Fur-ther research on its growth and proper inari-agement in natural forest is warranted

Major references Aubr6ville, 1970; Bolza &Keating, 1972; BurkiU, 1995; Keay, Hoyle &Duvigneaud, 1958; Takahashi, 1978; Vivien &Faure, 1985; White & Ahemethy, 1997.

Other references Aubr6ville, 1968; Haw-thorne & Jongkind, 2006; Lewis at a1. , 2005;Normand & Paquis, 1976; Sallenave, 1955;Sallenave, 1964; Voorhoeve, 1979.

Sources of illustration Aubr6ville, 1968Authors JR. Cobbinah & E. A. Obeng

CTENOLOPHONENGLERIANUS Mildbr.

Protologue Notizbl. Bot. Gart. Bermi-Dahlema 706 (1924)

Family CtenolophonaceaeOrigin and geographic distribution Cteno-

lophon errglerionus occurs from southern Nige-ria to Gabon and northern ingola.

Uses The wood, known as 'okip' in Gabon, isused for heavy construction in house building,also for poles, and for canoes. The buttresseshave been used for rifle-butts. The wood is

suitable for heavy flooring, mine props, vehicle

ers.

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Ctenolophon errglerionus - wild

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bodies, railway sleepers, sporting goods andagricultural implements. It is used as firewoodThe bark is used for walls of huts.

Production and international trade Thewood of Ctenolophon errglerionus is used locallyand rarely traded on the international market

Properties The heartwood is reddish brown,sometimes with purplish streaks, and iridis-tinctly demarcated from the sapwood. Thegrain is usually straight, texture flne. Thewood contains oildeposits and tylosesThe wood is heavy, with a density of 920-970kg/ina at 12% moisture content, hard andtough. The tendency of warping, checking andsplitting during drying is high and it is recoin-mended to dry the wood carefully and slowlyThe shrinkage rates are quite high, from greento oven dry 5.9-6.5% radial and 9.0-9.3% tan-gential. Once dry, the wood is moderately sta-ble in service. At 12% moisture content, themodulus of rupture is 182-240 N/min2, inodu-Ius of elasticity 14,600-16,850 N/min2, coin-pression parallel to grain 71-96 N/min2, shear9.5-10.5 N/min2, cleavage 14-27 N/min andChalais-Meudon side hardness 7.5-18.2The wood is rather difficult to saw and workwith both hand and machine tools. The blunt-

ing effect on saw teeth and cutting edges isquite severe. Mortising, moulding and planingoperations are difficult, although the resultsare usually satisfactory. Pre-boring is neededfor nailing and screwing. The wood is durable,being resistant to termite and Lyetus attacks,but liable to marine borers. It is resistant totreatment with preservatives

Description Small to medium-sized tree upto 25(-30) in tall; hole usually straight, up to80 cm in diameter, fluted at base and withsteep buttresses; bark surface smooth, brown,becoming scaly with irregular, small, peelingscales, inner bark pinkish; crown rounded,irregular; twigs with stellate hairs. Leavesopposite, simple and entire; stipules Interpetio-Iar, caducous; petiole 3-6 min long, slightlygrooved; blade ovate to elliptical, 3.5-8.5 cm x2-5 cm, curieate at base, short-acuminate atapex, leathery, glabrous, pinnately veined with6-8 pairs of lateral veins. Inflorescence a ter-minal panicle, short-hairy with stellate hairsFlowers bisexual, regular, 5-merous, sweet-scented; pedicel c. 2 mm long; sepals free,rounded, 2-4 min in diameter, minutely hairy;petals free, oblong, I-1.5 cm long, minutelyhairy outside, greenish yellow to pinkish; sta-mens 10, free, I-1.5 cm long, whitish; ovarysuperior, elongated, densely hairy, 2-celled,

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CTENOLOPHON 253

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Cteno!ophon errglerionus - I, buse of 601e, 2,flowering twig, ' 3, flowerRedrou!it grid odopted by Ishoh Syomsudin

style long, 2-10bed. Fruit a cylindrical capsule1.5-2 cm long, longitudinalIy striped, glossybrown, dehiscing with 2 valves, I'Seeded. Seedup to 1.5 cm long, with comb-like, fibrous anI

Other botanical information Ctenolophonis a small genus comprising 2 widely separatedspecies, one occurring in tropical Africa and theother in tropical Asia. Formerly, it has beenplaced in the family Linoceoe.

Anatomy Wood-anatonitcal description (IAWAhardwood codes):Growth rings: 2: growth ring boundaries lridis-tinct or absent. Vessels: 5: wood diffuse-porous;9: vessels exclusively solitary (90% or more);14: scalariform perforation plates; 16: scalari-form perforation plates with 10-20 bars; 17scalariform perforation plates with 20-40 bars;(21: intervessel pits opposite); 22: intervesselpits alternate; 25: intervessel pits small(4-7Urn); 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; 41: mean tangentialdiameter of vessellumina 50-100 grn; 48: 20-40 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheids

254 TIMBERS2

and fibres: 61: fibres with simple to minutelybordered pits; 62: fibres with distinctly bor-dered pits; (63: fibre pits common in both radi-al and tangential walls); 66: non-septate fibrespresent; 70: fibres very thick-walled. Axial pa-renchyma: 76: axial parenchyma diffuse; 77:axial parenchyma diffuse-in-aggregates; 78:axial parenchyma scanty paratrachea1; 92: four(3-4) cells per parenchyma strand; 93: eight(5-8) cells per parenchyma strand. Rays: 97ray width I-3 cells; (98: larger rays commonly4- to 10-senate); (102: ray height > I mm); 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;1/3: disjunctive ray parenchyma cell wallspresent; 1/5: 4-12 rays per min. Mineral inclu-

136: prismatic crystals present; 137:SIOnS

prismatic crystals in upright and/or square raycells; 138: prismatic crystals in procumbent raycells; (140: prismatic crystals in chamberedupright and/or square ray cells); 142: prismaticcrystals in chambered axial parenchyma cells.(E. E. Mwakalukwa, P. Baas & H. Beeckman)

Growth and development In Nigeria Cte-ito10phon errglerionus flowers from March toApril and from August to September, in Gabonfrom March to August. Fruits ripen soon afterflowering, and flowers and ripe fruits cansometimes be found on the same tree

Ecology Ctenolophon errglerionus occurs infreshwater swamp forest and riverme forest atlow altitudes.

Management Cteno!ophon errglerionus islocally gregarious. In some forests in westernGabon an average wood volume of 1.1 mayhahas been recorded

Genetic resources Ctenolophon errglerionushas a limited area of distribution, but it is 10-cally common in suitable habitats. It is notcommercially exploited and it does not seem tobe liable to genetic erosion at present.

Prospects Cteno!ophon errglerionus is a use-ful local source of wood, particularly valuedbecause of its durability. It does not seem tohave potential as a commercial timber sourcebecause it occurs only locally abundantly andoften has a poor hole shape with deep grooves.

Major references Badr6, 1973a; Bolza &Keating, 1972; Burki11, 1985; de Saint-Aubin,1963; Keay, 1989; Raponda-Walker & Sillans,1961; Tailfer, 1989; Takahashi, 1978.

Other references Badr6, 1972a; Keay, 1958b;Normand & Paquis, 1976; Sallenave, 1955;Sallenave, 1964

Sources of illustration de Saint-Aubin,1963; Keay, 198, .

Authors E. A. Obeng

CURTISiADENTATA (Burm. f. ) C. Asm

Protologue Journ. S. atr. For. Ass. 20: 50(1951)

Family Cornaceae (APG: Curtisiaceae)Chromosome number it= 13

Synonyms CurtisiofogineoAiton (1789).Vernacular names Assegaitree (En).Origin and geographic distribution Curti-

sio dentoto occurs in eastern Zimbabwe, south-ern Mozambique, eastern and southern SouthAfrica, and Swaziland.

Uses The wood has been used for furniture,heavy flooring, wagons, spokes and tool han-dles. It is suitable for interior trim, ship build-Ing, sporting goods, toys, novelties, agriculturalimplements, musical instruments, carving,pattern making and turnery.Bark extracts are used in traditional medicine,especially by Zulu people, to treat diarrhoeaand stomach complaints, to purify the blood,and as aphrodisiac. The bark is also used totreat pimples and in skin lighteners. It hasbeen used for tanning leather. Curtisio dentotois planted as ornamental in gardens and inhedges.

Production and international trade In 1998the annual trade of Curtisio dentoto bark in

KwaZulu-Natal Province (South Africa) wasestimated at 23.9 t.

Properties The heartwood is greyish brownto purplish red and not distinctly demarcatedfrom the slightly paler, fairly wide sapwood

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Curtisio dentoto - coild

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The grain is straight, texture fine and evenThe wood is heavy, with a density of 870-930kg/ina at 12% moisture content. It is difficult todry, often developing deep cracks on the sur-face. Very mild schedules are needed in kilndrying. The rates of shrinkage are rather high.Once dry, the wood is stable in serviceAt 12% moisture content, the modulus of rup-ture is 1/6-131 N/mm2, modulus of elasticity16,660-18,720 N/min2, compression parallel tograin 72 N/min2, shear 13.5-15 N/min2, Jankaside hardness 11,290-11,550 N and Janka endhardness 12,620-13,110 N. Although the woodis heavy, hard and tough, It saws and worksfairly well. It can be planed well and producesa smooth and glossy surface. Pre-boring is nec-essary in nailing and screwing. The wood gluesand turns well. It is fairly durable, also whenused in contact with water and soil.

The bark contains up to 21% of tannin, whichis probably responsible for the medicinal prop-erties against diarrhoea. Lupeol, betulinic acid,ursolic acid and 20-hydroxyursolic acid havebeen isolated from the leaves; the three lattercompounds showed pronounced antifungal ac-tivity, especially against Condid0 o16iconsLeaf extracts also showed considerable anti-bacterial activity against a wide panel of bacte-

Botany Evergreen shrub or small to medi-urn-sized tree up to 20 in tall; hole usually cy-11ndrical, up to 180 cm in diameter, withoutbuttresses; bark surface smooth and brown inyoung trees, becoming fissured and dark brownin older trees; twigs densely reddish brownhairy when young, later becoming more or lessglabrous. Leaves opposite, simple; stipulesabsent; petiole up to 2.5 cm long; blade ovate toelliptical or nearly circular, 2.5-10 cm x 2.5-5.5(-7.5) cm, curieate to rounded at base, acuteto rounded at apex, margins distinctly toothedespecially in upper part of leaf, densely reddishbrown hairy below when young, but becomingglabrous, pinnately veined with 6-10 pairs ofprominent lateral veins. Inflorescence a termi-nal panicle up to 12 cm long, densely hairy,many-flowered. Flowers bisexual, regular, 4-merous, sessile; calyx fused with the ovary,with broadly triangular teeth, hairy; petalsfree, ovate, I-1.5 min long, hairy outside; sta-mens free, alternating with the petals, c. I mmlong; ovary inferior, top-shaped, hairy, 4-celled,style short, stigma 4-lobed. Fruit an ovoid toglobose drupe 5-10 min in diameter, whitebecoming red when fully mature, crowned withthe remains of calyx and style, 4-seeded

Trees grow comparatively slow under naturalconditions, but may be fast-growing in cultiva-tion. They flower from October to MarchCurtisio dentoto is almost entirely self-pollinating. Fruits ripen 6-10 months afterflowering, but they often develop sporadicaUyThey remain on the tree for a long time and areeaten by birds, monkeys and wild pigs, whichmay act as seed dispersers; the IOUrie and thebush dove have been recorded as the main seed

dispersers.Curtisio comprises a single species. It has longbeen included in Comaceoe, but recently it hasbeen classified in its own family Curtisioceoe,mainly because of differences in anatomy of theovary. Phylogeny reconstruction based on ino-Iecular data grouped Curtisio close to Grubbio,a genus of 3 shrub species endemic to southernSouth Africa, and it has been suggested toplace both genera in Grubbioceoe, also becauseof inorphological similarities; however, thefruits are completely different

Ecology Curtisio dentoto occurs in ever-green forest, grassland on mountain slopes andcoastal scrub forest, from sea-level up to 1800in altitude. Trees develop best in medium-moist forest. Curtisto dentoto seems to be alater successional species in the forest insouthern South Africa, where it is often associ-ated with 01eo copensts L. , Podocorpus lotifo-Itus (Thunb. ) R. Br. ex Mirb. , AfrocorpL, s foieo-tus (Thunb. ) GNPage and Ocoteo bulloto(Burch. ) Baill.

Management The germination rate of freshseed is about 50%. Under natural conditionsseeds take 6-12 months to germinate after thefruits have fallen. When the fleshy fruit pulp isremoved, germination may start after 3-4weeks. For good growth, seedlings should beplanted in deep, fertile soil with plenty of coin-post, lightly covered with soil, and under lightshade. They are susceptible to drought; regularwatering is needed until the young trees areabout 3 years old. Once established, the treecan withstand moderate drought. Seedlingsand saplings are sensitive to frost.Curtisio dentoto occurs scattered in the forestand is never dominant or even co-dominant

Trees respond to harvesting by producing vig-orous coppice shoots, both from wounds in thebark as well as from stumps. Root suckers mayalso develop. The centre of holes is often defec-tive with purplish brown heart rot due to inclp-lent decay. Therefore, logs are often quarter-sawn before drying and the central portion ofthe hole is often discarded. Trees are often

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CURTISiA 255

256 TIMBERS2

heavily parasitized by fungi and insects. Theseedlings are liable to attack by damping-offfungi.

Genetic resources and breeding Curtisiodentoto occurs only very locally in tropical at-rica. In South Africa it has a wider distribu-tion, but taller specimens have become rarebecause of exploitation for its timber in thepast. The popularity of the bark as medicinealso contributes considerably to the existingpressure on populations of Curtisio dentata. InKwaZulu-Natal Province in South Africa, thespecies has already been considered as vulner-able and declining in 1988 because of over-exploitation of the bark.

Prospects Curtisio dentoto is too uncommonin tropical Africa to have prospects as a coin-mercialtimber tree. This seems also the case in

South Africa, where it is more widespread andcommon, because large trees have become rarebecause of over-exploitation and because of therather slow growth under natural conditions. Itseems to have much better prospects as orna-mental shrub or tree. It deserves more re-

search attention concerning its pharmacologi-cal properties. Recent investigations demon-strated pronounced antibacterial and antifun-gal properties, supporting the use of Curtisiodentoto in traditional medicine

Major references Bolza & Keating, 1972;Cannon, 1978c; Coates Palgrave, 1983; Notten,2004; Takahashi, 1978.

Other references Cunningham, 1990; Fan& Xiang, 2003; Grace at a1. , 2002a; Grace at al. ,2002b; Grierson & ato1ayan, 1999; Kubitzki,2004; Palmer & Pitman, 1972-1974; Shat ata1. , 2008a; Shai at a1. , 2008b; van Wyk, vanOudtshoorn & Gencke, 1997.

Authors R. H. M. J. Lemmens

.

Cussonio zimmermonnii - coild

ness and bleeding after childbirth, and also tofacilitate childbirth. Root decoctions are takenor used as a wash to treat fever and malaria,and administered against gonorrhoea. Themarrow of stem and branches is eaten as atreatment of epilepsy.

Properties The heartwood is whitish andindistinctly demarcated from the sapwood. Thetexture is moderately fine. The wood is light-weight, with a density of about 400 kg/ina at12% moisture content, soft and brittle. Thewood is easy to air dry, with little degrade. It iseasy to saw and work, and planes to a smoothsurface. The wood is not durable; it is suscepti-ble to fungal attacks such as blue stainExtracts of the root bark showed promisingresults in the Yarninobutyric acid type-A(GABAA) receptor binding assay, and alsoshowed in-vitro antiprotozoal activity againstfryponosomo brucet rhodesiense and PIOsmo-churn foietporum. Several polyacetylenes andstigmasterol have been isolated from the rootbark. Some of the polyacetylenes showed activ-ity against 77yponosomo bri, cat rhodesiense,Tryponosomo cruzi, Plusmodium foieiporumand Letshmonio donouoni. These results sup-port the use of Cussonio atminermonnii in tra-ditionalmedicine

Botany Small to medium-sized tree up to25(-45) in tall; hole straight; bark surface greyto greenish grey, fissured and scaly; crownrounded, dense; twigs glabrous. Leaves ar-ranged spiralIy, clustered at ends of branches,digitately compound with 5-7(-9) leaflets;stipules partly fused with petiole; petiole up to50 cm long; leaflets sessile, elliptical to obo-vate, 5-25 cm x 2-8 cm, curieate at base, acute

CussONIAziMMERMANNiiHarms

Protologue Bot. Jahrb. Syst. 53: 361 (1915)Family AraliaceaeVernacular names Mbomba mail, inpapayi

inwitu (Sw)Origin and geographic distribution Cusso-

it10 21mmermonnzz occurs in eastern Kenya,eastern Tanzania and northern Mozambique.

Uses The wood is used for coffins, drums,boats and carvings. It is suitable for furnitureand light interior construction. It is also usedas firewood, although of low quality for thispurpose.

Roots are used in the treatment of mental ill-

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to acuminate at apex, margins toothed to near-Iy entire, papery to leathery, glabrous, pinnate-Iy veined. Inflorescence a spike-like raceme 8-34 cm long, up to 14 together at ends ofbranches; bracts up to 4 min long, often manyclose together at base of inflorescence. Flowersbisexual, regular, usually 5-merous, 4-8 min indiameter, greenish white; pedice1 3-5 min long;calyx with short teeth; petals free; stamensalternating with petals, inserted on a disk;ovary inferior, 2-celled, styles 2, short, fused atbase. Fruit an obconical to globose drupe-likeberry up to 6 min long, greenish white, gla-brous or slightly hairy. Seeds ovoid-globose,slightly compressed, with ruininate endosperm.Cussonio comprises about 20 species and isrestricted to mainland tropical Africa. Thewood of several other species is used, but isusually considered of little value and onlyavailable in smaller sizes because the tree

holes are often smaller and more poorly shapedthan those of Cussonio zimmermo, tnn.

One of the most widely distributed species isCussonio orboreo Hochst. ex A. Rich. , of whichthe primary use is for medicinal purposes, as isthe case for Cussonio spiedto Thunb. occurringin mountain forest in East and southern Africa.

The whitish, soft wood of Cussonio boricoensisAubr6v. & Fellegr. , a medium-sized tree up to20(-30) in tall with a hole diameter up to 60(-100) cm occurring from Liberia to Nigeria (alt-hough it has been suggested that it is only na-tive to Ghana and planted elsewhere), is usedin Ghana for drums, utensils and tool handles.The bark showed antinociceptive and anti-inflammatory activities, possibly due to thepresence of terpenoids such as 23-hydroxyur-sonc acid. Saponins have also been isolatedfrom the bark, as wellas stigmasterolThe wood of Cussonio hoistit Harms ex Engl. isalso whitish and soft, and used for doors, bee-hives, utensils, tool handles and musical in-struments. Cussonio hoisttits a small to medi-

urn-sized tree up to 15(-20) in tall with a holediameter up to 60(-100) cm occurring from DRCongo to Ethiopia and Somalia, and south toTanzania. A bark decoction is used to expeltheplacenta after childbirth, to stop vomiting, andagainst diarrhoea in livestock. Bark and rootdecoctions are administered to improve thehealth of children and to treat blood diseases.

The leaves serve as forage for goats, donkeysand camels. A bark extract showed pronouncedactivity against Trichomonos Dogino!is; thepentacyclic triterpenoid bederagenin was ISO-lated as the active constituent.

Ecology Cussonio zimmermonniiis found incoastal evergreen forest and bushland, also inforest margins, up to 400 in altitude. It occursin rainforest as well as drier evergreen forest

Genetic resources and breeding Cusso-itto zimmermonnii has a rather limited distri-

bution in an area where there is much pressureon the forest. It may easily become threatenedby genetic erosion and protection measuresmay be necessary, as is the case for the WestAfrican Cussonio boricoensis, which is alreadyincluded in the IUCN Red List as vulnerable

Prospects The wood of Cussonio zimmer-monniiis of rather poor quality, as is the casefor other Cussonio spp. , but it will probablyremain of some importance for local applica-tions. Interesting pharmacological activities ofthe bark have been demonstrated. These war-

rant more attention, also in the view of thepromising results of pharmacological researchon other Cussortio spp. and their constituents

Major references Beentje, 1994; Chikamaiat al. , undated; Senn, 2006; Senn at a1. , 2007;Tennant, 1968.

Other references Cannon, 1978b; Gessler ata1. , 1994; He at a1. , 2003; Heme & Heme, 1988;Holingren at a1. , 2004; Jansen, 1981; Kokwaro,1993; Maundu & Tengnd. s (Editors), 2005;Maundu at a1. , 2001; Neuwinger, 2000

Authors R. H. M. J. Lemmens

GINIERA 257

ConERAMACROURAK. Schum

Protologue Bot. Jahrb. Syst. 33(2): 352(1903).

Family RubiaceaeSynonyms Cuuiero dialonensisAchev. (1920)Origin and geographic distribution CUDie-

ro macrouro is distributed from Guinea and

Sierra Leone eastward to Cameroon

Uses Larger holes are used for dug-out ca-whereas branches are used asnoes, yam

stakes

Botany Shrub or smalltree up to 12 in tall;spreading; branches with oppositecrown

spines between the nodes; twigs often hollowand ant-infested. Leaves opposite, simple andentire; stipules often with ear-shaped lobes,persistent; petiole short; blade oblong-ellipticalto narrowly oblong, more or less asymmetrical,up to 20 cm x 6 cm, base obliquely curieate torounded, margins wavy, leathery, glabrous,shiny, pinnately veined with 6-10 pairs of Iat-eral veins. Inflorescence an axillary cyme c. 7.5cm long, branched from the base; bracts

258 TIMBERS2

.

Cuuiero macrouro - wild

thread-like, glabrous. Flowers bisexual, regu-Iar, 5-merous, white turning yellow, fragrant;bracteoles subulate, c. I cm x I mm; calyx gla-brous, lobes subulate, c. I cm x I min; corolla c.2 cm long, lobes lanceolate, with thread-liketips at apex; stamens inserted in corolla tube,alternating with corolla lobes; ovary inferior, 5-celled. Fruit a nearly globose drupe c. 4.5 cmlong, 5-ridged.In Benin CUDiero mererouro flowers in Augustand fruits ripen in November-FebruaryCUDiero comprises about 30 species, all in trop-icalAfrica, but the genus is poorly known. It ispossibly close to Vongueriel!0, but the group ofVonguerieoe, in which CUDiero is classified, isin need of revision to elucidate the relation-

ships between taxaCt, uiero acuti/10ro DC. is a shrub or smalltreeup to 10 in tall distributed from Guinea andSierra Leone to Gabon and DR Congo. In SIer-ra Leone the wood is used for making housewattles. In Nigeria the leafis made into a laxa-tive for children

Ecology Cuuiero ingcrouro occurs in wetlocations in savanna, secondary forest, forestedges and swamp forest.

Genetic resources and breeding In viewof its wide distribution and wide range ofhabi-tats, CUDiero macrouro seems not threatenedwith genetic erosion

Prospects CUDiero mererouro is only locallyused as a source of wood for canoes and stakes

Information on its wood properties is lacking,but in view of its limited actual use and small

size its Importance is unlikely to increaseMajor references Ako6gninou, van der

Burg & van der Maesen (Editors), 2006; Au-

br6ville, 1959c; Burki11, 1997; Hepper & Keay,1963.

Other references Fauna & Flora Interna-

tional, undated; Hawthorne, 1990; Hawthorne,1995; Hawthorne & Jongkind, 2006; Irvine,1961; Tchout0, 2004.

Authors M. Brink

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CYNOMETRAALD<ANDRIC. H. Wright

Protologue H. H. Johnst. , Uganda Prot. I:325 (1902)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Vernacular names Tuna (Sw)Origin and geographic distribution Cyno-

metro o1exondri occurs from DR Congo east towestern Uganda and western Tanzania

Uses The wood, traded from Uganda as 'inu-himbi' or 'Uganda iron wood' is used for heavyconstruction, heavy flooring, ship building,vehicle bodies, mine props, railway sleepers,furniture, cabinet work, sporting goods, toys,novelties, agricultural implements, tool han-dles, draining boards, carving and turnery. Ithas a high resistance to abrasion and is verysuitable for heavy-duty flooring. The wood isalso used as firewood and for charcoal produc-tion

In Uganda Cynometro origxondriis planted tocontrol erosion and occasionally as an orna-mental shrub. Flowering trees are a source ofpollen for honey bees. The bark serves to makebaskets and is also used as an arrow poison. InDR Congo bark powder is applied to woundsCaterpillars feeding on the leaves are collectedin DR Congo to serve as baitforfishing

I.

Cynometr0 o1exondr, - colld

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Production and international trade The

wood of Cynometr0 o1exondriis used locallyand only occasionally traded on the interna-tional market, mostly as flooring blocks andstrips. Trade statistics are not available. MuchCynometr0 o1exondritimber has been exportedin the 1950s and 1960s from western Ugandato Europe

Properties The heartwood is pinkish brown,darkening upon exposure to reddish brownwith darker streaks. It is distinctly demarcatedfrom the 5-7.5 cm wide, greyish white sapwoodwhich turns pale yellow upon exposure. Thegrain is usually interlocked, texture fine.The wood is moderately heavy to heavy, with adensity of 760-940 kg/ina at 12% moisture con-tent. It air dries slowly with only slight distor-tion, but checking and end splitting may occur.Drying should therefore be done with care,especially kiln drying. The rates of shrinkageare moderate, from green to 12% moisture con-tent about 2.5% radial and 4.5% tangential,and from green to oven dry 4.3% radial and7.8% tangential. Once dry, the wood is onlymoderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 109-202 N/mm2, modulus of elasticity10,900-17,500 N/mm2, compression parallel tograin 51-72 N/mm2, shear 17-23 N/mm2,cleavage 22 N/mm, Janka side hardness11,300-11,830 N, Chalais-Meudon side hard-ness 5.4 and Janka end hardness 13,160 NThe wood is difficult to saw and work, and hassevere blunting effect on cutting edges and sawteeth; the use of tungsten-tipped cutting edgesis recommended. Resin may build up on sawteeth. In planing operations a 15' cutting angleis recommended to obtain a good finish, andcutting edges should be kept sharp. The woodtakes a fine polish. Boring and mortising prop-erties are less favourable with a tendency ofsphtting at exits. Pre-boring is needed for nail-ing and screwing, butthe wood holds nails andscrews well. It glues moderately well and canbe stained satisfactorily requiring little fillingIt turns well. The steam bending properties aremoderate. The wood is durable, being resistantto termite and Lyetus attacks, but occasionallyliable to pinhole borer and longhorn beetle at-tacks. It is also durable in water and resistant

to marine borers. The heartwood is resistant to

impregnation with preservatives, but the sap-wood is permeable

Description Evergreen medium-sized to largetree up to 50 in tall; hole branchless for up to25 in but often shorter, straight and cylindrical

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CryOMETRA 259

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Cynometro diexo, Idri - I, base of bale, 2, leg/ytwig, ' 3, flowering twig, ' 4, flower, 5, fruits.Redrown grid adopted by Ishok Syomsudin

or twisted, up to 150(-180) cm in diameter,with large, thin plank buttresses up to 5 inhigh; bark surface smooth but irregularly flak-ing, grey to reddish grey or pale brown, innerbark reddish brown; crown flattened withspreading branches; twigs short-hairy but be-coining glabrous. Leaves arranged spiralIy,panpinnately compound with 2(-3) pairs ofleaflets; stipules linear, 0.5-I cm long; petioleup to 0.5 cm long, rachis up to 3.5 cm long,widely grooved above; leaflets opposite, sessile,obliquely lanceolate to elliptical or ovate, I-10cm x 0.5-4 cm, upper ones largest, acuminateat apex, glabrous. Inflorescence an axillary orterminal panicle up to 6 cm long, short-hairy,with densely crowded flowers; bracts smallFlowers bisexual, nearly regular, whitish orsometimes pinkish, fragrant; pedicel up to 1.5cm long, glabrous, jointed near apex and per-SIStent after shedding of flowers; sepals 4(-5),c. 4.5 mm long, reflexed; petals 5, free, obovate-lanceolate, 5-6 mm long; stamens usually 10,free, 6-8 mm long, slightly unequal; ovary su-perlor, c. 2.5 min long, with stipe c. 2 min long,hairy, I-celled, style 3-4 mm long. Fruit an

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260 TIMBERS2

obliquely oblong to obovate, flat pod 5-10 cm x3-5 cm, with 0.5-I cm long stipe, rounded orshort-pointed at apex, smooth and brown, de-hiscent with 2 woody, twisting valves, few-seeded. Seeds rounded, flat, 1.5-2 cm in diame-ter, brownish. Seedling with epigeal germina-tion; hypocoty1 4-11 cm long, epicoty1 5.5-13cm long, short-hairy; first 2 leaves opposite,with 4 leaflets, subsequent leaves alternate.

Other botanical information Cynometrocomprises about 90 species and occurs in thetropical regions of the world. Approximately 25species can be found in mainland tropical Mri-ca and about 10 in Madagascar. Tropical Asiaand tropical America have about 25 specieseach

Cy, tometro webberi Baker f. also occurs in EastAfrica, where it is restricted to coastal dry ev-ergreen forest in south-eastern Kenya andeastern Tanzania. It is a smalltree up to 12 intall with short hole. The pinkish brown, heavyand durable wood is used for heavy construc-tion and furniture. Cynometro webberiis in-cluded in the IDCN Red List as vulnerable

Anatomy Wood-anatointcal description orAWAhardwood 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-Iygona1; 24: intervessel pits minute (S 4 pin);29: vestured pits; 30: vessel-ray pits with dis-tinct borders; similar to intervessel pits In sizeand shape throughout the ray cell; 41: meantangential diameter of vessellumina 50-100pin; 47: 5-20 vessels per square initlimetre; 48:20-40 vessels per square minimetre; 58: gumsand other deposits in heartwood vessels. 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);80: axial parenchyma allform; (81: axial paren-chyma lozenge-allform); (82: axial parenchymawinged-allform); 83: axial parenchyma conflu-ent; 86: axial parenchyma in narrow bands orlines up to three cells wide; (91: two cells perparenchyma strand); 92: four (3-4) cells perparenchyma strand; 93: eight (5-8) cells perparenchyma strand. Rays: 96: rays exclusivelyuniseriate; 97: ray width I-3 cells; 104: all raycells procumbent; (106: body ray cells procum-bent with one row of upright and/or squaremarginal cells); 1/6:;;: 12 rays per mm. Stonedstructure: 1/8: all rays stoned; 120: axial pa-

Tenchyma and/or vessel elements stoned; 121fibres stoned. Mineral inclusions: 136: pris-

matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells; (143prismatic crystals in fibres)(P. Mugabi, P. Baas & H. Beeckman)

Growth and development Young plantsrequire shade untilthey are established. Treesgrow slowly. In natural forest in westernUganda, it has been estimated that trees reacha hole diameter of 60 cm after about 80 years'Older trees have large, widely spreading but-tresses and spreading roots, making estab-lishment on shallow soils and slopes more easyTrees lose their leaves irregularly. In DR Con-go they flower in March-May. In the Budongoforest reserve in western Uganda, chimpanzeeshave a preference for Cynometro diexondritrees to build their nests. The presence of ve-SICu}ar-arbuscular mycorrhizae has been rec-orded

Ecology Cynometro dietondrioccurs in ever-green and semi-deciduous forest at 700-1200(-1400) in altitude. In Budongo forest reserve inwestern Uganda, where dense stands ofCynometro die"undri occur, the temperaturerange is 11-30'C, mean temperature 21'C andmean annual rainfall about 1500 mm, with apronounced dry season in January-FebruaryForest dominated by Cynometr0 o1exondriisconsidered a climax vegetation type. Cynome-tr0 o1exondriis susceptible to forest fires.

Propagation and planting Natural regen-eration of Cynometr0 o1exondriis good in natu-ral forest in western Uganda with differenthistories of logging and other disturbances.Cynometr0 o1exondriis normally propagatedby seeds and wildings. There are about 290seeds per kg. Seeds are usually collected dur-ing the raining season from the ground, andsown in pots. Fretreatment of seeds before sow-ing is not necessary. Storage of seeds is diffi-cult due to insect infestation; it is recommend-ed to add ash or apply insecticides.

Management In the 1950s and 1960s whenintensive logging took place in forest in west-ern Uganda, Cynometr0 o1exondri was oftenconsidered an undesired species and treatedwith arboricides. The reason was that in later

stages of forest succession, the slow-growingCynometro diexondri suppressed more highlyvalued timber species such as Louoo trichili-oides Harms, Mineio excelso orelw. ) C. C. Berg,Khoyo grithotheco (WeIw. ) CDC. and Errton-drophrogmo spp. However, in several forests,such as the Budongo forest reserve, Cynometro

o1exondriis stinthe most dominant tree spe-cies. Trees can be managed by coppicing, prun-ing and pollarding.

Harvesting Cynometr0 o1exondriis not easyto cut because of the often large and high but-tresses, the hard and heavy wood, and the of-ten hollow holes in large trees.

Genetic resources Cynometro diexondrLisfairly widespread and locally common to evendominant in the forest. It does not appear to beliable to genetic erosion

Prospects The low growth rate, the hard andheavy wood, and the often poor hole shape ofCynometro alexondri are serious drawbacks forcommercialization efforts. However, its fairregeneration rate in natural forest, and thehigh natural durability and high resistance toabrasion of the wood offer possibilities forcommercial timber production for special PUT-poses such as heavy-duty flooring, as long aslong rotation cycles are acceptable. Furtherresearch is recommended on the growth ratesunder favourable conditions and appropriatemanagementsystems

Major references ATIBT, 1986; Bolza &Keating, 1972; Brenan, 1967; Eggeling & Dale,1951; Fouarge, Quoilin & Roosen, 1970;Katende, BITnie & Tengnas, 1995; Takahashi,1978.

Other references Chikamaiet al. , undated;Dawe, 1910; Liben, 1970b; Mwavu & Witkowski,2009; Neuwinger, 2000; Plumptre, 1996; Shell,Jennings & SaviU, 2000; Tailfer, 1989; Terashima& Ichikawa, 2003; Terashima, Ichikawa &Sawada, 1988; Wilczek at a1. , 1952.

Sources of illustration Eggeling & Dale,1951.

Authors JR. Cobbinah & E. A. Obeng

.

CryOMETRA 261

Cynometro orionto- wild

wood of Cynometro orionto is used locally andonly rarely traded on the international market

Properties The heartwood is reddish brownwith darker streaks, and distinctly demarcatedfrom the 2.5-7.5 cm wide, yellowish to pinkishbrown sapwood. The grain is usually inter-locked, texture fineThe wood is heavy, with a density of 910-1000kg/ing at 12% moisture content. It air driesslowly with a tendency of checking and endsplitting. Drying should therefore be done withcare. The rates of shrinkage are moderatelyhigh, from green to oven dry 5.2% radial and9.3% tangential. Once dry, the wood is not verystable in service.

At 12% moisture content, the modulus of rup-ture is 126-187 N/min2, modulus of elasticity14,700-17,840 N/min2, compression parallel tograin 72-83 N/min2, cleavage 18.5-19.5 N/min,Janka side hardness 11,700 N, Chalais-Meu-don side hardness 8.5-14.8 and Janka end

hardness 12,940 N.The wood is difficult to saw and work, and hassevere blunting effect on cutting edges and sawteeth; the use of tungsten-tipped cutting edgesis recommended. In planing operations a 15'cutting angle is recommended to obtain a goodfinish, and cutting edges should be kept sharp.Pre-boring is needed for nailing and screwingThe corrosion rate of iron and steelin the wood

is high. The wood glues moderately well andcan be stained and polished satisfactorily re-quiring little filling. It turns well. The steambending properties are moderate. The wood isdurable, being resistant to termite and Lyetusattacks, but occasionally liable to pinhole borerand longhorn beetle attacks. The heartwood is

CYNOMETRAANANTA Hutch. & Dalziel

Protologue F1. W. trop. Mr. I(2): 331 (1928)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Origin and geographic distribution Cyno-

metro orionto is restricted to West Africa,where it occurs from Liberia east to Ghana.

Uses The wood, known as 'apom6' or 'arian-ta', is suitable for heavy construction, heavyflooring, bridge building, exterior Joinery, Inte-nor trim, ship building, vehicle bodies, mineprops, poles, railway sleepers, sporting goods,toys, novelties, agricultural implements, pat-tern making and turnery

Production and international trade The

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262 TIMBERS2

resistant to impregnation with preservatives,butthe sapwood is moderately permeableSome jinidazoline alkaloids have been isolated

from the leaves.

Description Evergreen medium-sized tofairly large tree up to 35(-45) in tall; bolebranchless for up to 18 in, straight to twisted,often angular and knotty, up to 120 cm in di-ameter, with thin plank buttresses up to 3(-4.5) in high; bark surface irregularly flakingwith small scales, grey with yellowish to red-dish marks, inner bark fibrous, reddish, becoin-ing reddish brown upon exposure, exuding areddish resin; crown rounded or widely spread-ing, heavily branched; twigs glabrous, withlenticels. Leaves arranged spiralIy, panpin-nately compound with I pair of leaflets; stip-tiles triangular, c. I min long, caducous; petiole0.5-I cm long, rounded; petiolules 2-4 mmlong; leaflets opposite, sickle-shaped, 3.5-15 cmx I-5 cm, acute to acuminate at apex, thin-leathery, glabrous. Inflorescence an axillary orterminal panicle up to 10 cm long, short-hairy,with densely crowded flowers; bracts small.Flowers bisexual, nearly regular, whitish; pedi-cel c. 0.5 cm long; sepals 4(-5), c. 4 mm long,

reflexed; petals 5, free, obovate, 3-5 mm long;stamens 10, free, 5-7 min long; ovary superior,c. I mm long, with stipe c. 0.5 mm long, hairy,I-celled, style 3-4 min long. Fruit an obliquelyoblong to obovate, flat pod 8-12 cm x 4-5 cm,with short stipe, short-pointed at apex, smoothand brown, dehiscent with 2 woody, twistingvalves, I-2-seeded. Seeds rounded to elliptical,flat, 2-2.5 cm long, pale brown. Seedling withepigeal germination; hypocoty1 3-9 cm long,epicoty1 4-18 cm long; cotyledons 1.5-2.5 cmlong, thick and fleshy; first 2 leaves opposite,with 2 leaflets, subsequent leaves alternate.

Other botanical information Cynometrocomprises about 90 species and occurs in thetropical regions of the world. Approximately 25species can be found in mainland tropical Mri-ca and about 10 in Madagascar. Tropical Asiaand tropical America have about 25 specieseach

Cynometru mego!ophyllo Harms, a small tomedium-sized tree up to 20 in tallwith hole upto 100 cm in diameter, is also restricted toWest Africa, occurring from C6te d'TVoire eastto Nigeria, mostly along rivers. Its reddishbrown, heavy, hard and durable wood is usedfor poles, posts and frames in house buildingBark decoctions are used in traditional medi-

cine to treat kwashiorkor, and leaf decoctionsto treat measles and chickenpox. Cynometromegolophyllu has been suggested to be usefulfor reafforestation of swampy areasCynometro Dogelii Hook. f. is a smalltree up to10(-20) in tallwith hole up to 100 cm in diame-ter, occurring from Senegal east to Nigeria. Itsreddish brown, hard wood is locally used fortool handles; it is also used as firewood. Thefoliage is fed to cattle. The seed is reportedIyedible.

Anatomy Wood-anatointcal description CAWAhardwood 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-Iygona1; 24: intervessel pits minute (S 4 pin);29: vestured pits; 30: vessel-ray pits with dis-tinct borders; similar to intervessel pits in sizeand shape throughout the ray cell; 42: meantangential diameter of vessellumina 100-200ILm; 46: S 5 vessels per square minimetre; 47:5-20 vessels per square minimetre; 58: gumsand other deposits in heartwood vessels. Tra-cheids and fibres: 61: fibres with simple to inI-nutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled; 70:

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Cynometro orionto - I, base of 601e, 2, Ieo/ytwig, . 3, flower, . 4, fruit.Redrown and oddpted by/shoh Syomsudin

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fibres very thick-walled. Axial parenchyma(76: axial parenchyma diffuse); (77: axial pa-renchyma diffuse-in-aggregates); 80: axial pa-renchyma anform; 82: axial parenchymawinged-allform; 83: axial parenchyma conflu-ent; 85: axial parenchyma bands more thanthree cells wide; (86: axial parenchyma in nar-row bands or lines up to three cells wide); (89:axial parenchyma in marginal or in seeminglymarginal bands); 92: four (3-4) cells per paren-chyma strand; (93: eight (5-8) cells per paren-chyma strand). Rays: 96: rays exclusively uni-senate; 97: ray width I-3 cells; 104: all raycells procumbent; (106: body ray cells procum-bent with one row of upright and/or squaremarginal cells); 1/5: 4-12 rays per mm. Stonedstructure: (1/8: all rays stoned); (120: axialparenchyma and/or vessel elements stoned);(122: rays and/or axial elements irregularlystoned). Mineral inclusions: 136: prismaticcrystals present; 142: prismatic crystals inchambered axial parenchyma cells.(L. Awoyemi, P. E. Gasson & E. A. Wheeler)

Growth and development Older trees havelarge, widely spreading buttresses and spread-ing roots, making establishment on shallowsoils and slopes more easy. Shedding of oldleaves is immediately followed by new flushes,which are initially brilliant red. In Liberiatrees flower in September-October, in C6ted'TVoire in October-December. In Liberia, C6ted'ivoire and Ghana fruits ripen in December-January, in C6te d'Ivoire also in June-July.The fruits are explosiveIy dehiscent, dispersingthe seeds over short distances

Ecology Cynometro orionto occurs mainly inhumid evergreen forest, only occasionally inmoist semi-deciduous forest. It is most abun-

dant in regions with an annual rainfallof morethan 1500 mm and a dry period of less than 4months. It usually occurs on sandy soils, but inC6te d'Ivoire it has been recorded to preferhumid, even slightly swampy soils.

Propagation and planting Seedlings andsaplings are often abundant near mother treesThey are shade tolerant, but in Liberia it hasbeen reported that saplings of more than 2 intall are uncommon. It seems that regenerationis better in logged-over forest. There are about650 seeds per kg, and the germination rate ishigh within 15 days after sowing.

Management Locally Cynometro orlonto hasgregarious stands and may even be semi-dominant. In forest in Liberia it has been re-

ported to occur at an average density of 03holes of more than 60 cm in diameter per ha,

but locally it is much more common, in someareas having an average density of 3 largeholes per ha. In C6te d'TVoire and Ghana it isalso locally abundant.

Harvesting Cynometro orionto is not easy tocut because of the often large and high but-tresses and the hard and heavy wood. In Gha-na the recommended minimum hole diameter

allowed for felling is 70 cmGenetic resources Cynometro orionto does

not appear to be liable to genetic erosion atpresent because it occurs locally in large standsand does not seem to be subject to selectivelogging. However, with the ongoing fragmenta-tion of evergreen forest in West Africa, moni-tonng of populationsis recommended

Prospects The hard and heavy wood andthe often poor hole shape of Cynometro oriontoare serious drawbacks for commercialization

efforts. However, its fair regeneration rate innatural forest, and the high natural durabilityand high resistance to abrasion of the woodoffer possibilities for commercial timber pro-duction for special purposes such as heavy-duty flooring, as long as long rotation cyclesare acceptable. Research is recommended onthe growth rates under favourable conditionsand appropriate management systems.

Major references Bolza & Keating, 1972;Burki11, 1995; de Koning, 1983; Holingren ata1. , 2004; 0teng-Amoako (Editor), 2006; Sal-Ienave, 1955; Sallenave, 1964; Takahashi,1978; Voorhoeve, 1979

Other references Arbonnier, 2004; Aubr6-vine, 1959b; Cooper & Record, 1931; de inMensbruge, 1966; Hall & Swame, 1981; Haw-thorne, 1995; Hawthorne & Jongkind, 2006;Irvine, 1961; firyn & Fobes, 1959; Kunkel,1965; Neuwinger, 2000; Normand & Paquis,1976; Taylor, 1960.

Sources of illustration Oteng-Amoako(Editor), 2006; Taylor, 1960; Voorhoeve, 1979

Authors C. Essien & A. A. Oteng-Amoako

CYNOMETRA 263

CryOMETRAHANKEIHarms

Protologue Notizbl. Bot. Gart. Berlin-Dahlem, App. 21(2): 39 (1911)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Origin and geographic distribution Cyno-metro hurtkei occurs from south-eastern Nige-ria and western Cameroon to DR Congo.

Uses The wood, known in Cameroon as 'nko-kom' or 'riganga', is used for heavy construc-

264 TIMBERS2

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Cynometro honhei- wild

tion, bridge building, heavy flooring, joinery,railway sleepers, tool handles and sliced ve-neer. It is suitable for ship building, vehiclebodies, mine props, sporting goods, toys, novel-ties, agricultural implements, carving andturnery.

Properties The heartwood is reddish brown,often with slightly shiny ribbon-like markingson quarter-sawn surfaces, and distinctly de-marcated from the up to 3 cm wide, pale brownsapwood. The grain is straight or interlocked,texture fine

The wood is heavy, with a density of (740-)870-980 kg/ina at 12% moisture content. It airdries slowly with a tendency of checking andend splitting. Drying should therefore be donewith care. The rates of shrinkage are high,from green to oven dry 5.1-5.7% radial and9.9-11.0% tangential. Once dry, the wood isnot stable in service

At 12% moisture content, the modulus of rup-ture is 150-213 N/mm2, modulus of elasticity15,500 N/mm2, compression parallelto grain 75N/mm2, cleavage 21-24 N/mm and Chalais-Meudon side hardness 7.2-10.3

The wood saws slowly but quite well. In plan-ing operations a 15" cutting angle is recoin-mended to obtain a good finish, and cuttingedges should be kept sharp. Pre-boring is need-ed for nailing and screwing, butthe wood holdsnails and screws well. Sanding gives smoothsurfaces and the wood polishes and turns wellIt is durable, being resistant to fungal, termite,Lyetus and marine borer attacks, but occasion-ally liable to pinhole borer attack. The heart-wood ls resistant to impregnation with pre-servatives, but the sapwood is moderately

permeable. Wood dust may cause irritation tothe respiratory tracts and eyes.Some lintdazole alkaloids have been isolated

from the bark and seeds

Botany Evergreen medium-sized to largetree up to 45 in tall; bole branchless for up to22 in, usually straight and cylindrical, up to150 cm in diameter, with thin buttresses up to8 in high; bark surface flaking with smallscales, grey with reddish brown marks, innerbark brown, becoming reddish brown uponexposure; crown dome-shaped, with ascendingbranches; twigs short-hairy, with reddish lenti-cels. Leaves arranged spiralIy, panpinnatelycompound with 7-15 pairs of leaflets; stipuleslinear, c. 0.5 cm long, caducous; petiole andrachis together 2-10 cm long, grooved aboveand often slightly winged, short-hairy; leafletsopposite, sessile, oblong, 0.5-3 cm x 0.2-I cm,rounded to slightly notched at apex, glabrousor slightly hairy below. Inflorescence an axil-Iary or terminal panicle up to 12 cm long,short-hairy; bracts small. Flowers bisexual,nearly regular, whitish; pedice1 0.5-1.5 cmlong, short-hairy, jointed near apex and persis-tent after shedding of flowers; sepals 4, ovateto rounded, 3-3.5 mm long, reflexed; petals 5,free, obovate to lanceolate, 3-4 min long; sta-mens 10, free, 5-6 min long, slightly unequal;ovary superior, ellipsoid, c. 2 min long, withshort stipe, hairy, I-celled, style 2.5-3 mmlong. Fruit a slightly obliquely oblong-obovate,flat pod 6-9 cm x 3-4 cm, with short stipe,short-pointed at apex, smooth and brown, de-hiscent with 2 woody, twisting valves, I-2-seeded. Seeds rounded, dark brown. Seedlingwith epigeal germination; hypocoty1 2.5-3.5 cmlong, epicoty1 3-6.5 cm long; cotyledons oblong,thick and fleshy, wrinkled; first 2 leaves OPPo-site, with many leaflets, subsequent leavesalternate

The plants described as Cynometro hanhet inDR Congo differ slightly from those in Nigeriaand Garnero0n and may represent another,undescribed species. The leaflets are oftensmaller and completely glabrous, the rachis isslightly winged between the pairs of leaflets,and the pedicels are longerCynometro comprises about 90 species andoccurs in the tropical regions of the world. Ap-proximateIy 25 species can be found in main-land tropical Africa and about 10 in Madagas-car. Tropical Asia and tropical America haveabout 25 species eachThe distribution area of Cynometro inarinii011v. is largely similar to that of Cynometro

..

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hankei, occurring from south-eastern Nigeriato south-western DR Congo, often along riversIt is a small to medium-sized tree up to 17 intall with short, irregular hole. The reddishbrown and hard wood is occasionally used, forinstance to make fish traps. In Congo barkdecoctions are taken to treat cough; the bark isconsidered purgative and emetic

Ecology Cynometro hunhei occurs in ever-green forest up to 1000 in altitude, usually onwell-drained and sandy soils. It is often foundin smallgroups

Management In western Cameroon the ave-rage density of Cynometro heritheitrees with ahole diameter of more than 60 cm is 04-0.5 perha, and the average timber volume 2.6-3.5inalba. The minimum hole diameter for bar-

vesting is 60 cm in Cameroon. In DR Congo atree of 35 in tallwith a hole of 18 in long and adiameter of 60 cm yielded 3.5 ina of wood.

Genetic resources and breeding Cyno-metro honheiis fairly widely distributed andnot uncommon, and does not seem to bethreatened by genetic erosion. However, thepopulations In south-western Nigeria andwestern Cameroon could easily become endan-gered with ongoing deforestation

Prospects The hard and heavy wood andthe often high buttresses of Cynometro honheiare serious drawbacks for commercialization

efforts. However, the high natural durabilityand high resistance to abrasion of the woodoffer possibilities for commercial timber pro-duction for special purposes such as heavy-duty flooring, as long as long rotation cyclesare acceptable. Research is recommended onthe growth rates under favourable conditionsand appropriate management systems. Thebiosystematics of populations in Nigeria andCameroon on the one hand and in DR Congoonthe other should be studied because they mayrepresent different species

Major references Bolza & Keating, 1972;Fouarge, G6rard & Sacr6, 1953; Fouarge, Sacr6& Mottet, 1950; Takahashi, 1978; Vivien &Faure, 1985

Other references Aubr6ville, 1968; Aubr6-vine, 1970; Burki11, 1995; Neuwinger, 2000;Raponda-Walker & Sillans, 1961; Tailfer, 1989;Waterman & Faulkner, 1981; Wilczek at al. ,1952

Authors R. H. M. J. Lemmens

CryoMETRALUJAE Dewild

Protologue min. Mus. Congo Belge, Bot. , s6r5, I: 250 (1906).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Origin and geographic distribution Cyno-metro Jujue occurs in southern Gabon, south-ern Congo, south-western DR Congo and Ca-binda (Angola)

Uses The wood is suitable for heavy andhydraulic construction, bridges, joinery, doors,window frames, vehicle bodies, furniture, rail-way sleepers, sporting goods, tool handles andprecision equipmentThe bark is used in traditional medicine in

Congo to treat toothache, cough and rheuma-tism

Properties The heartwood is reddish brownwith a purplish tinge and with yellowish brownmarkings, distinctly demarcated from the up to8 cm wide, brownish yellow sapwood. The grainis usually straight, sometimes interlocked,texture fine

The wood is heavy, with a density of (760-)855-995 kg/ina at 12% moisture content. Therates of shrinkage during drying are moderate-Iy highAt 12% moisture content, the modulus of rup-ture is 174-205 N/min2, modulus of elasticity15,000 N/mm2, compression parallel to grain55-90 N/min2, cleavage 21.5 N/min and Cha-Iais-Meudon side hardness 13.5

The wood saws slowly. Planing may occasional-Iy result in rough surfaces due to the presenceof interlocked grain. Pre-boring is needed fornailing and screwing, but the wood holds nailsand screws well. It is durable, being resistant

CYNOMETRA 265

.

Cynometro Iujoe - wild

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266 TIMBERS2

to fungal and termite attacks, but liable tomarine borer attack

Several jinidazole alkaloids have been isolated

from the bark and leaves.

Botany Evergreen small to medium-SLzedtree up to 20(-30) in tall; hole branchless for upto 12(-20) in, up to 45(-65) cm in diameter;bark surface smooth, greenish brown, innerbark brown; twigs short-hairy, with whitishlenticels. Leaves arranged spiralIy, panpin-nately compound with 5-11 pairs of leaflets;stipules linear, I-1.5 cm long, caducous; petioleand rachis together 3-15 cm long, groovedabove and slightly winged, short-hairy; leafletsopposite, sessile, oblong to elliptical, I-6 cm x0.5-2.5 cm, usually rounded at apex, slightlyhairy below. Inflorescence an axillary or termi-nal panicle up to 12 cm long, short-hairy;bracts sinaU. Flowers bisexual, nearly regular,whitish or slightly pinkish; pedice1 0.5-1.5 cmlong, short-hairy, Jointed near apex and persis-tent after shedding of flowers; sepals 4, ovateto oblong, 3.5-4 mm long, reflexed; petals 5,free, oblong, 4-4.5 min long; stamens 10, free,6-8 min long, slightly unequal; ovary superior,ellipsoid, c. 2 mm long, with short stipe, hairy,I-celled, style 3-4 min long. Fruit a slightlyobliquely oblong-obovate, flat pod 6.5-7 cm x3-3.5 cm, with short stipe, short-pointed atapex, smooth and brown, dehiscent with 2woody, twisting valves, I-2-seeded. Seedsrounded, dark brown. Seedling with epigealgermination; hypocoty1 2.5-3.5 cm long, epicotyl6.5-8 cm long; cotyledons oblong, thick andfleshy, wrinkled; first 2 leaves alternate, with8-10 leaflets.

Cynometro Jujue closely resembles Cynometrohonket Harms, which differs in its leaves gen-erally having more and smaller leaflets; more-over, the leaflets of Cynometro honhei are usu-ally notched at apex and glabrous.Cynometro comprises about 90 species andoccurs in the tropical regions of the world. Ap-proximateIy 25 species can be found in main-land tropical Africa and about 10 in Madagas-car. Tropical Asia and tropical America haveabout 25 species each

Ecology Cynometro Jujue occurs in evergreenlowland forest.

Management A hole harvested in southernDR Congo of 19.5 in long and 65 cm in diame-ter yielded 4 ms of wood

Genetic resources and breeding AlthoughCynometro Iujoe seems to have a rather limitedarea of distribution, there are no indicationsthat it is threatened by genetic erosion

Prospects The high natural durability andhigh resistance to abrasion of the wood offerpossibilities for timber production for specialpurposes. However, the often small size of thehole makes Cynometro Jujue less Interesting forcommercialization.

Major references Aubr6viUe, 1968; Fouarge& G6rard, 1964; Tchissambou, Benechie &Khuong-Huu, 1982; Wilczek at a1. , 1952.

Other references SImpson, 1996; Tailfer,1989.

Authors R. H. M. J. Lemmens

CYNOMETRAMADAGASCARIENSIS Baill.

Protologue Bull. Mens. SOC. Linn. Paris I:375 (1883).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Origin and geographic distribution Cyno-metro inoddgoscoriensis is endemic to easternMadagascar, where it occurs from Sambavasouth to Taolatiaro.

Uses The wood, known as 'mainpay' togetherwith the wood of several other Cynometro spp.from Madagascar, is used for posts and planksin house construction

Properties Ethanolic leaf extracts of Cyno-metro inodoguscoriensis showed potent antivi-ralactivity against herpes simplex

Botany Evergreen small to medlurn-sizedtree up to 20(-30) in tall; hole up to 50(-100)cm in diameter; twigs short-hairy, with lenti-cels. Leaves arranged spiralIy, panpinnatelycompound with (5-)6-16 pairs of leaflets; stip-ules threadlike, 0.5-I cm long, caducous; peti-o1e I-3 min long, thick, wrinkled, rachis I-10

.

Cynometro inoddgoscoriensis - wild

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cm long, grooved above, short-hairy; leafletsopposite, sessile, obliquely oblong to oblong-ovate, 0.5-2(-2.5) 0.2-0.6(-0.8) cm,notched at apex, glabrous. Inflorescence anaxillary raceme up to 1.5 cm long, short-hairy;bracts small. Flowers bisexual, nearly regular,whitish; pedice1 0.5-I cm long; sepals 4(-5), 2-3 mm long, reflexed; petals 5, free, narrowlyobovate, 3.5-5 mm long; stamens 10, free, c. 7mm long; ovary superior, with short stipe,hairy, I-celled, style 3-5 mm long. Fruit aslightly obliquely oblong-elliptic, flat pod I-2cm x 0.5-2 cm, with short stipe, warty, I-seeded. Seed with thin seed coat

Cynometro mudogoscoriensis has been subdi-vided into 3 subspecies, differing in numberand size of the leaflets and length of the leafrachis. Trees flower from August to DecemberCynometro comprises about 90 species andoccurs in the tropical regions of the world. Ap-proximateIy 25 species can be found in main-land tropical Africa and about 10 in Madagas-car. Tropical Asia and tropical America haveabout 25 species each.The wood of several other Cynometru spp. isused in Madagascar for similar purposes asthat of Cynometro ingdogoscoriensis. Cynome-tr0 o6rohomiiDU Puy & R. Rabev. is a medium-sized tree up to 30 in tall with hole up to 50 cmin diameter, widespread in deciduous woodlandand evergreen forest. Its wood Is used for postsin house construction and as roofsupportCynometro gunto R. Vig. is a small to medium-sized tree up to 25 in tall with hole up to 100cm in diameter, endemic to lowland north-eastern Madagascar. The hard wood is used forconstruction.

Cynometro coinmersoniono (DC. ) Baill. is asmalltree up to 12 in tallwith hole up to 30 cmin diameter, occurring in lowland evergreenforest in eastern Madagascar. The hard andresilient wood is used for construction and tool

handles, and as firewoodCynometro douphinensis DU Puy & R. Rabev. isa smalltree up to 15 in tall with hole up to 30cm in diameter, also occurring in lowland ever-green forest in eastern Madagascar. Its dark-coloured wood is used for poles and planks inhouse construction, for carpentry, and as fire-wood.

Cynometro soholouo DU Puy & R. Rabev. is asmallto medium-sized tree up to 25 in tallwithbole up to 40 cm in diameter, fairly widespreadin deciduous forest in northern and western

Madagascar. The hard wood is used for poles inhouse construction, for carpentry, and as fire-

cm x

wood.

Ecology Cynometro madogoscoriensis occursin evergreen coastal forest and hill forest up to600 in altitude, on sandy and latente soils

Genetic resources and breeding Cyno-metro mudogoscoriensis does not appear to beliable to genetic erosion at present because it iswidespread in eastern Madagascar. However,with the ongoing fragmentation of evergreenforest, monitoring of populations is recoin-mended.

Prospects The wood of Cynometro mudugos-curiensis and other Cynometro spp. will remainof some importance in Madagascar for localapplications, especially in house building. Lit-tle Is known about nearly all aspects, and re-search Is needed to do recommendations for

future applications of the wood and sustainableexploitation of the trees

Major references du Puy at a1. , 2002; Hud-son, Lee & Rasoanaiv0, 2000.

Other references Schatz, 2001.Authors R. H. M. J. Lemmens

CERTOGONONE 267

CYRTOGONONEARGENTEA(Pax) Pram

Protologue Bull. Misc. Inform. Kew 1911232 (1911).

Family EuphorbiaceaeOrigin and geographic distribution Cyrto-

gonone orgenteo occurs from eastern Nigeria toGabon.

Uses The wood, known as 'akikubu' in Cam-eroon, Is used for general carpentry, joineryand furniture. The seeds are eaten, often to-gether with sauce made from fruit kernels ofIruingio gobonensis (Aubry-Lecomte ex O'Rorke)

,.

Cyrtogonone urgenteo- wild

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268 TIMBERS2

Baill. The grated bark, mixed with sugar-canejuice and cooked with palm cabbage or ground-nuts, is taken against stomach-ache. It has anacrid taste and is strongly purgative. In west-ern Cameroon unspecified parts of the plantare used as anodyne.

Properties The wood is reddish white topale brown. The texture is fine, and the wood ismoderately hard and not durable. Roots of Ni-genan material have been found very effectiveas a molluscicide giving 100% mortalityagainst the fresh-water snail Bunnus globotusat an extract concentration of 100 ppm; leafextracts showed no activity. Extracts of theleaf, stem and fruit showed significant activityin a phorbol dibutyrate receptor binding assay,indicating that the extracts have tumour pro-rooting properties

Botany Dioecious, medium-sized tree up to30(-35) in tall; hole often fluted at base, up to35 cm in diameter; bark surface greyish brownto dark brown, smooth to rough with horizontallines, inner bark granular, brown, with yellowinnermost layer; twigs densely silvery scalyLeaves alternate, simple; stipules minute; peti-o1e 6-10 cm long, with 2 distinct glands at theapex; blade erriptical or obovate-oblong, some-times deeply 3-lobed, 15-35 cm x 7.5-12.5(-30)cm, base curieate to rounded, apex abruptlyacuminate, margin entire or occasionally with2 teeth or minute lobes near the apex, leathery,densely silvery scaly below, pinnately veined.Inflorescence a terminal, narrowly pyramidalpanicle up to 30 cm long, scaly, with manybranches up to 7.5 cm long, with 3-7-floweredcymes. Flowers unisexual, regular, pedicel upto 5 min long; male flowers with globose calyxin bud, up to 8 mm in diameter, at floweringirregularly (2-)3(-4)-lobed, densely scaly, pet-als 5(-6), free, orbicular, contorted, glabrous,white, stamens about 30, free, glabrous, diskglands many; female flowers with 4-5-10bedcalyx, petals as in male flowers, disk shallowlycup-shaped, ovary superior, 3-celled, with 2-ridstyles. Fruit a 3-lobed capsule up to 5 cm indiameter, brown hairy, splitting into 3 parts,each 2-valved and I'Seeded. Seeds globoseIn Nigeria trees flower in May and maturefruits can be found in July-SeptemberCyrtogonone comprises a single species andseems to be closely related to Crotonogyne andMonniophyton.

Ecology Cyrtogonone o78eitteo occurs inevergreen rainforest, often in secondary forest,up to 1000 in altitude.

Genetic resources and breeding Although

Cyrtogonone orgenteo is restricted to rainforestand has a limited area of distribution, it seemsto be quite common in most of its distributionarea, and does not seem to be threatened atpresent

Prospects Cyrtogonone orgenteo is likely toremain of local use only, unless research onthis poorly known species would reveal newopportunities.

Major references Burki11, 1994; Normand& Paquis, 1976; Radcliffe-Smith, 2001; Rapon-da-Walker & Sillans, 1961; Wilks & TSSemb6,2000.

Other references Anonymous, 2001; Beut-Ier at a1. , 2006; Brown, Hutchinson & Pram,1909-19/3; Keay, 1989; Neuwinger, 2000;Sunderland, 2005; Van der Veen & Bodingabwa Bodinga, undated; van WeIzen & Stuppy,1999; Westra & Koek. Noorman, 2004

Authors L. P. A. Oyen

DACTYLADENIASCABRIFOLIA (Hua) Prance& F. White

Protologue Brittonia 31: 486 (1979)Family ChrysobalanaceaeSynonyms Actoo SCObrifolto Hua (1897).Origin and geographic distribution Ducty-

IQdenio SCObrifolio occurs from Guinea to C6ted'TVoire.

Uses The wood is locally used for construc-tion, planks and utensils such as rice pestlesand mortars. It produces excellent charcoal. Aleaf decoction is taken to treat dysentery. Theseed Is reportedIy edible.

Properties The heartwood is pale brown toreddish brown, sometimes with narrow darker

I.

Ductylodenio SCObrifolio - wild

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streaks, and not distinctly demarcated fromthe paler sapwood. The texture is moderatelycoarse. The wood is fairly heavy, hard andmoderately durable.

Botany Small to medium-sized tree up to20(-27) in tall; bole usually straight, slightlyfluted, up to 50 cm in diameter, without but-tresses; bark surface scaly, grey to pale brown,inner bark hard, granular, pale brown to red-dish, with clear exudate; crown fairly dense,heavily branched with often drooping branches;twigs glabrous, with pale lenticels. Leaves al-ternate, simple and entire; stipules linear, c. 4min long, persistent; petiole 3-5 mm long;blade elliptical to oblong, 6-13(-15) cm x 2.5-5.5 cm, curieate at base, acuminate at apex,leathery, glabrous, upper surface rough to thetouch, with a few glands near base of lowersurface, pinnately veined with 4-8 pairs oflateral veins. Inflorescence a terminal or axil-

Iary raceme 6-10 cm long, slightly hairy. Flow-ers bisexual, zygomorphic, 5-merous, greenishwhite to pinkish; pedice1 6-14 mm long; recep-tacle tube narrowly obconica1, 4-8 min long,glabrous; sepals 4-6 mm long; petals 4-6 mmlong, caducous; stamens c. 20, fused, 1.5-2.5cm long; ovary superior, inserted at mouth ofreceptacle tube, I-celled, style slightly longerthan stamens, slightly 3-lobed at apex. Fruitan ovoid drupe 4-5 cm X 2-3 cm, smooth andglabrous, I-seeded. Seedling with epigeal ger-mination.

In Sierra Leone flowering trees of Doctyludenioscabrifo!to have been recorded throughout theyear; fruits are most commonly found in An-gust-SeptemberDocty!odenio comprises about 30 species and Isconfined to West and Central Africa. Doc-

tylodenio SCObrijolio is closely related to Doc-tylodenio burteri(Hook. f. ex 011v. ) Prance &F. White, which has some reputation as a fallowcrop in Nigeria, but the latter species has ahairy receptacle tube and more smooth leaves

Ecology Ducty!odenio SCObrtfolio is usuallyan understorey tree of primary and old second-ary rainforest, but also occurs in rrverine forestand in abandoned farmland. It is found up to1100 in altitude in Mount Nimba. It is locallyabundant

Management Doctylodenio SCObrifolio hasbeen considered a weed in managed forest, andtrees have been poisoned

Genetic resources and breeding Doctylo-dento SCObrifolio is locally abundant in differ-ent types of forest and does not seem to be un-der threat of genetic erosion. However, many

other Doctylodenio spp. seem to be uncommon

Prospects Trees of Docty!odenio SCObrifolioare usually too small to be of Importance fortheir timber. Very little is known about thisspecies, but research could be worthwhile inthe light of the good prospects of the relatedDoctylodenio barteri in agroforestry systemsand in fuelwood plantations managed by cop-PICing

Major references Burki11, 1985; Ladipo &Kang, 2002; Prance & Sothers, 2003b; Savill &Fox, 1967

Other references Aubr6ville, 1959b; Haw-thorne & Jongkind, 2006; ITvine, 1961; Kryn &Fobes, 1959

Authors R. H. M. J. Lemmens

or rare.

DANIELLIA 269

DANIELLIAKLAINEIPierre exA. Chev.

Protologue Bois on Gabon: 172 (1917).Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Origin and geographic distribution Doniel-

jig h!Qinei occurs in Cameroon, EquatorialGuinea, Gabon, DR Congo and Cabinda (ingo-Ia).

Uses The wood, traded as 'faro' togetherwith the wood of other Doniellio spp. , Is suita-ble for light construction, light flooring, joinery,interior trim, canoes, furniture, toys, novelties,boxes, crates, agricultural implements, vats,draining boards, turnery, veneer, plywood,hardboard and particle board. Cylindrical box-co are made from the bark of young trees. Theresin from the wood yields gum copal. In Gabonthe gum is externalIy applied to treat scabies,

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Doniellio kidinet - coild

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270 TIMBERS2

ringworm and lice on the head.Production and international trade The

timber of Donielliu hloineiis only used on alocal scale.

Properties The heartwood is pinkish grey topale yellow-brown, with irregular green-brownmarkings, and fairly well demarcated from thewide, whitish or pale yellow sapwood. Thegrain is slightly interlocked, texture moderate-Iy coarse. Quarter-sawn surfaces show a nicefigure. The wood is scented and has a lustroussurface

The wood is lightweight to medium-weight,with a density of about 510 kg/ms at 12% ino1s-ture content, moderately soft but tough andstrong. It usually air dries well without de-grade. In thick boards there might be slightdistortion and collapse, but the defects are neg-Iigible. Once dry, the wood is stable in serviceAt 12% moisture content, the modulus of rup-ture Is 65-101 N/mm2, compression parallel tograin 31-42 N/mm2, cleavage 8.5-13 N/mmand Chalais-Meudon side hardness 1.9-2.8.The wood saws easily and is easy to work withboth hand and machine tools. It planes to anice finish, but needs careful filling before-hand. It nails and glues well, but is rather dif-ficult to split with tools. It joints well and canbe readily peeled into veneer. The steam bend-Ing properties are very poor.The wood Is not durable, being susceptible toattacks by fungi, termites, pinhole borers, Lye-tus and marine borers. The heartwood is inod-

erately resistant to impregnation, but the sap-wood is easier to treat with preservatives.The wood contains 47% cellulose, 25.5% 11gnin,21% pentosan, 1.0% ash and 0.02% silica. Thesolubility is 4.9% in alcohol-benzene, 3.5% inhot water and 25.6% in a I% NaOH solution

The heartwood contains a dark brown, oily,sticky, resinous gum which can be saponified,although it has a low saponification value. Oneof the main compounds of the gum is the diter-pene o1eoresin daniellic acid (illurinic acid), aswellas ozic acid and the alcoholozolThe essential oil distilled from the roots con-tamed as major component myrcene (55%), andas minor components orbumulene (8.1%), ger-macrene-D (6.1%), u-pinene (5.4%), finalo01(4.9%)and ^-, aryophyllene (3.6%). Th. oil sh, wedantioxidant and radical scavenging activities,as well as inhibition of lipid peroxidation. Italso showed in-vitro antibacterial activity.

Botany Deciduous, medium-sized tree up to30 in tall; hole straight and cylindrical, up to100 cm in diameter, without buttresses; bark

surface smooth, greenish grey, with lenticels inhorizontals rows, becoming scaly, inner barkthick, hard, fibrous, pinkish brown with finelines; crown flattened; twigs glabrous. Leavesalternate, paripinnately compound with 4-7pairs of leaflets; stipules up to 8 cm x I cm,soon falling; petiole (2-)2.5-4(-9) cm long, ra-chis 11-20(-30) cm long, shallowly channelledat base, with a pair of glands at insertion ofbasal pair of leaflets; petiolules 4.5-8 mm long;leaflets opposite, oblong-lanceolate to lanceo-late, up to 22(-38) cm x 7(-9) cm, basal andapicalleaflets smaller than middle ones, baserounded, asymmetrical, apex acuminate, mar-gins slightly wavy, papery to leathery, gla-brous, with many translucent glandular dots,pinnately veined with 13-29 pairs of lateralveins. Inflorescence an axillary or terminalcompound raceme 20-31 cm long, glabrous toshort-hairy, with 7-9 lateral branches. Flowersbisexual, zygomorphic; pedice1 (I-)1.5-2(-2.5)mm long, glabrous, enlarging in fruit, near themiddle with 2 caducous bracteoles c. 1.5 cm

long; sepals 4, oblong-ovate, (I-)1.5-2 cm long,glabrous but margins hairy, with few glanddots on external surface; petals 5, violet, une-qual, oblong-ovate to ovate-elliptical, 3 petalsI-1.5 cm long and 2 up to 0.5 cm long, glabrousto short-hairy; stamens 10, 3-4 cm long, 9united at base and I free; ovary superior, Ian-ceolate, 0.5-I cm long, glabrous but coveredwith gland dots, with stipe up to 0.5(-I) cmlong, style c. 3 cm long. Fruit an obliquely ob-long, flattened pod (8-)10-12 cm x 4-5 cm,with stipe of c. I cm long, glabrous, green be-coming brown, dehiscing with 2 woody valves,I-seeded. Seed oblong, flattened, 4-5 cm long,smooth, deep red to brown, attached to one ofthe valves by a c. I cm long funicle. Seedlingwith epigealgermination.Doniellio hintneiflowers from August to De-cember and fruits from October to February.Dontellio comprises 10 species, which are near-Iy all confined to the forest areas of West andCentral Africa, with only ISPecies extending toSudan and Uganda. The wood of some otherDoniellio spp. is used for similar purposes asthat of Doniel!to hintnei.

Donielli0 o18teeniono PADuvign. is a medi-urn-sized tree up to 25 in tall with hole up to150 cm in diameter, occurring in Gabon, DRCongo, Zambia and Angola, mainly in savannaIn Zambia the wood is used in house buildingand for canoes, veneer and plywood. It is alsoused as firewood and for charcoal making. Thewood yields gum copal. The inner bark from

saplings is made into string.Doniellto oblong0 011v. is poorly known, occur-ring in Cameroon, Equatorial Guinea and Ga-bon, possibly also in Benin and south-easternNigeria. It is a large tree up to 40 in tall withbole up to 150 cm in diameter, and is closelyrelated to Doniellio ogeo (Harms) Rolfe ex Hol-land and often confused with it. The wood isused for boats, canoes and railway sleepers,and is also suitable for boxes, crates and ve-neer. The wood yields gum copal. Donielliooblongo is threatened by habitat loss and ex-PIOitation, and has been classified as vulnera-ble in the IUCN Red List of threatened speciesDonie!!to py, toertii De Wild. is a large tree upto 40 in tallwith hole up to 240 cm in diameter,occurring in the Central African Republic, Ga-bon, Congo and DR Congo. The wood is proba-bly used for similar purposes as other Donie!Jigspp. In DR Congo the inner bark Is eaten bymen to increase virility. The wood yields gumcopal.Doniellio soyauxii(Harms) Rolfe is a fairly largetree up to 35 in tall with hole up to 120 cm indiameter, occurring in Gamero0n, EquatorialGuinea, Gabon, DR Congo and Cabinda (Ango-Ia). Its wood has similar uses as that of otherDoniellio spp

Ecology Doniellio hidinet occurs in evergreen,primary as well as secondary forest, also ingallery forest and swampy localities, from sea-levelup to 250(-800) in altitude.

Management The minimum bole diameterfor harvesting is 60 cm in Garnero0n and 70 cmin Gabon. Logs should be processed quicklyafter felling or treated with preservatives be-cause the wood is susceptible to discolorationby blue-stain fungiand to insect attacks

Genetic resources and breeding Donie!-Ito hintnei has a relatively large area of distri-bution, but has been classified as lowerrisk/near threatened in the IUCN Red List ofthreatened species as it is under pressure byhabitatloss and exploitation

Prospects International demand for sawnwood of Doniellio hloinei may remain limitedbecause of the poor durability of the wood, butthe straight and cylindrical holes are suitablefor rotary peeling for veneer and plywood pro-duction.

Major references Bolza & Keating, 1972;CTRAD Forestry Department, 2008; de in BS-trella at a1. , 2010; Koudou at a1. , 2008; Taka-hashi, 1978.

Other references Burki11, 1995; Chilufya &Tengnas, 1996; G6rard at a1. , 1998; Ngavoura,

1990; Onanga at a1. , 1999; Raponda-Walker &Sillans, 1961; Savard, Besson & Morize, 1954;Vivien & Faure, 1985; Wilczek at a1. , 1952.

Authors G. H. Schmelzer

DANIELLiAOGEA (Harms) Rolfe ex Holland

Protologue Bull. Misc. Inform. Kew, additser. 9: 268 (1911)

Family Caesalpiniaceae (I. eguminosae - Cae-salpinioideae)

Vernacular names ACcra copal, Benin gumcopal, gum copaltree (En). Arbre a encens, farod'Agboville (Fr). Incenso (Po).

Origin and geographic distribution Doniel-lid ogeo occurs in West and Central Africa,from Senegal east to Cameroon and south toGabon.

Uses The wood, traded as 'faro', 'ogea' or'daniellia', is suitable for light construction,light flooring, joinery, furniture, toys, novelties,boxes, crates, agricultural implements, vats,draining boards, veneer, plywood, hardboardand particle board. It can be used for papermaking, and is suitable for firewood and char-coalproductionThe quantity of gum, known as 'aguJa gum or'gum copal' in the wood is limited and coin-mercial tapping has not been successful. Thegum is used in cosmetics, to scent clothes andfumigate huts, and also for medico-magicaltreatments. It is used to repair broken potteryby placing it along the broken edges and set-ting fire to it to make the reparation strongerIt is also used as varnish. The gum, mixed withsoot and oil, is burnt and is used for tattooing.In Senegal the gum is applied to skin coin-

DAIJIELLIA 271

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Doniellio ogeo- wild

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272 TIMBERS2

plaints. In Ghana the gum, pounded togetherwith a seed ofDiocleo reflexo Hook. f. and somekaolin, is taken in water to treat cough andasthma. The gum is also applied to Buruli ul-cers, caused by Mycobocterium ulcerons. InBenin leaf infusions are taken to treat dysen-tery. In Ghana and Nigeria root decoctions ormacerations are taken to treat malaria and

gonorrhoea. The gum is chewed as a purgativeand together with other ingredients applied tosnakebites. In Nigeria the seeds are considerededible

Production and international trade In 1983

C6te d'Ivoire exported 73,000 ina of logs of faro(Doniel!to spp. ). Currently Donie!jig ogeo andother Doniellio spp. have no importance on theinternational timber market, but their wood iscommonly used locallyIn 1876 one of the first exports of 500 t gumcopalwas made from Ghana to Europe. Lumpsof gum copal are still sold in local markets,although they are often contaminated with soil

Properties The heartwood is pale pinkishbrown to reddish brown with dark streaks, andIs distinctly demarcated from the up to 15 cmwide, whitish or pale yellow resinous sapwood.The grain is interlocked, texture mediumcoarse. The wood Is scented and has a lustroussurface.

The wood is lightweight to medium-weight,with a density of about 515 kg/ina at 12% roots-ture content, moderately soft but tough and ofmedium strength. The shrinkage rates aremoderate, from green to oven dry about 2.6%radial and 5.6-8.2% tangential. Kiln drying isfairly fast with little degrade. In thick boards,there might be slight distortion and collapse,butthe defects are not severe. At 12% moisturecontent, the modulus of rupture is 77-84N/min2, modulus of elasticity 8820-9700N/mm2, compression parallel to grain 40-47N/min2, shear 7-11 N/mm2, Janka side hard-ness 3160 N and Janka end hardness 4310 N.The wood is easy to work with both hand andmachine tools, with little blunting effect oncutting edges. Although the wood planessmoothly, tearing of quarter-sawn surfacesmay occur due to interlocked grain, and thin,sharp cutting edges are more useful in obtain-ing a good finish than a reduced cutting angleThe wood stains satisfactorily but needs a fillerbefore polishing. It nails well and glues correct-Iy. It can be readily peeled into veneer. Thesteam bending properties are very poor. Thewood is not durable, being susceptible to at-tacks by fungi, borers, termites, pinhole borers,

Lyetus, longhorn beetles and marine borersThe heartwood and inner sapwood are moder-ately resistant to Impregnation, but the outersapwood is easy to treat with preservativesThe cellulose content of the wood is 46-48%.The wood fibres are strong and about 1.5 minlongThe gum from the heartwood is dark brown,oily and sticky, and can be saponified, althoughit has a low saponification value. One of themain compounds of the exudate is the diter-pene o1eoresin daniellic acid (illurinic acid), aswell as ozoic acid and the alcohol ozol. Chemi-cal analyses revealed that the gum contained6.0% water, 1.3% protein, 13.0% fat, 42.8%crude fibre and 2.0% ash. It had 8.0% foamingcapacity, and oil and water absorption capacitywere 12.0% and 24.3%, respectivelyThe seed contains low levels of protein (10.1%)and fat, but high levels of total carbohydrate(74%) and minerals. It contains much of the

amino acid cysteine. The seed oilcontains highamounts of linoleic acid (27.7%), followed by11gnoceric acid (9.2%), palmitic acid (7.5%),oleic acid (6.6%) and behenic acid (3.3%).A range of sesquiterpenoids were isolated fromthe leaf oil, with as main component caryo-phyllene oxide (20.1%), followed by burnuleneoxide (7.0%), u-burnulene (3.8%) and ^-sennene(3.8%)

Adulterations and substitutes The woodof Doniellio ogeo is similar to that of Donielliothurifero Benn. and Doniel!to hloinei Pierre exAchev. and has similar uses.

Description Deciduous, medium-sized tolarge tree up to 40(-50) in tall; hole straightand cylindrical, up to 125 cm in diameter,without buttresses; bark surface smooth, grey-ish white to greenish grey, with elongated hor-Izontal patches in various colours, inner barkthick, hard, fibrous, pinkish brown with finelines; crown rather small, flattened. Leavesalternate, panpinnately compound with 5-9pairs of leaflets; stipules up to 8.5 cm long,soon falling; petiole 2-4(-4.5) cm long, rachis(IIJ15-24(-27) cm long, shallowly channelledat base, quadrangular at apex, with a pair ofglands at insertion of basal pair of leaflets;petiolules 3-9 min long; leaflets opposite, ellip-tical-lanceolate to lanceolate, up to Ile15) cmx 4 cm, basal and apicalleaflets smaller thanmiddle ones, base curieate to rounded, asym-metrical, apex acuminate, margins entire, pa-pery to leathery, glabrous, with many translu-cent glandular dots, pinnately veined with 8-16 pairs of lateral veins.

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glabrous; cotyledons fleshy, c. 2.5 cm long,rounded; first leaves alternate, with I-2 pairsof leaflets.

Other botanical information Doniellio

comprises 10 species, which are nearly all con-fined to the forest areas of West and Central

Africa, with only I species extending to SudanandUganda

Anatomy Wood-anatomical description (IAWAhardwood 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; (26: intervessel pits medium (7-10pin)); 27: inter, essel pits large (^ 10 ILm); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 43: mean tan-gential diameter of vessellumina z 200 ILm; 46:S 5 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;(68: fibres very thin-walled); 69: fibres thin- tothick-walled. Axial parenchyma: 78: axial pa-renchyma scanty paratrachea1; 79: axial paren-chyma vastcentric; (89: axial parenchyma inmarginal or in seemingly marginal bands); 91two cells per parenchyma strand; (92: four (3-4) cells per parenchyma strand). Rays: 97: raywidth I-3 cells; 98: larger rays commonly 4- to10-senate; 106: body ray cells procumbent withone row of upright and/or square marginalcells; (107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells); 1/5: 4-12 rays per min. Stonedstructure: 1/8: all rays stoned; 120: axial pa-renchyma and/or vessel elements stoned; 121:fibres stoned. Secretory elements and cambialvariants: 129: axial canals diffuse. Mineral

inclusions: 136: prismatic crystals present; 142:prismatic crystals in chambered axial paren-chyma cells(L. Awoyemi, E. Ebanyenle, P. E. Gasson &E. A. Wheeler)

Growth and development Doniellio ogeois considered a pioneer, and seedlings of morethan 50 cm tall are light demanding and foundin forest gaps. The tree usually sheds its leavesat the end of the rainy season, and floweringoccurs when the tree is leaness. Doniellio ogeohas been found flowering from September toMay and fruiting from November to March.

Ecology Doniellio ogeo occurs in evergreenand semi-deciduous forest, also in swampy

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Doniellio ogeo - I, legj 2, flowers und fruits, 3,flower with one sepol und one petol reinoued, . 4,fruit uo!ue with seedRedro"it und udopted by 111 We88e!-Brond

Inflorescence an axillary or terminal compoundraceme (7-)15-20(-22) cm long, densely woollyhairy, with 5-12 lateral branches. Flowersbisexual, zygomorphic; pedice1 9-15(-17) minlong, glabrous, enlarging in fruit, near themiddle with 2 caducous bracteoles 0.5-I cm

long; sepals 4, oblong, I-1.5(-2) cm long, short-hairy, with gland dots on externalsurface; pet-als 5, blue to lilac, unequal, oblong to oblong-elliptical or oblong-lanceolate, 3 petals 0.5-I(-1.5) cm long and 2 up to 0.2 cm long, glabrousto short-hairy; stamens 10, 2-3(-3.5) cm long, 9united at base and I free; ovary superior, ob-long to rounded, c. 0.5 cm long, densely woollyhairy, with stipe 0.5-I cm long, style 1.5-2(-3)cm long. Fruit an obliquely oblong, flattenedpod 6-7(-9) cm x 2.5-3.5(-5) cm, with stipe0.5-I cm long, densely short-hairy mainly atmargin, green becoming brown, dehiscing with2 woody valves, I'Seeded. Seed oblong, flat-tened, c. 3 cm long, smooth, dark brown, at-tached to one of the valves by a c. 1.5 cm longfunicle. Seedling with epigeal germination;hypocotyl c. 9 cm long, epicotyl c. 6 cm long,

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DANIELLIA 273

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274 TIMBERS2

localities near water courses, from sea-level upto 800 in altitude. In Nigeria it occurs in oldtree plantations after felling of the plantationtrees

Management In forest management, thereis generally no focus on Donie!jig ogeo becauseit is not a preferred timber tree. In Sierra Leo-ne the average density of trees with a bole di-ameter of more than 60 cm has been estimated

to be 0.06 per haHarvesting The minimum bole diameter

allowed for harvesting is 60 cm in C6te d'TVoireand Cameroon, 70 cm in Liberia and 90 cm inGhana

Yield In Sierra Leone it has been reportedthat trees with a hole diameter of 90-100 cm

yield 6-9 ina of usable wood.Handling after harvest Cross-breaks may

occur near the heart of the hole. Logs should beprocessed quickly after felling or treated withpreservatives because the wood is susceptibleto discoloration by blue-stain fungi and attacksby insects.

Genetic resources Doniellioogeohas a largearea of distribution, but occurs sparsely. How-ever, it does not seem to be threatened by ge-netic erosion as its wood is generaUy mainlylocally used and not subject to commercial ex-PIOitation. Seeds are stored at the 11TA Genebank

Prospects Donie!lid ogeo has not been con-sidered a timber tree of commercial importancebecause its wood is susceptible to fungal at-tacks. However, the wood may have good pro-spects for the production of veneer and plywoodby rotary peeling. More information is neededon growth rates and methods to improve re-generation to be able to draw conclusions aboutpossibilities for sustainable exploitation fromnatural forest. Further research of the exudate

is needed on the possible usage in the varnishand cosmetic industry.

Major references Asekun & Ekundayo,2004; Bolza & Keating, 1972; Burki11, 1995; dein BStrella at a1. , 2010; Ezeagu at a1. , 1998;Ezeagu at a1. , 2002; Neuwinger, 2000; 0teng-Amoako (Editor), 2006; Takahashi, 1978

Other references Abbiw, 1990; NIOnkhai,1988; Ajibesin at a1. , 2008; Allahi at a1. , 2011;Bevan, Ekong & Okogun, 1968; CTFT, 1955a;de in EstreUa, Aedo & Velayos, 2009; Ezeagu,2005; Ezeagu at a1. , 1996; G6rard at a1. , 1998;Hawthorne, 1995; Hawthorne & Jongkind,2006; Keay, 1989; 01adip0, 2005; Petzke at al. ,1997; Sandermann, Ehlers & Feht, 1963; Savill& Fox, 1967

Sources of illustration Voorhoeve, 1965Authors G. H. Schmelzer

DANIELLIAOLIVERl(Rolfe) Hutch. & Dalziel

Protologue Bull. Misc. Inform. Kew 9: 382(1928).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Chromosome number2n=22

Vernacular names West African copaltree,African copaiba balsam, norm balsam, ACcracopal, Benin gum copal(En). Arbre a vernis(Fr). Pau-incenso (Po).

Origin and geographic distribution Doniel-Jig onDen occurs from Senegal east to SudanandUganda.

Uses The wood, traded as'WestAfrican gumcopal' and 'daniellia', is used for light flooring,joinery, interior trim, furniture, boat building,toys, novelties, cattle-troughs, drums, bowls,rice-mortars, packing cases, draining boards,carvings, veneer, plywood, hardboard and par-ticle board. However, the gum makes it rathertoo gummy for high-quality joinery and carv-Ings. It is popular as firewood, as it has a fra-grant smoke and is slow-burning. It is alsomade into charcoal. The wood has been used

for paper making. The wood ash is sometimesused in soap productionLarge strips of bark are made into beehives.The resin or gum, traded as 'gum copal' and'African copaiba balsam', is fragrant and isused as torch or as incense in religious ceremo-nies and to fumigate houses and clothes tochase insects. The gum is furthermore used tofix spearheads to the shaft. The powdered gum

I.

Duniellio onueri- wild

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is applied to cloth the make it shiny, and it isapplied to furniture as a polish or varnish, tomake it shiny and more resistant to termites.It is not suitable for commercial furniture

polish as it is not soluble in turpentine or alco-hol. However, it can be esterified to produce alacquer. In Nigeria drums are soaked in a solu-tion of the gum, a treatment which deepens thetone

The gum and stem bark, and to a lesser extentthe roots and leaves, are widely used in tradi-tional medicine. The gum, bark and leaves areburnt and the smoke inhaled to treat headache

and migraine; the smoke is also used as inOS-quito repellent. The gum is chewed and swal-lowed as a purgative to treat diarrhoea. In C6ted'Ivoire the gum is considered aphrodisiac anddiuretic, and it is chewed to treat cough, head-ache, tachycardia and painful menstruation.The gum is externalIy applied to treat itchingskin and skin diseases. The gum and bark aretaken in various preparations, internally andexternalIy, and sometimes with other plantparts, to treat venereal diseases, ulcers andsores, circumcislon wounds, leprosy, dysentery,colic, menstrual problems, cough, colds, angina,bronchitis, tuberculosis, kidney problems, ap-pendicitis, headache, back-ache, rheumatism,fever pains, hernia, tooth-ache and snakebitesThe root is considered diuretic and a decoction

is taken to treat venereal diseases, absence ofmenses, anxiety, insanity, food poisoning andskin diseases. Leafy twigs are put in baths andused as vapour baths to treat fever and jaun-dice, and also as a tonic. A decoction of theleafy twigs with salt is taken as a purgative, totreat constipation and stomach-ache. A leafbud infusion is taken to treat migraine andfeverish pain in teething children. Poundedyoung leaves are applied to wounds, burns andabscesses to ripen them; they are also appliedto painful parts of the body. A leaf decoction isgargled to treat toothache and drunk to treatcolic. The large stipules are used to cover sores,ulcers and injuries, and are also used as tam-pon for menstruating women. In Burkina Fasosheep and goats are given a stem bark decoc-tion to treat intestinal worms. In Nigeria thegum is applied to the skin of horses to treatmange

The young leaves are eaten with condiments orcooked in soup, as famine food. The seeds andfruits are eaten occasionally as well. Cattlebrowse the leaves readily, and the leaves areused as fodder. From the roots excellent chew-

sticks are made. The gum is sometimes chewed

for its sweetness, or used as a filling in dentalpractices. It is also made Into false amberbeads. The bark is added to accelerate fermen-

tation in brewing beer. Grilled branches areadded to palm-wine to improve digestIbilityThe burnt bark is used to blacken the interior

of pots. The tree is used in reforestation pro-grammes, and is also planted as a windbreakand avenue tree. The leaves are used for

mulching. The flowers are much visited byhoney bees for nectar. In Ghana Doniellio on-Dents considered an indicator of undergroundwater. Children wear a piece of gum aroundtheir neck as a charm against bad luck.

Production and international trade Doniel-

Ito OILueriis not important on the internationaltimber market, but it is commonly used locallythroughout its distribution area, similar toother Doniellio spp.

Properties The heartwood is red-brown, greyor red with dark streaks and is moderatelyclearly demarcated from the up to 8 cm wide,whitish sapwood with a pink or brown tingeThe grain is interlocked, texture medium tocoarse. The wood is lustrous and contains a

yellow or brown o1eoresinThe wood is lightweight to medium-weight,with a density of 510-680 kg/ina at 12% inois-ture content, moderately soft but tough andstrong. The shrinkage rates of the wood aremoderate, from green to oven dry are 2.8-3.5(-5.6)% radial and 3.7-7.0(-9.3)% tangential. Airdrying is fairly fast with little degrade. Oncedry, the wood is moderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 56-114 N/mm2, modulus of elasticity6280-9700 N/mm2, compression parallel tograin 25-63 N/min2, shear 5-15 N/min2, cleav-

12-28 N/mm and Chalais-Meudon sideage

hardness 1.2-3.4(-5.8).The wood is easy to saw and work with bothhand and machine tools, with little bluntingeffect on saw teeth and cutting edges. Thewood stains satisfactorily but needs a fillerbefore polishing. In general it nails and screwswell and glues correctly. It can be readilypeeled into veneer, but the veneer may warpduring drying. The wood is not durable, beingsusceptible to attacks by fungi, termites, pin-hole borers, Lyetus and marine borers. Theheartwood is resistant to impregnation withpreservatives and the sapwood has a variablepermeabilityThe wood contains 40.5-42% cellulose, 27.5-28% 11gnin, 18-19% pentosan and 0.8-1.0% ash.The solubility is 3.6-7.3% in alcohol-benzene,

DANIELLIA 275

276 TIMBERS2

2.3-4.2% in hot water and 19.4-20.2% in a I%

NaOH solution

The gum from the heartwood is yellow to darkbrown, oily and sticky. It is rich in essentialoils (30-35%), and one of the main compoundsis the diterpene o1eoresin daniellic acid (illur-inIC acid), as well as ozic acid and the alcoholozol. The essential oils from the leaves contain

mainly sesquiterpenoids, with as major coin-pounds 6-cadinene (24.2-31.1%) and or-CODaene(7.0-8.3%). The flavonoid glycosides rutin,quercitrin and quercimeritrin were isolatedfrom the leaves. The dried leaves contain c.

5.5% protein. The essential oil from the barkyielded only sesquiterpenoids, with as majorcompounds orcopaene (6.0-12.0%), germacreneD (4.5-79.5%) and 6-cadinene (25.5-29.8%)Tannins, saponins, and cyanogenetic and car-diac glycosides were also isolated from thebark. The seeds have a very high protein con-tent of 33.4% and a moderately high carbohy-drate content of 44.6%. The most abundant

mineral in the seeds is Ca (263 ing/100 g), theleast abundant minerals are Zn, Cu and Mn.Several compounds considered anti-as

nutritional were also found to be present in lowamounts, including cyanide, polyphenols, phyt-ate and oxalate

Hexane and methanol extracts of the bark were

tested for analgesic, antipyretic and anti-inflammatory activities. Hexane extracts ex-bibited a dose-related analgesic activity inmice, and methanol extracts inflammatoryproperties in rats. Metabolic cage studiesshowed that the ethanol extract significantlydecreased body weight, food intake, urine andstool output of rats. This extract also exhibited

histamine-a competitive antagonism on

induced contraction of the guinea pig ileumand a non-competitive inhibition of acetylcho-line-induced contraction of the frog rectus ab-dominis muscle. in aqueous extract of the barkshowed significant and dose-dependent anti-nociceptive activities when intraperitoneallyinjected in rats and mice. The extract showed anon-dose dependent anti-lrillammatory activi-ty. The effect was significant at doses of 100and 200 ing/kg. The L050 of a bark decoction insheep was found to be 3500 ing/kg. The cardiacglycosides in the methanolbark extract showedto be non-competitive antagonists for musca-rinic receptors. The methanolic leaf and barkextracts were found to possess neuromuscularblocking properties in ratsDifferent water and ethanolic extracts of

leaves, bark and roots showed moderate to

significant antibacterial activity in vitroagainst a range of pathogenic bacteria. An eth-anonc leaf extract also showed significant anti-fungal activity against Tricophyton rubrum. Acrude bark extract showed significant anddose-dependent arithelmintic activity againstthe gastro-intestinal parasite Hoemonchus con-tortus in vitro. A pot experiment showed thatmulching with Donielli0 o11uerileaves showedan inhibitory effect on the germination of soy-bean, cowpea, maize, sorghum and millet

Adulterations and substitutes The wood

of other West African Donie!Ito spp. is used forsimilar purposes as the wood of Donie!lid on-uerj

Description Deciduous, medium-sized treeup to 25(-35) in tall; bole straight and cylindri-cal, up to 200 cm in diameter, without but-tresses; bark surface smooth, greyish white,becoming scaly in older trees, flaking off inlarge, circular patches, inner bark thick, deepred; crown dense, Inversely cone-shaped; twigsglabrous. Leaves alternate, panpinnately coin-pound with (3-)6-11 pairs of leaflets; stipulesup to 6.5(-9) cm x 0.5 cm, soon falling; petiole(2-)2.5-3.5(-5.5) cm long, rachis (17-)24-33(-

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52) cm long, shallowly channelled, becomingquadrangular towards apex, sparsely short-hairy, with glands at insertion of leaflets; pe-tiolules 3-15 mm long; leaflets opposite, ob-long-ovate to lanceolate, up to 15(-21) cm x 7(-10) cm, basal and apicalleaflets smaller thanmiddle ones, base curieate, asymmetrical, apexacuminate, papery to leathery, margins slight-Iy wavy, sparsely to densely short-hairy tonearly glabrous, with few translucent glandu-Iar dots, pinnately veined with 9-17 pairs oflateral veins. Inflorescence an axillary or ter-minal compound raceme 15-20(-25) cm long,glabrous to densily hairy, with 6-16 lateralbranches. Flowers bisexual, zygomorphic,scented; pedice1 7-13 min long, glabrous, en-Iarging in fruit, below the middle with 2 cadu-cous bracteoles c. 0.5 cm long; sepals 4, oblong,c. 1.5 cm long, glabrous but margins hairy;petals 5, white to creamy, unequal, ovate-elliptical, 4 petals I-1.5(-3) mm long and one cI cm long, nearly glabrous, with few glandulardots; stamens 10, free, (2-)3-4 cm long; ovarysuperior, oblong-lanceolate, 0.5-I cm long, gla-brous, with stipe c. 0.5 cm long, style up to 3cm long. Fruit an obliquely lanceolate, flat-tened pod 6-10 cm x 3-4.5 cm, with stipe c. Icm long, glabrous, green becoming brown, de-hiscmg with 2 papery valves, I'Seeded. Seedsobovoid-ellipsoid, flattened, 2-2.5 cm long,smooth, dark brown, attached to one of thevalves by a I-2 cm long funicle. Seedling withepigealgermination.

Other botanical information Doniellio

comprises 10 species, which are nearly all con-fined to the forest areas of West and CentralAfrica, with only Doniellto onDen extending toSudan andUganda

Anatomy Wood-anatointcal 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; 26: intervessel pits medium (7-10 pin);27: intervessel pits large ^ 10 pin); 29: ves-tured pits; 30: vessel-ray pits with distinct bor-ders; similar to lritervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 pin;43: mean tangential diameter of vessellumina^ 200 prn; 46: s 5 vessels per square minimetre;47: 5-20 vessels per square minimetre; 58gums and other deposits in heartwood vesselsTracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibres

present; 69: fibres thin. to thick-walled. Axialparenchyma: (79: axial parenchyma vasicen-tric); 80: axial parenchyma allform; 81: axialparenchyma lozenge-allform; 83: axial paren-chyma confluent; 89: axial parenchyma inmarginal or in seemingly marginal bands; 91two cells per parenchyma strand; 92: four (3-4)cells per parenchyma strand. Rays: 97: raywidth I-3 cells; 98: larger rays commonly 4- to10-senate; 106: body ray cells procumbent withone row of upright and/or square marginalcells; (107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells); 1/5: 4-12 rays per mm. Stonedstructure: 1/8: all rays stoned; 120: axial pa-Tenchyma and/or vessel elements stoned; (121fibres stoned). Secretory elements and cambialvariants: 127: axial canals in long tangentiallines; 128: axial canals in short tangentiallines.

(L. Awoyemi, P. E. Gasson & E. A. Wheeler)Growth and development Donielliu o1iueri

is considered a relatively fast-growing tree. InMallseedlings reached 18 cm tall 10-15 weeksafter sowing, and in C6te d'TVoire and BurkinaFaso 45 cm and 32 cm, respectively, 32 monthsafter planting. Saplings had an average heightof 90 cm in C6te d'Ivoire when 4.5 years oldand 130 cm after 5.5 years, with a survival rateof 57%. In Maliroot suckers reached 2.4 in tall

after 5.5 years. In C6te d'Ivoire and Benin av-erage increase in hole diameter is 6 min/year,but in Burkina Faso growth rings up to 20 mmwide have been observed. Donie!lid onDen

flowers during the first half of the dry seasonfrom October to March, usually when trees areleaness or are developing new leaves, andfruits from January to June

Ecology Doniellio onDen occurs in tree sa-vanna, bush savanna and in more open grass-land, on any type of soil, but often on sandysoils, from sea-level up to 1500 in altitude. It isoften one of the dominant trees in more humidsavanna and may locally constitute nearly purestands, e. g. in temporarily flooded localities inthe Sahel region. It is sometimes found on ter-mite mounds. It is resistant to fires ifthese are

not too frequent and not too late in the dry

Propagation and planting Donie!lid on-ueriis a light-demanding species. Natural re-generation is often well. There are about 600seeds per kg. The seed is liable to insect attackand ash should be added during storage to pre-vent damage. It should be soaked in water for72 hours before sowing. The germination rate

DANIELLIA 277

season

278 TIMBERS2

is 75-95% in 2-3 weeks. Growth in the nurseryIs slow and irregular. Transplanting of seed-lings is not very successful; tap roots areformed rapidly, and frequent pruning of theroots is essential for successful planting. InC6te d'Ivoire and Burkina Faso the best sur-vival rate 3 years after planting was 60%, butin most cases it was less than 20%. Seeds mayalso be sown directly into the field. Donie!JigonDer" can also be propagated by root suckers.

Management Weeding after transplantingIs needed because of the slow growth of seed-lings. As Donie!Ito onueri produces many rootsuckers, many small trees are often found to-gether. It can be coppiced. In Ghana it is beingplanted experimentalIy to provide fodder forcattle. It can be pruned regularly for fodder

Diseases and pests Dunielli0 o1iuert is analternative host of leaf spot of shea butter tree(Vitellorio porodoxo CF. Gaertn. ), caused byPesto!otio spp. Leaves are frequently eaten byelephants and baboons

Harvesting The resin or gum is extractedby removing a part of the bark or by cutting ahole to the heart of the hole, after which theresin starts flowing. When fire is made in thehole, the process is accelerated, but this prac-tice can kill smalltrees

Yield In natural woodland in C6te d'Ivoire,trees with a mean hole diameter of 80 cm had

an average yield of 2.25 ms of log wood and 3.4ina of firewood. The firewood yield and the prof-itability of a traditional Donie!jig onDen short-rotation coppice on fallow lands in northernBenin increased significantly when weedingwas done three times during 42 months, whentotal biomass reached on average 3.67 tma ofdry matter in weedy plots and 11.63 tma inweed-free plots. Most of the biomass was mar-ketable in local markets.

Handling after harvest Logs should be pro-cessed soon after felling or treated with pre-servatives to avoid fungal and insect attacks

Genetic resources Doniel!to onDen has a

large but irregular distribution, and is locallycommon, often gregarious. It Is therefore notconsidered to be threatened by genetic erosion,although no information is available on its cur-rent exploitation for timber and firewood, andas medicinal plant.

Prospects Doniellio onDents an importanttimber tree in the drier forest zone of WestAfrica, as it is fast-growing, fairly large and itswood is easy to work. It has a range of otheruses, and especially its gum and bark are ex-tensively used medicinalIy. More research is

needed, however, on the pharmacology of thegum and bark to evaluate their potential. Atthis moment the antibacterial and arithelmin.

tic activities seem promising. More informationis also needed concerning its potential as afodder plant

Major references Arbonnier, 2004; Avohouat a1. , 2011; Baerts & Lehmann, 2012; Bolza &Keating, 1972; Burki11, 1995; de in BStrella ata1. , 2010; Katende, Birnie & Tengnas, 1995;Neuwinger, 2000; 0teng-Amoako (Editor),2006; Takahashi, 1978

Other references Adama at a1. , 2009; Adu-biar0, 01aofe & Akintay0, 2011; chinadu at al. ,2003; chinadu at a1. , 2004; Balogun & Adebayo,2007; Bellefontaine at a1. , 2000; CTFT, 1955a;EI Mahmood, Doughari & Chanji, 2008; Ha-yashi & Carsky, 1997; Kabor6 at a1. , 2010;Louppe & Ouattara, 1996; Louppe, M'Bla &Coulibaly, 1994; Menut at a1. , 1994; Onwu-kaeme, 1995; Onwukaeme, Lot & Udoh, 1999;OSakwe, Steingass & Drochaner, 2004; Schwobat a1. , 2008; SEPASA1, , 2012; Taita, 2000;Wilczek at a1. , 1952.

Sources of illustration Brenan, 1967;Eggeling & Dale, 1951.

Authors G. H. Schmelzer & D. Louppe

DANIELLIATHURIFERABenn

Protologue Pharm. Journ. Trans. 14: 253(1854)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Vernacular names GOPal tree, frankin-cense tree, gum copal, Niger copal, Sudan copal(En). Acacia copaldo Sudao (Po).

Origin and geographic distribution Doniel-Jig thurifero occurs from Senegal east to west-ern Nigeria

Uses The aromatic wood of Donielliu thu-rifero, traded as 'faro' and 'daniellia', is usedfor light construction, planks, light flooring,joinery, Interior trim, boat building, furniture,toys, novelties, boxes, crates, agricultural jin-PIements, draining boards, turnery, hardboardand particle board. It is suitable for veneer,plywood and pulpwood, as well as firewood andcharcoal productionBeehives are made from the bark. The twigsare used as chew sticks. The tree yield a resinor gum, known as 'gum copal' and used forcosmetics, perfume, pomades, varnish, soapand fumigation. The gum 1STubbed on the bodyfor its fragrance, usually mixed with clay, and

I.

Doniellio thurifero - wild

it can be used to mend broken pottery. Theseeds yield a clear oil, which is made in Liberiainto a hair dressingIn Ghana a decoction prepared from the POW-dered gum is taken by the firobo people totreat cough. The gum is used as embrocation totreat skin diseases, including ringworm. InSierra Leone Doniellio thurifero is planted asornamental tree along roadsides. The gum Isburnt to keep away evilspirits.

Production and international trade In1983 C6te d'TVoire exported 73,000 ing of logs oftaro (Doniellio spp. ). Currently Doniellio thu-rifero and other Doniellio spp. have no jin-portance on the international timber market,but their wood is commonly used locally inWest Africa

Properties The heartwood is pale pinkishbrown to reddish brown with dark streaks and

is distinctly demarcated from the wide, whitishor pale yellow sapwood. The grain Is Inter-locked, texture medium coarse. The wood isstrongly scented and has lustrous and oily sur-faces.

The wood is lightweight to medium-weight,with a density of 490-660 kg/ina at 12% inois-ture content, moderately soft but tough andstrong. It air dries easily, although in thickboards there might be slight distortion andcollapse. The shrinkage rates are moderate,from green to oven dry 2.6-4.5% radial and5.6-8.2% tangential. Kiln drying is fairly fastwith little degrade. Once dry, the wood is inod-erately stable in service. At 12% moisture con-tent, the modulus of rupture is 66-122 N/min2,modulus of elasticity 8130-9220 N/min2, coin.pression parallel to grain 30-46 N/min2, shear

6.5-7 N/mm2, cleavage 11-18 N/min, Jankaside hardness 5560 N, Janka end hardness6850 N and Chalais-Meudon side hardness1.2-3.8.

The wood is easy to saw and work with bothhand and machine tools, with little bluntingeffect on saw teeth and cutting edges. Workedsurfaces tend to be woolly, so cutting edgesmust be kept sharp. Although the wood planessmoothly, tearing of quarter-sawn surfaces dueto interlocked grain may occur and a reducedcutting angle of 20'is recommended. The woodstains satisfactorily but needs a filler beforepolishing. It nails well and glues correctly. Itcan be readily peeled into veneer, but gluing ofveneer can be difficult due to warping duringdrying. The wood is not durable, being suscep-tible to attacks by fungi, termites, pinhole bor-ers and Lyetus. The heartwood is moderatelyresistant to impregnation, but the sapwood iseasy to treat with preservativesThe wood contains 42-43% cellulose, 27.5-29%Itgrim, 19-19.5% pentosan and 1.0-1.1% ashThe solubility is 2.5-2.8% in alcohol-benzene,2.5-3.1% in hot water and 18.0-20.4% in a I%

NaOH solution

The resin from the heartwood is dark brown,oily and sticky, and can be saponified, althoughit has a low saponification value. One of themain compounds of the gum is the diterpeneo1eoresin daniellic acid (illurinic acid), as wellas ozic acid and the alcoholozol

Adulterations and substitutes The wood

of Doniellio thurifero is similar to that of Do.ritel!to ogeo (Harms) Rolfe ex Holland and Do-ritellio h!Ginei Pierre ex Achev. , and has SImi-Iar uses.

Description Deciduous, medium-sized tofairly large tree up to 35(-45) in tall; holestraight and cylindrical, up to 150 cm in diam-eter, without buttresses; bark surface smoothbecoming scaly in older trees, greyish white orgreenish with horizontal brownish bands, innerbark thick, orange brown to reddish brown;crown dense, wide, flattened; twigs glabrous.Leaves alternate, panpinnately compound with5-9 pairs of leaflets; stipules up to IOC15) cmx Tel. 5) cm, soon falling; petiole 1.5-3.5(-5.5)cm long, rachis (10-)13-17(-27) cm long, shal-lowly channelled, at least at base, with smallglands at insertion of basal pair of leaflets;petiolules 4.5-8 min long; leaflets opposite,ovate-elliptical to oblong-lanceolate, up to 12(-18) cm x 6 cm, basal and apicalleaflets smallerthan middle ones, base curieate, asymmetrical,apex acuminate, papery to leathery, glabrous,

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DANiELLIA 279

280 TIMBERS2

long, glabrous; cotyledons fleshy, oblong, 2.5-3cm long, erect; first leaves alternate, with 2-3pairs of leaflets

Other botanical information Doniellio

comprises 10 species, which are nearly all con-fined to the forest areas of West and Central

Africa, with only I species extending to Sudanand Uganda

Anatomy Wood-anatorritcal description CAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(9: vessels exclusively solitary (90% or more));13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 27: intervessel pits large (^ 10 prn); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to Intervessel pits in size andshape throughout the ray cell; 43: mean tan-gentialdiameter of vessellumina z 200 pin; 46S 5 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;(68: fibres very thin-walled); 69: fibres thin- tothick-walled. Axial parenchyma: 78: axial pa-Tenchyma scanty paratrachea1; 85: axial paren-chyma bands more than three cells wide; (89:axial parenchyma in marginal or in seeminglymarginal bands); 91: two cells per parenchymastrand; 92: four (3-4) cells per parenchymastrand. Rays: 97: ray width I-3 cells; 106: bodyray cells procumbent with one row of uprightand/or square marginal cells; 107: body raycells procumbent with mostly 2-4 rows of up-right and/or square marginal cells; 1/5: 4-12rays per mm. Stoned structure: 1/8: all raysstoned; 1/9: low rays stoned, high rays non-stoned; 120: axial parenchyma and/or vesselelements stoned; 121: fibres stoned. Secretoryelements and cambial variants: 127: axial ca-nals in long tangentiallines; 129: axial canalsdiffuse.

(L. Awoyemi, P. E. Gasson & E. A. Wheeler)Growth and development Seedlings reach

about 12 cm tall after 1.5 months and 15 cmtall after 2.5 months. In Guinea growth in thefield has been reported to be quite slow; seed-lings reached 35 cm in height after I year, 60cm after 2 years and 150 cm after 5 years. InLiberia fruit bearing Doniellio th!, rifero treesof about 10 in tallwere observed. The tree usu-

ally sheds its leaves at the end of the rainyseason, and flowering occurs when the tree isleafless, mainly from August to JanuaryFruits ripen about 5 months after flowering.

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with many translucent glandular dots, pin-nately veined with 10-25 pairs of lateral veinsInflorescence an axillary or terminal compoundraceme (6-)9-Ile15) cm long, glabrous, with6-10 lateral branches. Flowers bisexual, zygo-inorphic, slightly scented; pedice1 8-Ile17)min long, glabrous, enlarging in fruit, near themiddle with 2 caducous bracteoles; sepals 4,oblong, 1.5-2 cm long, glabrous but marginshairy, with few gland dots on external surface;petals 5, white to yellowish, unequal, oblong totriangular, 3 petals I-1.5 cm long and 2 petalsI-3 mm long, short-hairy to nearly glabrous,with few glandular dots; stamens 10, 3-4.5 cmlong, 9 united at base and I free; ovary superi-or, oblong to rounded, 0.5-I cm long, glabrous,with stipe c. 0.5 cm long, style up to 2.5 cmlong. Fruit an obliquely oblong, flattened pod7-9 cm x 3.5-5 cm, with stipe up to I cm long,glabrous, green becoming brown, dehiscingwith 2 woody valves, I'Seeded. Seed oblong,flattened, 1.5-3 cm long, smooth, deep red tobrown, attached to one of the valves by a c. 1.5cm long funicle. Seedling with epigeal germina-tion; hypocoty1 4-5 cm long, epicotyl c. 6 cm

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When the fruit dries, the two valves start tocurl, releasing the seed, which remains at-tached to one of the valves by a long funicle.When the valve with the seed falls, it rotatesrapidly and may be spread by wind far fromthe mother tree.

Ecology Donie!lid thurifero occurs in scat-tered groups in evergreen forest, in primary aswell as secondary forest, sometimes also ingallery forest and wooded savanna, from sea-levelup to 400 in altitude. It prefers damp are-as but avoids swampy conditions

Propagation and planting Natural regen-eration of Doniellio thurifero is abundant inforest gaps and disturbed forest, as it is a pm-neer and light demander. There are 600-800seeds per kg. The germination rate of freshseeds is usually high, 50-95%, and germinationstarts about 2 weeks after sowing. Shadingshould be removed soon after germination. Theseedlings are ready for planting into the fieldafter one year in the nursery. In Guinea themortality of seedlings planted in full sun was50% after 5 years, but this was probably due toa lack of weeding after planting

Management In evergreen forest in C6ted'TVoire, the average density of trees with abole diameter of more than 90 cm was locally10 trees/ha, and of saplings with a hole diame-ter of 10-20 cm about 20 per ha. In Sierra Leo-ne plantations of Doniellio thurifero have beenestablished. Systematic thinning of the forestwas found to be beneficial in terms of recruit-

merit and growth. However, the difference be-tween heavy and light thinning was negligible,and the maximum recommended thinning in-tensity was 28% of the initial basal area (5-6in2ftja removed).

Diseases and pests Logs should be pro-cessed quickly after felling or treated with pre-servatives because the wood is susceptible todiscoloration by blue-stain fungiand attacks byinsects. Fallen seeds are often eaten by ro-dents.

Harvesting Trees are harvested by the se-Iective logging system; larger trees are occa-sionally defective. Some caution is needed dur-ing harvesting operations because logs mayhave brittle heart

For harvesting of the gum a deep hole is cutnear the base of the tree, and the exudate Iscollected. However, Doniel!io thurifero doesnot yield commercial quantities of gum.

Genetic resources Doniellio thurifero isfairly widespread and there are no indicationsof overexploitation; therefore it does not seem

to be threatened by genetic erosion. However,it is probably much confused with other Do-rite!!to spp. , and therefore its exact status isnot known

Prospects Donie!lid thurifero has not beenconsidered a timber tree of commercial jin-

portance because its wood is susceptible to fun-gal attacks and the gum may Interfere withprocessing. However, it may have good pro-spects for the production of veneer and ply-wood. More information is needed on growthrates and methods to improve regeneration tobe able to draw conclusions about possibilitiesfor sustainable exploitation from natural for-est. Further research of the exudate is needed

on the possible usage in the varnish and cos-metic industry

Major references Bolza & Keating, 1972;Burki11, 1995; CTRAD Forestry Department,2008; Cooper & Record, 1931; de to EstreUa ata1. , 2010; Hawthorne & Jongkind, 2006; Hu-hert, undated; Irvine, 1961; Kryn & Fobes,1959; Takahashi, 1978.

Other references ATIBT, 1986; CTFT,1955a; de in Mensbruge, 1966; Durrieu de Ma-dron at a1. , 1998a; G6rard at a1. , 1998; Haw-thorne, 1995; Hawthorne & Gyakari, 2006;Keay, 1989; Sandermann, Ehlers & Febt, 1963;Savard, Besson & Morize, 1954; Savill & Fox,1967; Voorhoeve, 1965

Sources of illustration de in BStrella at al. ,2010

Authors N. Nyunai

DASYLEPiS 281

DASYLEPISINTEGRAWarb. ex Engl.

Protologue Abh. Preuss. Akad. Wiss. : 44(1894)

Family F1acourtiaceae (APG: AChariaceae)Synonyms DOSylepis leptonhyllo GIIg (1908).Origin and geographic distribution DOSy-

Iepis tritegro is restricted to south-easternKenya and north-eastern Tanzania

Uses The wood is locally used for furniture,utensils, clubs and tool handles, and also asfirewood.

Properties The wood is moderately hardBotany Evergreen shrub or smalltree up to

15(-35) in tall; bole slender; bark surfacesmooth, becoming flaky, grey-brown, Innerbark orange-red. Leaves alternate, simple;stipules early caducous; petiole 5-7(-10) minlong; blade elliptical to lanceolate or oblong, 8-16 cm x 2.5-6 cm, curieate to obtuse at base,short-acuminate at apex, margins entire to

282 TIMBERS2

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DOSylepis tritegro - wild

remoteIy toothed, papery to thin-leathery, gla-brous, pinnately veined with 5-9 pairs of loop-ing lateral veins. Inflorescence a drooping axil-Iary raceme up to 12 cm long, glabrous, laxlyflowered. Flowers bisexual or male, regular;pedicel slender, 5-8(-20) mm long; sepals 3-4,slightly fused at base, rounded to elliptical, c. 5mm long, pinkish-reddish, glabrous; petals 7-8,similar to sepals but slightly longer, more whit-ish and with basal hairy appendage inside;stamens 15-20, c. 5 min long; ovary superior,glabrous, I-celled, style very short, with 3 shortbranches. Fruit a globose capsule 1.5-2 cm indiameter, short-pointed, with 3-4 longitudinalgrooves, glabrous, pale purple, tardily dehis-cent with 3 valves, I-2(-5)-seeded. Seedsovoid-angular, 8-9 min long, glabrous.Flushes of young leaves are reddishDOSylepis comprises 6 species and is related toErythrospermum, Rowsonio and Scottellio.DOSylepis tritegro resembles DOSylepis race-inOS0 011v. , which is a small to medium-sizedtree up to 20 in tall with hole up to 45 cm indiameter with a much larger area of distribu-tion, from Guinea east to western Uganda andsouth to Gabon and DR Congo. Although usesof DOSylepis rocemoso have not been recorded,its wood is possibly locally used for similarpurposes as that of DOSylepis tritegro.

Ecology DOSy!epts tritegro usually occurs inthe understorey of moist forest at 900-2250 inaltitude. It is locally co- dominant

Genetic resources and breeding DOSyle-pts tritegro has a restricted area of distribution,which is severely fragmented. Therefore, it hasbeen included in the IUCN Red List as vulner-able.

Prospects The wood of DOSy!epis tritegrowill remain of limited importance for local ap-PIications. The small size of the hole makesthat it has Do prospects for the internationaltimber market, and focus should be primarilyOn Its protection.

Major references Lovett & Clarke, 1998;Lovett, Ruffo & Gereau, 2003; 0bama &Breteler, 2004; SIeumer, 1972a; SIeumer, 1975.

Other references Beentje, 1994; Chase ata1. , 2002; InsideWood, undated; Miller, 1975

Authors R. H. M. J. Lemmens..

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DEINBOLLLAGRANDiFOLiA Hook. f.

Protologue Hook. , Niger F1. : 249 (1849).Family SapindaceaeOrigin and geographic distribution Dein-

bonier grundtfolio occurs from Sierra Leone toTogo and also in Garnero0n, Chad, the CentralAfrican Republic and Gabon

Uses The wood of Deinbollio grondifo!io isused in West Africa for house building and forplanks. The fleshy fruit pulp is edible and theslightly oily seeds are eaten as well. Juice fromthe mashed inner bark is used as eye-drops totreatjaundice.

Properties The wood of Deinbollio grondifo-lid is pinkish white and not durable.

Botany Shrub or small palm-like tree up to8(-15) in tall; hole up to 30 cm in diameter;bark surface grey, scaly; twigs often zigzag,reddish with small white lenticels. Leaves al-ternate, panpinnately compound, up to 100 cmlong, with (5-)7(-10) pairs of leaflets; stipulesabsent; petiole 10-27 cm long, rachis up to 60cm long, grooved; petiolules c. I cm long; leaf-

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Deinbo!lid grondifolio - wild

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lets often alternate, sometimes opposite, oblongto narrowly obovate, 13-30 cm x 5-8 cm, curie-ate and asymmetrical at base, acuminate atapex, papery, minutely hairy or scaly, pinnate-Iy veined with 12-16 pairs of lateral veinsInflorescence an axillary panicle 10-20(-50) cmlong, yellowish or brownish short-hairy. Flow-ers unisexual, regular, 5-merous, greenish yel-low; pedice1 1.5-2 mm long; sepals free, round-ed, up to 5 min long, nearly glabrous; petalsfree, ovate, up to 4 mm long, halry at margins,inside with 2 small scales at base; stamens 8-17; disk annular; ovary superior, 2-3-lobed,style up to 5 mm long. Fruit a globose to ovoidberry up to 1.5 cm long, fleshy, glabrous, or-ange when ripe, indehiscent, I'Seeded. Seedovoid, c. I cm long, brown to blackish. Seedlingwith hypogeal germination; epicotyl c. 6 cmlong, hairy; first 2 leaves opposite, with 2 ellip-ticalleaflets up to 3.5 cm x 1.5 cm.Deinbollio comprises 30-40 species and occursin tropical mainland Africa and MadagascarThe wood of many species developing into atree is used. Deinbo!lid curieifolio Baker isfound from Guinea to C6te d'Ivoire and inGamero0n and Gabon. Although it is only asmalltree up to 4 in tall, the wood is very hardand used for building. The fruit pulp is eatenA steam bath, to which crushed bark andleaves are added, is prepared to overcome fa-tigue and as a cure for painful feversDeinbolliu Iuluo-tomentel!o Baker f. (synonym:Deinbo!lid montuscu!a Radlk. ), a small tree ofup to 9 in tall, occurs in the Central AfricanRepublic, Gabon, DR Congo, Uganda, Tanzaniaand Angola. In DR Congo its wood is used forconstruction

At least 2 of the 7 endemic Deinbollio spp. ofMadagascar are sought after for their wood.The wood of Deinbollio macrocorpo Capuron isused in boat building and for firewood. Deinbol-jig peruil!ei (Blume) Radlk. is appreciated forthe quality and red colour of its wood, which isalso used in boat building. It is also used as asupport in vineyards and as firewood

Ecology Deinbollio grondtfolio is usuallyfound in the undergrowth of deciduous forest.It is quite shade tolerant.

Genetic resources and breeding As Dein-bouto grondifolio is widespread and not inten-SIvely exploited there seem to be no seriousthreats

Prospects The small stature of Deinbo!jiggrondifolio and other Deinbollio spp. limitstheir usefulness as timber-producing trees.

Major references Burki11, 2000; Hawthorne

& Jongkind, 2006; Menninger, 1977.Other references Bolteau, Bolteau & A1-

10rge-Boiteau, 1999; Brown at a1. , 2009; Davies& Verdcourt, 1998; de in Mensbruge, 1966;Fouilloy & Hall6, 1973a; Hauman, 1960; Li-sowski, 2009; Martin at a1. , 2009; Schatz, 2001

Authors C. H. BOSch

DETARIUMMACROCARPUMHarms

DETERiuM 283

Protologue Bot. Jabrb. Syst. 30: 78 (1901)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Origin and geographic distribution Detori-

urn macrocorpum occurs in south-eastern Ni-gena, Cameroon, Central African Republic,Equatorial Guinea and Gabon.

Uses The wood, often traded as 'mambode','enouk' or 'alen' from Gabon, 'amouk' fromGarnero0n and 'enuk' from Equatorial Guinea,is used for flooring, joinery, Interior panelling,stairs, furniture, cabinet work, turnery andsliced veneer.

In Cameroon the bark is an ingredient of hunt-Ing poison.

Production and international trade Detori-

urn macrocorp!, in timber Is traded in smallamounts on the international market, but sta-tistics are not available.

Properties The heartwood is reddish brownto coppery brown, often with dark brownstreaks, and distinctly demarcated from the 7-10 cm wide, pinkish sapwood. The grain isstraight or interlocked, texture mediumThe wood is medium-weight, with a density of480-660 kg/ina at 12% moisture content. It airdries slowly, with minimal risk of distortion

.

Detorit, in macrocorpum - wild

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284 TIMBERS2

and checking. The rates of shrinkage fromgreen to oven dry are 3.3-4.3% radial and 4.8-6.0% tangential. Once dry, the wood is moder-ately stable in service. At 12% moisture con-tent, the modulus of rupture is 83-1/5 N/min2modulus of elasticity 11,100-13,100 N/mm2compression parallel to grain 46-64 N/mm2and Monnin side hardness 3.2-4.6.

The wood saws and works satisfactorily withmoderate blunting effect on saw teeth and cut-ting edges. However, the presence of resin maycause some difficulties. The use of a filler is

needed to obtain a good finish. Pre-boring isrecommended in nailing and screwlrig. Thewood glues satisfactorily except when muchresin is present. It is moderately durable, beingonly moderately resistant to fungi, termitesand marine borers. The sapwood is susceptibleto dry-wood borers. The wood is moderatelypermeable to preservatives

Botany Medium-sized to very large tree upto 60 in tall; hole branchless for up to 20 in,usually straight, cylindrical, up to 150(-200)cm in diameter, with smallbuttresses or slight-Iy swollen at base; bark surface fissured, be-coming scaly, silvery grey to dark grey or grey-ish brown, inner bark hard, fibrous, brownoutside, pinkish inside; crown dome-shaped;twigs glabrous. Leaves alternate, panpinnatelyor Imparipinnately compound with 8-20 leaf-lets; stipules minute, early caducous; petiolules5-7 min long; leaflets alternate, ovate to ellip-tica1, 4-8 cm x 2-4.5 cm, slightly unequal atbase, acuminate at apex, papery, with translu-cent dots, glabrous, pinnately veined withmany lateral veins. Inflorescence an axillary,lax panicle c. 8 cm long, glabrous. Flowers bi-sexual, slightly zygomorphic; pedice1 2-3 mmlong; sepals 4, ovate to lanceolate, c. 5 mmlong, I slightly broader than other 3, whitish,glabrous outside, hairy inside; petals absent;stamens 10, free, c. 5 mm long; ovary superior,ellipsoid, c. 2 min long, densely hairy, I-celled,style c. 3 min long, curved. Fruit a drupe-like,globose to ovoid pod 7-8(-10) cm in diameter,slightly flattened, indehiscent, smooth andolive-green, pulp pale green and fibrous, stoneup to 6 cm in diameter, wrinkled, I-seeded.Seedling with epigeal germination; hypocotyl6-14 cm long, epicoty1 13-28 cm long; cotyle-dons thick and fleshy, spoon-shaped; firstleaves alternate, with 8-10 leaflets.The fruits usually ripen in the dry season. Ele-phants eat them and disperse the stones. Dui-kers feed on the fruit pulp, but usually do notswallow the stones and thus do not play an

important role in seed dispersal, although theyellow-backed duiker may swallow the stones.Gorillas, chimpanzees, mandrills and pigs canbreak the tough wall of the stone and eat theseed

Detorium comprises 3 species and is confinedto Africa. It is related to GOPoifero. The 3 spe-cies are inorphologically quite similar, but eco-logically different.

Ecology Detorium inderoco, pum occurs inhumid lowland forest, also in gallery forest. InEquatorial Guinea and Gabon it is usuallyfound inland and not close to the coast. The

annual rainfall in the area of distribution is

about 1400 mm. Detorium ingcrocorpum usual.Iy occurs on well-drained sites.

Management In general, larger trees ofDetorium mererocorpum occur scattered and inlow densities in the forest. In south-western

Garnero0n the average density of holes of morethan 60 cm in diameter is 0.07 per ha, with anaverage wood volume of 0.73 ing/ha. In Gabonthe average wood volume has been recorded as0.08 mama only. Freshly harvested logs arereported to float in water, and can thus betransported by river. The durability in the for-est is moderate, and it is recommended to treatthe logs with preservatives or to process themrapidly.Elephants often feed on the bark and the holeIs often damaged and swollen at the baseHowever, the bark fibres break apart, prevent-ing the elephants from removing large slabs

Genetic resources and breeding Detori-urn inderocorpum is not very widespread andoccurs scattered in the forest, usually in lowdensities. This may make it liable to geneticerosion and caution is needed in case of in-

creasing exploitationProspects Detortum inderocorpum is a val-

uable timber tree that, however, occurs in toolow density to be commercially important. Vir-tually no technical information exists on manyaspects, including growth rates, propagationand management in natural forest and planta-tions, and much research is still needed to as-sess its possibilities for exploitation on a sus-tainable basis

In view of the presence of edible fruits andseeds and widespread medicinal applications inother Detorium spp. , phytochemical and phar-macologicalinvestigations of Deterrium ingcro-corpum are recommended.

Major references CIRAD Forestry Depart-merit, 2008; Neuwinger, 1996; Neuwinger,1998a; Vivien & Faure, 1985; White & Aber.

nethy, 1997.Other references Aubr6ville, 1968; Aubr6-

vine, 1970; Christy at a1. , 2003; de Saint-Aubin,1963; Keay, 1989; Leonard & Doucet, 1997;Lewis at a1. , 2005; Normand & Paquis, 1976;Tailfer, 1989; Wilks & TSSemb6, 2000

Authors R. H. M. J. Lemmens

DETARluMSENEGALENSE J. F. Ginel.

Protologue Syst. nat. 2(I): 700 (1791)Family Caesalpiniaceae or, eguminosae - Cae-

salpinioideae)Synonyms Detorium heudelotio, turn Baill

(1866).Vernacular names Tallow tree, dattock,

detah (En). Grand d6tar (Fr)Origin and geographic distribution Detori-

urn senego!errse occurs from Senegal and Gain-bia east to Sudan, and south to northern DRCongo. It is planted as fruit tree and as orna-mental shade tree within its area of distribu-

tion and has been planted in the Caribbeanregion.

Uses The wood is used in house construction

for planks, posts, poles and piles, and for furni-ture, fences, mortars, tool handles, boat-ribsand canoes. It is suitable for flooring, joinery,interior trim and framework, mine props, vehi-cle bodies, boxes, crates, veneer and plywood. Itis also used as firewood

The greenish and sweet-acidulous fruit pulp isedible, and can be eaten raw or cooked. It isalso used to prepare sweetmeat or as an ingre-dient of ice-cream. However, it may also betoxic, and caution is needed. The seed is oilyand edible, and pounded seed is used as cattle

feed. In Nigeria the seed flour is used tradi-tionaUy as a thickening agent in foods. Thegum from the bark is used to fumigate clothingand houses. Roots have been used to preparebird-lime and root sap as coating in brass cast-Ing

Detorittm senegolense is an important medici-nal plant. Several plant parts are used in tra-ditional medicine. Bark decoctions or macera-

tions are taken in case of heavy loss of blood, totreat digestive disorders, bronchitis, pneumo-nia and stomach-ache, and to expelthe placen-to after childbirth. Bark powder is applied towounds, burns and skin complaints, and barkpulp is eaten against tuberculosis and as atonic. The bark is also used as arrow poisonand soap substitute, and is added to palm wineto accelerate fermentation and to make it more

bitter. Root decoctions are administered as

anodyne and to treat intestinal complaints,marasmus, debility and anaemia. Leaf andshoot decoctions are used to treat fever, trypa-nosomiasis, dysentery, anaemia, conjunctivitis,arthritis, inflammations, fractures, boils andskin complaints. Fruit pulp is applied for thetreatment of kidney pain, spinal tuberculosis,syphilis, cough, rheumatism and leprosy, andin mixtures with other fruits as a stimulant

Seeds are taken as antidote against arrow poi-son and as emetic, and the smoke of burntseeds as mosquito repellent. Detorium servego-lense is planted as an ornamental shade tree.

Production and international trade In the

past there has been some export of the woodfrom West Africa to the United Kingdom andthe United States. In Gambia Detorium sene-

golense belongs to the most important timbertrees. The wood is sold on local markets in Su-

dan. The fruits are commonly sold on localmarkets in West Africa. However, statistics arenot available, neither for the wood nor for thefruits

Properties The heartwood is pale yellow,becoming reddish brown towards the centre ofthe hole, and distinctly demarcated from thepaler, thick sapwood. The grain is straight tointerlocked, texture medium to fine. The woodis lustrous and has a pleasant odourThe wood is moderately heavy, with a densityof (600-)710-850(-900) kg/ms at 12% moisturecontent. It should be air dried carefully becauseit has a tendency to surface checking and split-ting. The rates of shrinkage are moderate, fromgreen to oven dry 3.7-6.2% radial and 5.7-8.4%tangential. Once dry, the wood is only moder-ately stable in service. At 12% moisture con-

DETERiuM 285

.

Detor, urn senego!errse - wild

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286 TIMBERS2

tent, the modulus of rupture is 84-148 N/mm2modulus of elasticity 11,380-15,290 N/mm2compression paraUelto grain 38-54(-63) N/min2,cleavage 11-19 N/mm, Janka side hardness8220 N, Janka end hardness 9340 N and Cha-Iais-Meudon side hardness 1.7-3.3

The wood is rather difficult to saw and work,but with care it can be finished well. It is some-

times difficult to plane due to the presence ofinterlocked grain. It nails satisfactorily withoutsplitting, and it glues well. It is durable, beingresistant to termites and moderately resistantto pinhole borers and marine borers. The sap-wood is susceptible to Lyetus attackThe fruits are edible, but those of some treesare toxic. The poisonous fruits can be recog-nized in the forest because they are left be-neath the mother tree by animals, while theedible ones are soon eaten or taken. A toxic,bitter compound has been isolated from thefruit, as well as an acid-tasting compoundcalled detaric acid.

The composition of the fruit pulp per 100 gedible portion is: water 67 g, energy 485 kJ(1/6 kcal), protein 1.9 g, fat 0.4 g, carbohydrate29.6 g, fibre 2.3 g, Ca 27 ing, P 48 ing, thiamine0.14 ing and ribonavin 0.05 ing. The fruit pulpis rich in ascorbic acid, 1000-2000 ing per 100g

All arithocyanidin alkaloid has been isolatedfrom the bark. This compound showed antibac-tenal activity against a panel of pathogenicbacteria. Leaf extracts showed antiviral activi-

ty against a panel of human and animal virus-es. Flour prepared from the seeds containssignificant amounts of water-soluble non-starch polysaccharides. In tests the flour re-duced postprandial blood glucose and insulinconcentrations in humans. The main polysac-charide is a xyloglucan

Adulterations and substitutes The bark,roots, leaves and fruits of Detorium micro-euroum Gulll. & Perr. are used for similar me-dicinal purposes as those of Detorium senego-lense and the fruits are also eaten. The wood is

SLmilar, but Detorium microcorpum is usuallya smaller tree and therefore its major use is asmedicinal plant. The wood of GOPotfero spp.may be confused with that of Deterrium senego-lense.

Description Medium-sized to fairly largetree up to 35(-40) in tall; hole branchless up to12(-15) in, straight or irregular, cylindrical, upto 60(-100) cm in diameter, without buttressesbut sometimes swollen at base; bark surfacefinely fissured, becoming scaly, greyish to

I

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Detorium senegalerrse - I, flowering twig, . 2,sepals, . 3, flower with sepols reinoued, . 4, fruit.Source. F10re orig!ytique du Bentn

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blackish, with large, round lenticels, innerbark thick, fibrous, red-brown, with somesticky gum; crown large, dark green, withspreading branches; twigs slightly hairy toglabrous. Leaves alternate, panpinnately orimparipinnately compound with (6-)8-12 leaf-lets; stipules minute, early caducous; petioleand rachis together 10-23 cm long, swollen atbase; petiolules 3-6 min long; leaflets alter-nate, ovate to elliptical, 3-8 cm x 2-4 cm,slightly unequal at base, rounded to slightlynotched at apex, papery, with translucent dots,minutely hairy beneath, pinnately veined withmany lateral veins. Inflorescence an axillary,lax panicle up to LOG15) cm long, short-hairy.Flowers bisexual, slightly zygomorphic, nearlysessile; sepals 4, ovate, 3.5-4 min long, Islightly broader than other 3, whitish, sparselyhairy to glabrous outside, hairy inside; petalsabsent; stamens 10, free, 4-5 min long; ovarysuperior, ellipsoid, c. 2 min long, densely hairy,I-celled, style 3-4 min long, curved. Fruit adrupe-like, globose to ovoid pod 4-6(-7) cm indiameter, slightly flattened, indehiscent,smooth and greenish, eventually blackish, pulp

I:, I

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, 3

4

greenish and very fibrous, stone up to 4.5 cm indiameter, I'Seeded. Seedling with epigeal ger-mination; hypocoty1 2-6.5 cm long, epicotyl a15 cm long; cotyledons thick and fleshy, spoon-shaped; first leaves alternate, with 8-10 leaf-lets

Other botanical information Detorium

comprises 3 species and is confined to Africa. Itis related to GOPoifero. The 3 species are inor-phologically quite similar, but ecologicalIy dif-forent. Detorium microcorpum Gulll. & Perr. istypically for dry savanna areas and differsfrom Detorium senego!errse in Its smaller treesize, usually fewer, larger and more leatheryleaflets, more compact inflorescences, sepalshairy outside and slightly smaller fruits.Detorium ingcrocorpum Harms occurs in hu-inId forest

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: I: growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; 13: sim-PIe perforation plates; 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 F1m); 27:intervessel pits large (z 10 F1m); 29: vesturedpits; 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; 43: mean tangentialdiameter of vessellumina Z 200 prn; 46: S 5vessels per square minimetre; 47: 5-20 vesselsper square minimetre. Tracheids and fibres61: fibres with simple to minutely borderedpits; 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); 85:axial parenchyma bands more than three cellswide; 86: axial parenchyma in narrow bands orlines up to three cells wide; 91: two cells perparenchyma strand; 92: four (3-4) cells perparenchyma strand. Rays: (97: ray width I-3cells); 98: larger rays commonly 4- to 10-senate; 104: all ray cells procumbent; (106body ray cells procumbent with one row of up-right and/or square marginal cells); 1/5: 4-12rays per mm. Secretory elements and cambialvariants: 127: axlal canals in long tangentiallines. Mineral inclusions: 136: prismatic crys-tals present; 142: prismatic crystals in chain-hered axial parenchyma cells(R. Shanda, A. A. Oteng-Amoako & P. E. Gas-son)

Growth and development Trees are usual-Iy leaness for about 3 weeks, and flowering

occurs when young leaves have developed, of-ten towards the end of the dry season. In C6ted'Ivoire trees flower in February-May and inSudan in February-August. Fruits take about6 months after flowering to mature. They areeaten by elephants, which disperse the stones.Chimpanzees also eat the fruit pulp, and insome regions they crack the fruit stone by us-ing stones and eat the seed. Trees suckerabundantly.

Ecology Delorium senegalerrse occurs inlowland dry forest and gallery forest in savan-na areas. It is often found along watercoursesThe tree has some resistance to fire

Propagation and planting Detorium sene-golense is usually propagated by stones. Thereare about 40 stones per kg. They germinateafter 6-10 weeks, but the germination rate islow. Grafting is possible. In tests in Senegal,apical grafts (cleft graft and splice graft) gavethe best results. For grafting, the end of thedry season is the besttime.

Management Locally Detortum senegolenseis very common, but in other regions it occursonly scattered or is even rare. In Sierra Leonethe average density of Detorium senego!errsetrees of over 60 cm hole diameter has been rec-

orded as only 0025-0.15 per ha.In Senegal Detorium senegolense is used to-gether with Neocoryo macrophy!10 (Sabine)Prance in agroforestry programmes, which areimportant for local human populations. Theaverage density in these agroforestry systemsis 17 holes per ha. The fruits are consideredvaluable and this makes it worthwhile for local

populations to harvest in a sustainable way.Yield In agroforestry systems in Senegal,

the average fruit production is 3.1 t per ha.Genetic resources Detorium senegalerrse is

quite widespread and locally common and thusdoes not seem to be in direct danger of geneticerosion. However, in several regions within Itsdistribution area it is uncommon or even rare

and has been over-exploited or subject to regu-Iar forest fires. Detor, urn senego!errse is valuedin agroforestry systems in Senegal and is assuch protected

Breeding A domestication programme hasbeen carried out in Senegal, aiming at selectingand cloning trees of a superior phenotype be-fore introducing them to local populations

Prospects Detorium senegolense is a multi-purpose species with several opportunities forincreased use. It is useful as a timber tree, butalso produces edible fruits and good-qualityseed flour, and is considered a valuable medic-

DETERiuM 287

288 TIMBERS2

inalplant. It is worth domesticating at a largerscale. The selection of superior trees for differ-ent purposes, e. g. good hole shape and size fortimber production and optimal fruit quality,should be stimulated, and more research isneeded on proper methods of vegetative propa-gation and on optimization of growth rates.Phytochemical analyses are needed to supportthe uses in traditional medicine and to be able

to judge the risk of toxic compounds in thefruits

Major references Arbonnier, 2004; Bolza &Keating, 1972; Burki11, 1995; Duke, 2001; Jan-ick & Paull(Editors), 2006; Niane & Moctar,2002; Pousset, 2004; Savill & Fox, 1967;Takahashi, 1978; Wilczek at a1. , 1952

Other references Abubakar at a1. , 2007;Aubr6ville, 1959b; Bouquet & Debray, 1974; ElAmin, 1990; Hawthorne, 1995; Hawthorne &Jongkind, 2006; Hegnauer & Hegnauer, 1996;Inngjerdingen at a1. , 2004; Keay, 1989; Ker-baro & Adam, 1974; Kudi & Myint, 1999; Lew-is at a1. , 2005; Neuwinger, 2000; Normand &Paquis, 1976; Okwu & Uchegbu, 2009;Onyechi, Judd & Ellis, 1998; Richter &Danwitz, 2000; Soloviev & Gaye, 2004; Tailfer,1989; Warig at a1. , 1996

Sources of illustration Ako6gninou, vander Burg & van der Maesen (Editors), 2006

Authors H. H. El-Kamali

.

DLALIUMAUBREVILLEIPellegr.

Protologue Bull. SOC. Bot. France 80: 463(1933).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Chromosome number2n=28

Origin and geographic distribution Digit-urn oubreuillei occurs in Guinea, Sierra Leone,Liberia, C6te d'TVoire and Ghana.

Uses There are few recorded uses of the

wood of Digitum erubreuillei. It is said that the

use of the wood is limited because the holes areoften short and crooked and because the woodis very hard. In Sierra Leone the wood is usedfor tool handles, but undoubtedly it is used forother purposes as well, similarly to other Di-ajium spp. It Is suitable for heavy constructionsuch as bridges, stairs and for poles and postsin house building, heavy flooring, mine props,ship building, vehicle bodies, railway sleepers,ladders, sporting goods, toys, novelties, vatsand agricultural implements. In Sierra Leonethe wood is used for charcoal production.

Digitum uubreui!lei - wild

The slightly acidulous fruit pulp is edible.Properties The heartwood is pale brown to

yellowish brown, often with reddish brownstreaks, and distinctly demarcated from thegreyish white to greyish yellow, up to 7 cmwide sapwood. The grain is usually straight,texture fine. The wood is heavy, with a densityof 900-1050 kg/in" at 12% moisture content,hard and tough. It air dries slowly and dryingshould be done with care to avoid warping andexcessive surface and end checking. Properstacking is needed, and it is recommended toquarter-saw logs for best results. The rates ofshrinkage are quite high, from green to ovendry 5.2% radial and 10.4% tangential. At 12%moisture content, the modulus of rupture is165-203 N/mm2, modulus of elasticity 20,600-25,500 N/min2, compression parallel to grain96-128 N/min2 and Janka side hardness

10,900-15,100 N.The wood is difficult to saw due to the presenceof silica; saw blades often heat up and stain thewood. The blunting effect on saw teeth andcutting edges is very high. The use of stellite-tipped saw teeth and tungsten-carbide cuttingedges is recommended. It is difficult to workwith both hand and machine tools, but planesto a good finish with a nice polish without theuse of a filler. The wood has a tendency of split-ting upon nailing and screwing, and pre-boringis recommended. It is durable, usually beingvery resistant to fungi and marine borers, andresistant to termite attacks. The heartwood is

resistant to treatment with preservatives. Thesaw dust may cause irritation to mucous mein-branes rowood workers.

Description Evergreen medium-sized to

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superior, ovoid, sessile, hairy, I-celled, stylecurved. Fruit a nearly globose, flattened pod1.5-2 cm in diameter, glabrous, black, withspongy pulp, indehiscent, I(-2)-seeded. Seedsrounded, flattened, c. I cm long, brown, finelyveined. Seedling with epigeal germination;hypocoty1 4-6 cm long, epicoty1 2-3.5 cm long;cotyledons fleshy, elliptical, up to 1.5 cm long,greenish; first leaves opposite, simple.

Other botanical information Digitum

comprises nearly 30 species and is pantropical.Approximately 15 species occur in mainlandtropical Africa and 3 in Madagascar, whereastropical Asia has about 7 species and tropicalAmerica one.

Apart from Digitum oubreuillei in West Africaand Digitttm perchyphy!Ium Harms in CentralAfrica, several other Diglium spp. produce use-ful wood. For some of these, the use of theiredible fruits is more important than that of thewood. This is the case for DidIturn gumeenseWind. from West and Central Africa, Digitumorigo!errse WeIw. ex Onv. and Didjinm errgleri-onurn Henriq. from Central and southern Mri-ca, and Digitum schlechteri Harms from south-ern Africa

Didjinm binindense Harms (synonym: DidIturnflewryiPenegr. )is a medium-sized tree up to 35in tall with hole up to 100 cm in diameter, oc-curring in Gamero0n, Equatorial Guinea andGabon. Its reddish brown to dark brown wood,which is heavy, with a density of about 1070kg/ms at 12% moisture content, and very hard,is suitable for similar purposes as that of Di-gnum dubreuillei. It is also used as firewood

and for charcoal production.Diglium corbisieri Staner is a medium-sized to

fairly large tree up to 35 in tall with hole up to100 cm in diameter, occurring in Congo and DRCongo, and possibly Gabon. Its pinkish brownwood, which is heavy, with a density of about1020 kg/ina at 12% moisture content, and veryhard, is suitable for similar purposes as that ofDialium dubreui!!at. It is also used as firewood

and for charcoal productionDigitum dinhlogei Harms is a small to medi-urn-sized tree up to 20(-30) in tallwith hole upto 50(-90) cm in diameter, occurring fromGuinea and Sierra Leone eastward to Gabon

and western DR Congo. Its wood is used forutensils; it is similar to that of Diolium dubre-ui!lei and suitable for the same purposes. It isalso used as firewood and for charcoal produc-tion. The fruit pulp is edible; a refreshing drinkis made by maceration of the fruit pulp in wa-ter. The acid-tasting leaves are eaten as vege-

3

Digitum oubreuillei - I, flowering twig, 2, flow-er, ' 3, fruits.Redrown ond adopted by G. WE. uon denBerg

\

DIALluM 289

fairly large tree up to 35(-65) in tall; holebranchless for up to 15(-30) in, straight orcrooked, angular, up to 90(-120) cm in diame-ter, with smallto fairly large buttresses up to 2in high; bark surface scaly with small, Irregu-Iar scales, greyish with yellowish brown patch-es, inner bark thin, brittle, yellowish outside,reddish inside, with a reddish, sticky exudate;crown rounded, dense; twigs with scatteredlenticels, soon glabrous. Leaves alternate, jin-panpinnately compound with (3-)5 leaflets;stipules 2-3 mm long, caducous; petiole I-3 cmlong, rachis 3-5 cm long; petiolules c. 0.5 cmlong; leaflets alternate, sometimes nearly op-POSite, obovate to elliptical, 4-13(-16) cm x 2-6cm, curieate to rounded at base, acuminate atapex, margins recurved, thick-papery, gla-brous, pinnately veined with 4-9 pairs of Iat-eral veins. Inflorescence a terminal or axillarypanicle up to 15 cm long, short-hairy. Flowersbisexual, zygomorphic, fragrant; pedice1 2-3min long; sepals 5, free, triangular to broadlyelliptical, c. 2.5 min long, hairy; petal I, spatu-late, c. 2.5 mm long, pale yellow; disk c. 1.5 mmin diameter, finely hairy; stamens 2; ovary

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290 TIMBERS2

table and to treat complaints of the respiratoryorgans, whereas leaf decoctions are taken as atreatment for jaundice, haematuria and feverBark infuslons are applied as purgative andexternalIy to treat arthritis.DidIturn zenkeri Harms (synonym: DidIturn so-yourit Harms) is a small to large tree up to 40in tall with hole up to 100 cm in diameter, oc-curring in Garnero0n, Equatorial Guinea, Ga-bon, Congo and DR Congo. Its dark pinkishbrown wood, which is fairly heavy, with a den-sity of about 800 kg/in3 at 12% moisture con-tent, and fairly hard, is suitable for similarpurposes as that of Dialium dubreui!lei. It isalso used as firewood and for charcoal produc-tion. Diglium zenheri is possibly conspecificwith DidIturn gumeense WindDiorlium excelsum Steyaert is a medium-sizedto very large tree up to 50 in tall with hole upto 150 cm in diameter, occurring in DR Congoand Uganda. Its dark brown wood, which isheavy, with a density of about 910 kg/in3 at15% moisture content, and hard, is suitable forsimilar purposes as that of Digitum aubreuillei.Digitum pentondrum Steyaert is a medium-sized to large tree up to 40 in tall with hole upto 100 cm in diameter, restricted to DR CongoIts pinkish brown wood, which is heavy, with adensity of about 930 kg/in3 at 12% moisturecontent, and hard, is suitable for similar pur-poses as that of Did!min oubreuillei.Digitum holtzii Harms (called 'inpepeta' inSwahili) is a shrub or small to medium-sizedtree up to 25 in tall, occurring in dry evergreenforest and riverIne forest in eastern Tanzania

and northern Mozambique. The heavy (density785-835 kg/ms at 12% moisture content), hard,brown wood with darker streaks is used for

poles in construction, boat ribs, mortars andtool handles, and as firewood and for charcoalproduction. The whitish fruit pulp is edible; arefreshing drink is made by maceration of thefruit pulp in water with sugarDigitum orientole Baker f. (also called 'inpepe-ta' in Swahili) is a small tree up to 12(-18) intall with short hole, occurring in dry evergreenforest and wooded grassland in southern Soma-11a, eastern Kenya and north-eastern Tanza-nia. The heavy (density about 930 kg/ms at12% moisture content), hard, reddish brownwood is used for poles in construction, lo1nery,furniture, boat ribs, mortars and tool handles,and as firewood and for charcoal productionThe whitish fruit pulp is edible; a refreshingdrink is made by maceration of the fruit pulpin water with sugar, and the pulp is also used

for flavouring porridge and beer.In Madagascar 3 Digitum spp. are found, andthe wood of anthese is used. Dialium ingdo-

gascoriense Balll. is a medium-sized tree up to30(-40) in tallwith hole up to 100 cm in diame-ter, occurring in humid evergreen forest ineastern Madagascar. The heavy and hard woodis used in construction of houses and bridges,and for carpentry; the holes are used to makedug-out canoes. In traditional medicine, barkand roots are used in preparations to reducetension

Digitttm occidentale (Capuron) DU Puy &R. Rabev. is a medium-sized tree up to 20 in tallwith hole up to 90 cm in diameter, occurring inseasonalIy dry deciduous forest in western andcentral Madagascar. The heavy and hard woodis used for construction and carpentryDigitum unifoliolotum Capuron is an ever-green medium-sized tree up to 30 in tall withhole up to 100 cm in diameter, occurring inhumid evergreen forest in eastern Madagascar.Its wood is also used for construction.

Growth and development In Liberia treesof Digitum oubreuilleiflower from July to Oc-toher, and fruits ripen about 6 months afterflowering. The sweetish fruit pulp is eaten bymonkeys, which possibly disperse the seeds.

Ecology Digitum uubreuil!ei occurs scatteredin evergreen and moist semi-deciduous forest,up to 500 in altitude. It is occasionally found insecondary forest. It is most common on hillsand seems to avoid swampy locations. It pre-fors regions with an annual rainfall of 1500-2000 min and base-poor acid soils.

Propagation and planting Natural regen-eration is most abundant in undisturbed forest.

Seedlings have been reported to prefer shadeand mineral soils, and are often abundant inthe vicinity of parent trees. There are about4000 seeds per kg. Seeds may take 2 weeks to 3months to germinate and the germination rateis moderate

Management In forest in Liberia, the aver-age number of DidIturn trees with a bole diam-eter of more than 60 cm has been estimated at

0.1-0.3 per ha, whereas the wood volume hasbeen estimated at 0.7 mama for trees with a

hole diameter of more than 50 cm. In Ghana

Digitum uubreuilleiis locally quite commonHarvesting The minimum hole diameter

allowed for harvesting in Ghana is 70 cm.Handling after harvest Conversion of the

logs after harvesting is not easy because of thehardness and high silica content of the woodSpecial equipment is needed including band-

saws with stellite-tipped teeth. Even then,sawing operations give much waste and coin-paratrvely low yields of sawn wood.

Genetic resources Digitum uubreuil!at seems

tolrly common in at least some parts of itsrange, e. g. in Liberia and Ghana. Although itoften occurs scattered and seems to prefer un-disturbed forest, there are at present no indica-tions of threats by genetic erosion. However,with the ongoing logging offorest in West Mri-ca, it may become threatened in the near fu-ture

Prospects Diolium aubreuil!ei has been sug-gested as a substitute of azob6 (LOphtr0 o10toBanks ex P. Gaertn. ) because the wood is suita-ble for heavy construction for exterior use, butthis seems to be unrealistic because azob6 tim-

her is more widely available and has superiorcharacteristics. Digitum oubreuil!ei does not

seem to have good prospects as a timber tree ofcommercial importance with sustainable pro-duction because it does not regenerate easily indisturbed forest.

The taxonomy of DidIturn in mainland Africa isquite confusing, and a revision is badly needed

Major references Aubr6ville, 1959b; Burkill,1995; de in Mensbruge, 1966; Dudek, F6rster &missenbauer, 1981; Hawthorne, 1995; Holm-gren at a1. , 2004; 0teng-Amoako (Editor), 2006;Roj0, 1982; Taylor, 1960; Voorhoeve, 1979.

Other references Brenan, 1967; Brummittat a1. , 2007a; Bryce, 1967; Chikamai at al. , un-dated; CIRAD Forestry Department, 2008; duPuy at a1. , 2002; Fouarge, Quoilin & Roosen,1970; Fouarge, Sacr6 & Mottet, 1950; Haw-thorne & Jongkind, 2006; Lovett at a1. , 2007;Maundu & Tengn6. s (Editors), 2005; Normand& Paquis, 1976; Okai, 2001; Raponda-Walker& Sillans, 1961; Ruffo, BITnie & Tengn6. s, 2002;Sallenave, 1955; Sallenave, 1964; Vivien &Faure, 1996

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

.

DIALiuM 291

Dioliumpochyphyllum - wild

Uses In Gabon the wood, known as 'omvong',is used for posts in house building and in DRCongo for turnery. It is suitable for heavy con-struction including hydraulic works, heavyflooring, joinery, interior trim, mine props, shipbuilding, vehicle bodies, furniture, railwaysleepers, ladders, sporting goods, toys, novel-ties, vats, agricultural Implements, tool han-dles, turnery and sliced veneer. In DR Congo, itis valued for charcoal productionIn Congo ground young leaves with salt aretaken to treat cough, whereas bark macera-tions are administered for the same purpose inGabon. In DR Congo bark decoctions are ad-ministered as enema to treat sterility. Gumresin from the bark is considered poisonousand has been used in DR Congo as ordeal poi-son and to prepare arrow poison; however, inCongo it is applied to healwounds

Properties The heartwood is brown to pink-ish brown, sometimes with darker streaks andmarkings, and is distinctly demarcated fromthe yellowish white or pale brown, wide sap-wood. The grain is usually straight, sometimesslightly interlocked, texture fine. The wood isresinous. It is heavy, with a density of 890-1060 kg/in' at 12% moisture content, hard andtough. Drying may cause distortion and split-ting, and should be done with care. The rates ofshrinkage are high, from green to oven dry 5.3-5.8% radial and 10.4-11.0% tangential. It isrecommended to quarter-saw logs for best re-SUIts. Once dry, the wood is unstable in service.At 12% moisture content, the modulus of rup-ture is 172-263 N/min2 modulus of elasticity20,100-25,200 N/mm2, compression parallel tograin 73-1/6 N/min2, compression perpendicu-

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DIALIUMPACHYPHYLLUMHarms

Protologue Bot. Jah"b. Syst. 53: 468 (1915)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Sumonynrs Digitum inneronthum Achev. (19/6),

Dialium yombotoense Vermoesen (1923)Origin and geographic distribution Digit-

urn pochyphyllum occurs from south-easternNigeria east to the Central African Republicand south to northern Angola.

292 TIMBERS2

Iar to grain 4-17 N/min2, shear 10 N/min2Janka side hardness 14,000 N, Janka endhardness 14,100 N and Chalais-Meudon sidehardness 11.4-14.7

The wood is very difficult to saw due to thepresence of silica; saw blades blunt extremelyrapidly and often heat up and stain the woodIt is difficult to work with both hand and ina-

chine tools, but planes to a good finish with anice polish without the use of a filler. The woodholds nails and screws well, but pre-boring isrequired. It glues well, turns well and is suita-ble for sliced veneer. The wood is durable; it isusually resistant to fungal, dry-wood borer,termite and marine borer attacks. The sapwoodis susceptible to Lyetus attack. The heartwoodis extremely resistant to linpregnation withpreservatives. The sawdust may cause irrita-tionto mucous membranes in wood workers

Description Small to medium-sized tree upto 20(-30) in tall; hole branchless for up to 15in, usually straight and cylindrical or slightlyfluted at base, up to 90 cm in diameter, withsmall to fairly large buttresses; bark surfacescaly with small, irregular scales, yellowishbrown to reddish brown, inner bark thin, brit-

tle, yellowish white to pinkish, with a reddish,sticky exudate; crown rounded, dense, withsinuous branches; twigs with numerous lenti-cels, soon glabrous. Leaves alternate, Impari-pinnately compound with 3-5 leaflets; stipuleslinear, caducous; petiole and rachis together 8-17 cm long; petiolules c. 0.5 cm long; leafletsalternate to nearly opposite, oblong-elliptical,8-23 cm x 3-8 cm, curieate to rounded at base,acuminate at apex, leathery, glabrous, pinnate-Iy veined with about 10 pairs of lateral veinsInflorescence a terminal or axillary panicle upto 20 cm long, yellowish brown hairy. Flowersbisexual, zygomorphic, fragrant; pedicel short;sepals 5, free, triangular, c. 4 mm long, hairy;petal I, spatulate, c. 4 min long, yellowish; diskpentagonal, c. 2 mm in diameter, dark brownhairy; stamens 2; ovary superior, ovoid, sessile,hairy, I-celled, style curved. Fruit a globose toobovoid, slightly flattened pod c. 2.5 cm x 1.5cm, densely dark brown hairy, with greenishwhite pulp, sepals persistent at base, indehis-cent, I(-2)-seeded. Seed flattened ellipsoid, c. Icm long, dark brown to black

Other botanical information Digitum

comprises nearly 30 species and is pantropical.Approximately 15 species occur in mainlandtropical Africa and 3 in Madagascar, whereastropical Asia has about 7 species and tropicalAmerica one

Diolium pochyphyl!urn is close to Digitum on-golense WeIw. ex 011v. and possibly conspecific.

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; 26: intervessel pits medium (7-10 pin);(27: intervessel pits large (Z 10 ILm)); 30: vessel-ray pits with distinct borders; similar to inter-vessel pits in size and shape throughout theray cell; 42: mean tangential diameter of vesselIumina 100-200 prn; 47: 5-20 vessels persquare minimetre. Tracheids and fibres: 61fibres with simple to minutely bordered pits;66: non-septate fibres present; 69: fibres thin-to thick-walled. Axial parenchyma: 78: axialparenchyma scanty paratrachea1; 86: axialparenchyma in narrow bands or lines up tothree cells wide; 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;104: all ray cells procumbent; 1/5: 4-12 raysper mm. Storied structure: 1/8: all rays sto-ried; 120: axial parenchyma and/or vessel ele-

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merits stoned. Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells; 159:silica bodies present; 161: silica bodies in axialparenchyma cells01. Shanda, A. A. Oteng-kirioako & P. E. Gasson)

Growth and development In Gabon treesmost commonly flower from November to Jan-uary, and fruits ripen about 6 months afterflowering. The sweetish fruit pulp is eaten bymonkeys, which possibly disperse the seeds.

Ecology Digitum pochyphyllum occurs indense evergreen rainforest, up to 700 in alti-tude. It is found on well drained as well as

swampy localitiesManagement In forest in Gabon, the aver-

age wood volume ofDioliumpochyphyl!urn hasbeen recorded to be 6 ms per ha. In a transectin Equatorial Guinea, a mean density of 0.6trees with a hole diameter of more than 70 cm

has been found perhaHarvesting In Gabon the minimum hole

diameter allowed for harvesting is 70 cmGenetic resources Diolium pochyphyllum

has a fairly wide distribution area and seemsfairly common in many parts of its range. Alt-hough it often occurs scattered in the forestand seems to prefer primary forest, at presentthere are no indications of threats by geneticerosion

Prospects Although Didjinm pochyphy!Iumprovides a useful timber that is particularlyvalued for heavy construction, there is littleinformation available on many aspects. Fur-ther research on its growth, development andproper management in natural forest is war-ranted. Difficulties in sawing and working ofthe wood hamper exploitation for domesticusage

Major references Aubr6ville, 1970; Bolza &Keating, 1972; Burki11, 1995; de Saint-Aubin,1963; Neuwinger, 1996; Roj0, 1982; Tailfer,1989; Takahashi, 1978; Vivien & Faure, 1985;Wilczek at a1. , 1952.

Other references Allen & Allen, 1981; Au-br6ville, 1968; CTRAD Forestry Department,2008; Fouarge, Quoilin & Roosen, 1970; Keay,1989; Konda ku Mbuta at a1. , 2010; Neuwinger,2000; Nkeoua & Boundzanga, 1999; Pauwels,1993; Raponda-Walker & Sillans, 1961; Sal-Ienave, 1964; van Reeth & Ngom0, 1997; Wilks& Issemb6, 2000

Sources of illustration Aubr6ville, 1970;Wilks & Issemb6, 2000

Authors J. K. Mensah

DIDELOTIAIDAE J. Leonard, Oldeman & de Wit

Protologue Blumea 12(2): 227 (1964).Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Origin and geographic distribution Didelo-

tio idoe is distributed in Sierra Leone, Liberia,C6te d'Ivoire and Ghana

Uses The wood of Didelotio idoe, traded as'broutou' 'gombe' or 'bondu', is used in Liberiafor planks. It is suitable for light construction,light flooring, joinery, interior trim, ship build-ing, vehicle bodies, furniture, ladders, toys,novelties, boxes, crates, tool handles, drainingboards, turnery, veneer, plywood, hardboardand particle board

Production and international trade Trade

statistics of Didelotio idoe are not available

Other Didelotio species such as Dide!otio ofri-cong Bam. , Didelotio breujponiculoto J. Leonardand Dide!otio letouzeyi Fellegr. are also tradedas 'gombe' or 'bondu'. Cameroon exported18,000 ms of 'gombe' logs in 1997, 5000 ms in1999 and 2300 ina in 2000, and Gabon exported25,000 ina of 'gombe' logs in 1997, 8900 ina in1999, 5000 ina in 2000, 4350 in3 in 2001, 4450in31n 2002, 8900 in3 in 2003, 8150 ina in 2004and 8450 ms in 2005. However, as Didelotiuidoe is not present in these countries, theseexport figures must refer to other Didelotiospecies.

Properties The heartwood is pinkish red,turning reddish brown upon drying, and isdistinctly demarcated from the up to 7.5 cmwide, yellowish brown to reddish brown sap-wood. The grain is straight or slightly inter-locked, texture coarse and even.The wood is medium-weight, with a density of

DIDELOTiA 293

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Dide!otio idoe - wild

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294 TIMBERS2

about 650 kg/in3 at 12% moisture content. Itmust be dried with care to avoid checking andsplitting, but mild schedules give excellentresults. It takes 5-6 days to kiln dry boards of2.5 cm thick to 12% moisture content. The

rates of shrinkage are moderately high, fromgreen to oven dry 4.2% radial and 8.1% tangen-tial. Once dry, the wood is moderately stable inservice. At 12% moisture content, the modulusof rupture is 107 N/mm2, modulus of elasticity12,400 N/mm2, compression parallelto grain 55N/min2, Brinell side hardness 19 N/mm' andBrinellendhardness 39 N/mm'

The wood worksfairly wellwith both hand andmachine tools. It planes to a good finish ontangential surfaces, but for radial surfaces areduced cutting angle of 20'1s recommended toprevent picking up due to the presence of inter-locked grain. A filler is necessary to obtainnicely polished surfaces. The wood holds nailsand screws well, and glues satisfactorily. Thewood has good moulding properties and theslicing and peeling properties are satisfactory;sliced veneer is decorative. The heartwood is

only moderately durable, being susceptible toattacks by fungi, termites and pinhole borersThe sapwood is susceptible to Lyetus attacksThe heartwood is resistant to impregnationwith preservatives, but can be treated satisfac-tonly by pressure; the sapwood is permeable.Tests showed that the wood is suitable for kraftpulp

Adulterations and substitutes Didelottoidoe can be substituted with other Didelotiospecies, such as Didelott0 o17icono, Didelotio67euiponiculuto and Didelotio letouzeyi. It hasbeen traded in mixtures with 'sipo' (Errtun-drophrogmo uti!e (Dawe & Sprague) Sprague)timber

Description Evergreen, medium-sized tovery large tree up to 55 in tall; hole branchlessfor up to 27 in, straight, cylindrical, up to 150cm in diameter, without buttresses but withslightly swollen base; bark surface smooth orslightly rough, with horizontal ridges, grey-green to dark brown, inner bark hard, fibrous,pink to bright red turning brown upon expo-sure, exuding a sticky, red gum; crown deltoid,fairly open, with ascending branches; twigsslender, hairy but becoming glabrous. Leavesdistichously alternate, with a single leaflet;stipules narrowly ovate, 2-5 mm long, withbifid apex, caducous except for the base; petioleand petiolule together 2-6 min long, with 2caducous, I-2 mm long stipels at the transi-tion; leaflet ovate to ovate-oblong or elliptical,

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(2-)6.5-10(-22) cm x (1.5-)4-6(-11) cm, basecurieate to rounded, apex usually acute, mar-gins entire, leathery, glabrous, pinnatelyveined with 6-8 pairs of lateral veins of which2-3 pairs from near the base of the leaflet. In.norescence an axillary or terminal, pendulouspanicle 6-15 cm long, with branches c. 2 cmlong along a slender rachis, densely hairyFlowers bisexual, nearly regular, 5-meTous;pedicel slender, 4-9 min long, at apex with 2broadly elliptical to ovate, concave, brownishred bracteoles c. 4 mm x 4 mm; sepals free,broadly ovate, c. I mm x I min, hairy at mar-gins; petals free, linear, I-2 mm long; stamensfree, (6-)8-12 mm long, pale purple, risingfrom between disk lobes; ovary superior, slight-Iy rectangular, I-celled, slightly hairy, style 6-8 min long. Fruit an oblong pod up to 12 cm x 5cm, flattened, pointed at apex, with I-2 longi-tudinal veins, smooth, yellowish brown, dehis-cent with 2 thin-woody, curling valves, few-seeded. Seeds disk-shaped or elliptical, I-2 cmx I-1.5 cm, flattened, smooth, dark brown.Seedling with epigeal germination; hypocotyl

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5-10 cm long, epicoty1 4-6.5 cm long, hairy;cotyledons thick and fleshy, c. 12 mm x 6 mm,sessile; first 2 leaves opposite.

Other botanical information Didelotiocomprises about 10 species, with about 7 spe-cies in Central Africa and 3 in West Africa. It is

particularly characterized by its 5 fertile sta-mens. The wood of several other Didelotio sppis used for similar purposes as that of Dide!otioidoe

Didelotio of neono Baill. is a medium-sized tolarge evergreen tree up to 40 in tall, with astraight hole branchless for up to 25 in and upto 100(-200) cm in diameter, distributed inNigeria, Gameroun, the Central African Re-public, Equatorial Guinea and Gabon. The palereddish brown or pinkish wood is suitable forconstruction, light flooring, joinery, interiortrim, vehicle bodies, furniture, sporting goods,musical instruments, toys, novelties, carvings,pattern making, turnery, veneer and plywood.Dide!oti0 o121e!it Taub. is an evergreen, small tomedium-sized tree up to 25 in tall with a holeup to 60 cm in diameter, distributed in Guinea,Sierra Leone, Liberia and Ghana, but it hasalso been recorded from Cameroon and Gabon.In Sierra Leone the reddish brown, moderatelyhard and heavy wood has been used for makingboards for the 'warn' game and is used formaking charcoal; seeds have been used ascounters for the warn game. A decoction ofyoung leaves is applied as a remedy for pilesDidelotto byeuiponiculoto J. Leonard (synonymTOMboot, ote 67euiponiculoto (J. Leonard) Au-br6v. & Fellegr. ) is an evergreen, large to verylarge tree up to 55 in tall with a straight, cy-11ndrical hole branchless for up to 25 in and upto 150 cm in diameter, distributed in Liberia,C6te d'Ivoire, Cameroon, Equatorial Guinea,Gabon and Congo. The reddish brown, medi-urn-weight wood, with a density of 640-750kg/ms at 12% moisture content, is suitable forconstruction, light flooring, joinery, interiortrim, ship building, vehicle bodies, furniture,ladders, sporting goods, toys, novelties, boxes,crates, tool handles, draining boards, turnery,veneer, plywood, hardboard, particle board andpulpwood.Didelotio letouzeyi Fellegr. is a medium-sizedto large tree up to 40(-60) in tall, with astraight and cylindrical bole branchless for upto 25 in and up to 150 cm in diameter, distrib-uted in Gamero0n, Equatorial Guinea, Gabonand Congo. The reddish pink, medium-weightwood, with a density of 600-700 kg/ina at 12%moisture content, is suitable for light construc-

tion, light flooring, joinery, Interior trim, shipbuilding, vehicle bodies, sporting goods, toys,novelties, boxes, crates, draining boards, turn-ery, veneer, plywood, hardboard, particle boardandpulpwood

Anatomy Wood-anatointcal description (IAWAhardwood 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 pin); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; (42: mean tan-gentialdiameter of vessellumina 100-200 prn);43: mean tangential diameter of vesselluminaa 200 pin; 46: s 5 vessels per square minimetre;(47: 5-20 vessels per square minimetre). Tra-cheids and fibres: 61: fibres with simple to ini-nutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled. Axialparenchyma: 79: axial parenchyma vasicentric;80: axial parenchyma allform; 81: axial paren-chyma lozenge-allform; (83: axial parenchymaconfluent); 92: four (3-4) cells per parenchymastrand; 93: eight (5-8) cells per parenchymastrand. Rays: 96: rays exclusively uniseriate;106: body ray cells procumbent with one row ofupright and/or square marginal cells; 107: bodyray cells procumbent with mostly 2-4 rows ofupright and/or square marginal cells; 1/5: 4-12rays per min. Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells; 143:prismatic crystals in fibres.(R. Shanda, P. E. Gasson & E. A. Wheeler)

Growth and development Natural regen-eration of Didelotio idoe is often abundant,with the seedlings tolerating dense shade. InSierra Leone flushes of young leaves appear inAugust-September, at the peak of the rainyseason. Flowering is in May-June, but does notoccur annually; fruits are ripe in August-November. In Liberia flowering has been rec-orded in the rainy season, in May and August,and fruits have been found in October-

January. Didelotio tdoe does not regrow fromstumps

Ecology Dide!otio idoe occurs in evergreenforest, in valleys and on slopes. It is also foundin coastal thickets. It occurs from sea-level upto 350 in altitude, and its occurrence Increaseswith rainfallup to about 2600 min per year. InSierra Leone and Liberia it does not grow inmarshy locations and near water, but in Ghana

DIDELOTIA 295

296 TIMBERS2

it is mostly found on riverbanks and wet partsof evergreen forest. In Liberia it occurs scat-tered, often in evergreen forest dominated byTetroberlinio tubmoniuno J. Leonard, and den-sities of over 40 trees with a diameter over 60cm per kin2 have been recorded. In Sierra Leo-ne it is locally gregarious and even dominant,but in Ghana it is fairly rare

Propagation and planting The 1000-seedweight is about 670 g. The germination rate isusually high. In Sierra Leone seeds sown inJanuary gave 90% germination in 15 daysInitial growth in the nursery is often slow, withthe seedlings only being 30 cm tallafter a year.

Genetic resources Didelotio idoe is includ-ed in the IUCN Red List, where it is consideredto be at lower risk although near threatenedHowever, its status needs revision. It is locallycommon, but human activities such as mining,logging and commercial forestry have caused asignificant decrease in the extent of its occur-

Origin and geographic distribution Dilo-beto thouorsii is endemic to Madagascar,where it is widespread in the eastern parts ofthe island and occurs locally in the central re-gions.

Uses The wood, most commonly known as'vivaona', Is widely used in house construction,and for joinery and railway sleepers. It waspopular for parts of wagons such as hubs, fel-10es and spokes. It is suitable for hydraulicworks in fresh water, mine props, vehicle bod-Ies, furniture, sporting goods, toys, novelties,agricultural Implements and turnery. It is notrecommended for flooring because of its highshrinkage rates. Branches are used as firewoodand the wood produces a good charcoal.Oil is extracted from the seeds. It is used ascooking oil, but only in times of scarcity of reg-ular cooking oil because it has a slightly disa-greeable taste. It is more commonly used ascosmetic, in soap production and for illumina-tion. The bark is used in traditional medicine.It has emetic properties and bark decoctionsare administered to treat gonorrhoea, and asdiuretic and vermifuge

Production and international trade The

wood of Dilobeio thouorsiiis commonly usedlocally, but not traded on the internationaltimber market

Properties The heartwood is yellowishbrown to pale reddish brown and indistinctlydemarcated from the narrow and slightly palersapwood. The grain is straight to slightly inter-locked, texture coarse and rather unevenFresh wood is somewhat oily to the touchThe wood is heavy, with a density of 940-1070kg/ina at 12% moisture content. It air driesslowly and great care is needed to avoid exces-

rence

Prospects The wood of Didelotio idoe hasgood strength properties and is decorative, andcan be used for a wide range of purposes. Itsdurability Is relatively low, however, making itless suitable for exterior uses. It is not knownto what extent the wood of Didelotio adoe iscurrently commercially exploited and exported,but the wood of other Didelotio species withsimilar properties is exported from tropicalAfrica

Major references Bolza & Keating, 1972;Burki11, 1995; Fox, 1968; Gottwald at a1. , 1968;Hawthorne & Jongkind, 2006; Oldeman, 1964;Poorter at a1. , 2004; Savill & Fox, 1967;Takahashi, 1978; Voorhoeve, 1979.

Other references Allen & Allen, 1981;anonymous, 1988; ATIBT, 2002; ATIBT, 2005;Aubr6ville, 1968; Aubr6vil}e, 1970; Chudnoff,1980; CIRAD Forestry Department, 2008; deSaint-Aubin, 1963; Doumenge, 1992; G6rard ata1. , 1998; Hawthorne, 1990; Hawthorne, 1995;Hawthorne & Gyakari, 2006; Leonard, 1996a;Sallenave, 1964; Vivien & Faure, 1985; Wilks& TSSemb6, 2000; World Conservation Monitor-ing Centre, 1998

Sources of illustration Voorhoeve, 1979Authors E. A. Obeng

DILOBEiATHOUARSllRoem. & Schult.

Protologue Syst. veg. 3: 476 (1818).Family ProteaceaeChromosome number2n= 52

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SIve distortion and splitting. It is recommendedto quarter-saw logs before drying. The rates ofshrinkage are high to very high, from green tooven dry 6.8-9.5% radial and 9.8-15.0% tan-gential. Once dry, the wood is unstable in ser-vice. At 12% moisture content, the modulus ofrupture is 196-255 N/mm2, modulus of elastici-ty 18,030-22,850 N/min2, compression parallelto grain 82-104 N/min2, shear 7-11 N/mm2cleavage 20-32 N/min and Chalais-Meudonside hardness 6.9-11.4.

The wood is rather difficult to saw and work

because of its high density; machine tools areneeded. It can be planed to a smooth finish andpolishes well. The nailing and screwing proper-ties are moderate; the wood is liable to splitand pre-boring is needed. The use of a filler isrecommended to obtain a good finish. The glu-ing properties are usually satisfactory andturning properties good. The wood paints satis-factorily, and is not suitable for slicing andpeeling. It is fairly durable, being moderatelyresistant to fungi and resistant to termites, butsusceptible to marine borer attack. The sap-wood is liable to Lyetus attack. The heartwoodcan be treated with preservatives under pres-sure with satisfactory results.Wood dust may Cause Irritation to eyes andmucous membranes; this is reportedIy due tothe presence of a whitish irritant substance inthe vessels

The seeds contain 60-65% of a brownish yellowoil with a characteristic strong smell that dis-appears rapidly upon cooking. The oil has asolidification point of 15'C. The main fattyacids are oleic acid (about 72%), stearic acid(12%) and palmitic acid (6%)

Description Dioecious, evergreen, small tomedium-sized tree up to 20(-30) in tall; holebranchless for up to 15 in, usually straight, upto 80 cm in diameter; bark surface scaly, red-dish brown, inner bark reddish brown withblackish spots; branches thick, with distinctleaf scars, twigs densely reddish to greyishshort-hairy. Leaves arranged spiralIy near tipsof branches, simple; stipules absent; petiole upto 11 cm long; blade heart-shaped, 7.5-20(-25)cm x 5-15(-19) cm, curieate at base, with 2large, rounded lobes at apex, with a gland be-tween the lobes at the end of the inIdrib, Ieath-ery, glabrous, inIdrib branching into 3 mainveins at 1.5-5 cm from the base, finer venationdistinct at lower leaf surface. Inflorescence

axillary, densely short-hairy, male one a many-flowered panicle up to 15 cm long, female one ashort raceme up to 6 cm long. Flowers unisex-

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ual, regular, with 4 free elliptical-oblong tepals4-6 mm long, hairy outside, caducous; maleflowers sessile, with 4 free stamens oppositethe tepals and rudimentary ovary; female flow-ers with short pedice1, 4 rudimentary stamensand superior, globose, hairy, I-celled ovary 2.5-4 mm in diameter, style very short, stigmalarge and deeply 2-lobed. Fruit an ellipsoiddrupe 3-4 cm x 2-2.5 cm, with 4 low ridges,pale brown short-hairy but becoming glabrouswhen mature, with thin layer of fleshy pulpand bony stone, I-seeded. Seed with thick, oilycotyledons

Other botanical information Dilobeio coin-

prises only 2 species, with Dilobeio tenuineruisBosser & R. Rabev. only known from forest nearTaolaiiaro. The heart'shaped persistent leavesare very characteristic for Dilobeio thouorsii

Anatomy Wood-anatomical description CAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;9: vessels exclusively solitary (90% or more);

298 TIMBERS2

13: simple perforation plates; 22: intervesselpits alternate; 24: intervessel pits minute (S 4Urn); 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 prn); 43:mean tangential diameter of vessellumina Z200 prn; 46: s 5 vessels per square minimetreTracheids and fibres: (61: fibres with simple tominutely bordered pits); (62: fibres with dis-tinctly bordered pits); (63: fibre pits common inboth radial and tangential walls); 66: non-septate fibres present; 70: fibres very thick-walled. Axial parenchyma: (76: axial paren-chyma diffuse); 80: axial parenchyma anform;82: axial parenchyma winged-allform; 83: axialparenchyma confluent; 84: axial parenchymaunilateral paratrachea1; 92: four (3-4) cells perparenchyma strand; 93: eight (5-8) cells perparenchyma strand. Rays: 97: ray width I-3cells; (98: larger rays commonly 4- to 10-senate); 104: all ray cells procumbent; 106:body ray cells procumbent with one row of up-right and/or square marginal cells; 1/5: 4-12rays per mm.

(FD. Kamala, P. Baas & P. E. Gasson)Growth and development Leaves of young

plants of Dilobeio thouorsii are quite distinctfrom those of older trees; they have 2 or 4 nar-row lobes up to 30(-50) cm long. Trees flowerfrom October to March and fruits can be found

nearly throughout the year, although theyseem to ripen only in November to January

Ecology Di!o6eio thouorsii occurs in humidevergreen forest from sea-level up to 1800 inaltitude, but most commonly at 700-1300 in. Itis locally abundant.

Harvesting Felhng the trees with their denseand hard wood requires special equipment, andthe heavy logs are not easy to transport.

Handling after harvest The fruit stonesare pounded with a mortar and the fragmentsare boiled in a small amount of water. The oil

is collected from the surface of the liquid.Genetic resources Dilobeio thouorsii is

widespread in eastern Madagascar and IOCaUycommon. There are no indications that it is in

immediate danger of genetic erosion, althoughthe splitting up of forest in its area of distribu-tion may threaten it in the near future

Prospects Dilobeio thouorsiiis valued bythe local people, mainly for its strong and du-rable wood and for its oil-rich seeds. Little is

known aboutits growth rates and propagation,and research is warranted to be able to draw

up recommendations for sustainable exploita-

tion.

Major references Bolza & Keating, 1972;Bosser & Rabevohitra, 1991; Decary, 1946;Gu6neau, Bedel & Thie1, 1970-1975; Parant,Chichignoud & Rakotova0, 1985; Ralaibi-banson, 1995; Schatz, 2001; Takahashi, 1978.

Other references Boiteau, Boiteau & A1-10rge-Boiteau, 1999; Brown at a1. , 2009;Gu6neau & Gu6neau, 1969; Kubitzki, 2007;Lanyon, 1979; Rakotovao at al. , en preparation;Sallenave, 1955; Sallenave, 1971.

Sources of illustration Schatz, 2001.Authors S. Rakotonandrasana

DIOSPYROSABYSSiNiCA (Hiern) Finlite

Protologue Bull. Jard. Bot. Etat 26: 241(1956)

Family EbenaceaeSynonyms Moboobyssinico Hiern (1873).Vernacular names Black bark, giant di-

OSpyros (En). Mueluili, indaa-inwitu (Sw).Origin and geographic distribution Diospy-

ros o6yssinico is widespread, occurring fromGuinea and Mali eastward to Eritrea, Ethiopiaand Kenya, and south to Angola, Zimbabweand Mozambique

Uses The wood of Diospyros o6yssinico isused for heavy flooring, poles, interior trim,mine props, furniture, cabinet making, mastsof dhows, agricultural implements, musicalinstruments, tool handles, ladders, toys, novel-ties, pestles, mortars, golf club heads, sticks,carving and turnery. It is also in demand forloom shuttles in weaving sisal cloth in KenyaThe wood is commonly used as firewood and forcharcoal production.

Diospyros obyssinico - wild

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Various parts of the plants are used in tradi-tional medicine. In Ghana bark and roots are

used by the Krobo people against various dis-eases. In Malileaf and roots decoctions are

used to treat malaria and dysentery, and topromote wound healing. In Tanzania a rootdecoction Is taken to treat leprosy.

Production and international trade In Tan-

zania the wood is considered suitable for the

handles of pick-axes and other tools. A yearlyproduction of up to 20,000 axe handles hasbeen recorded in the 1960s

Properties The heartwood is whitish yellowto pale grey-brown, often with irregular blackstreaks or entirely black in the centre; it is notdistinctly demarcated from the wide, whitish toyellowish sapwood. The grain is generallystraight, sometimes interlocked, texture usual-Iy fine. Freshly cut wood has an unpleasantsmell

The wood is hard, tough and moderately heavywith a density of 720-860 kg/in" at 12% ino1s-ture content. The rates of shrinkage are inod-erately high, from green to 12% moisture con-tent 3.7% radial and 6.6% tangential. The woodair dries satisfactorily, with little distortion orsplitting, but with a risk of severe cupping inover 5 cm thick boards. Boards of 5 cm thick

take about 5 months to air dry. The wood isliable to distortion when kiln dried and low

temperature schedules are recommended. At12% moisture content, the modulus of ruptureis 1/4-1/6 N/min2, modulus of elasticity 10,800N/min2, compression parallel to grain 51-55N/min2 shear 17 N/mm2 and Janka side hard-

ness 5380 N.

The wood saws well, but considerable power isrequired and packing of large amounts of wetsawdust around the rim should be avoided. The

wood works satisfactorily with hand tools. Thewood planes well with standard 30' cuttingangles, but for quarter-cut material with Irreg-ular grain smaller cutting angles are recoin-mended to avoid tearing and picking-up. Thewood moulds and sands satisfactorily. Nailingis difficult, and pre-boring is recommendedMortising and boring are somewhat difficult;boring causes some checking at the exits. Thewood has a low durability and is susceptible toattacks by fungi and pinhole borers and inod-erately susceptible to termite attack. The ser-vice life in contact with the ground is only 8-12months. The wood is moderately resistant toimpregnation with preservatives. The finesawdust may cause dermatitis and irritation tonose and eyes, and the use of a dust extractor

is recommended

Extracts of the root bark are potent inhibitorsof 15-lipoxygenase and showed distinct radicalscavenging activity. The triterpenoids betulin,betulinic acid and Iupeol have been isolatedfrom Diospyros o6yssinico. These compoundsare known to have anti-inflammatory activity.The bisnaphthoquinones diospyrin and isodio-spyrin have been isolated from the bark. Theyshowed activity against protozoan parasitesand tumours in mice, as well as inhibition ofblood platelet aggregation.

Description Evergreen, dioecious, small tofairly large tree up to 35(-40) in tall; bolebranchless for up to 18 in, straight and slender,up to 60(-75) cm in diameter, often with smallbuttresses; bark surface smooth in young trees,later becoming rough and scaling off in stripsor oblong plates, grey to dark green, darkbrown or black, inner bark thin, fibrous, yel-low-brown, darkening upon exposure; crownoften small and narrow, with branches at rightangles to main stem; twigs pendulous and usu-ally glabrous. Leaves alternate, simple andentire; stipules absent; petiole 4-8 min long;blade elliptical to oblong-elliptical, 3-15.5 cm x

DIOSPYROS 299

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I-5.5 cm, curieate to rounded at base, obtuse toslightly acuminate at apex, thinly leathery,shining, nearly glabrous, pinnately veined with5-12 pairs of lateral veins. Inflorescence inaxils of leaves or on older branchlets, in maletrees a 10-18-flowered contracted Cyme, In

female trees a (I-)3-5(-8)-flowered fascicle.Flowers unisexual, regular, 3-4-merous,creamy white to yellowish, scented; pedice1 I-2min long; male flowers with cup-shaped calyx c.2 mm long, corolla 5-6 mm long, deeply lobedwith lobes broadly elliptical, stamens 10-15, 2-4 mm long, ovary rudimentary; female flowerswith cup-shaped calyx c. 6 mm long, dividednearly to the base, corolla similar to that ofmale flowers, rudimentary stamens 3-4, ovarysuperior, conical, c. 4 min long, 6-celled, styleshort. Fruit an ellipsoid to nearly globose berry8-14 mm x 8-9 min, glabrous, yellow to or-ange, with persistent calyx at basis and styleat apex, IC2) seeded. Seed globose to ellipsoid,c. 9 min x 6 min, with smooth endosperm

Other botanical information Diospyros isa large pantropical genus of about 500 species;in tropical Africa about 90 species occur andseveral produce valuable timber or ediblefruits. There Is quite some variation in theshape, size, venation and indumentum ofleaves over the enormous eco-geographicalrange of Diospyros o6yssinico, and some of thisvariation resulted In the recognition of 4 sub-species. Subsp. o6yssintco covers the wholedistribution area of the species, the other 3subspecies are restricted to small regions inEast and southern Africa.

Anatomy Wood-anatomical description CAWAhardwood 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-nate pits polygonal); 25: intervessel pits small(4-7 prn); 30: vessel-ray pits with distinct bor-ders; similar to intervessel pits in size andshape throughout the ray cell; (40: mean tan-gential diameter of vessellumina s 50 prn); 41:mean tangential diameter of vessellumina 50-100 ILm; 48: 20-40 vessels per square minime-tre; (49: 40-100 vessels per square minimetre);58: gums and other deposits in heartwood ves-sels. Tracheids and fibres: 61: fibres with SIm-

PIe to minutely bordered pits; 66: non-septatefibres present; 70: fibres very thick-walled.Axial parenchyma: 77: axial parenchyma dif-

fuse-in-aggregates; 78: axial parenchymascanty paratrachea1; 86: axial parenchyma innarrow bands or lines up to three cells wide;92: four (3-4) cells per parenchyma strandRays: 96: rays exclusively urnseriate; (97: raywidth I-3 cells); 106: body ray cells procum-bent with one row of upright and/or squaremarginal cells; 107: body ray cells procumbentwith mostly 2-4 rows of upright and/or squaremarginal cells; 116: z 12 rays per min. Stonedstructure: (1/8: all rays stoned); 122: raysand/or axial elements irregularly stoned. Min-eralinclusions: 136: prismatic crystals present;142: prismatic crystals in chambered axial pa-renchyma cells.(E. E. Mwakalukwa, P. Baas & H. Beeckman)

Growth and development In generalDios-pyros o6yssinico is reported to grow slowly, butin Uganda it seems to be able to grow fasterwhen planted in farmland. Trees have a fullcrown of leaves through the dry season, butmost individuals drop their leaves during thewet season when new flushes appear. Fruitsripen about 3 months after flowering. They areeaten by various animals, such as baboons inGhana, bulbuls in Malawi and hornbills inZimbabwe; these may serve as seed dispersers.

Ecology Diospyros o6yssinico can be foundin a wide dlversity of forest types, and also inwoodland and thickets. In its entire distribu-

tion area the altitudinal range is 200-2500 inand the mean annual rainfall range 650-2050mm. However, its most characteristic habitatismoist semi-deciduous forest with well. dis.tributed rainfall between 1250 and 1500

min/year. Its presence in drier areas dependson the availability of water along watercourses,from runoff from rocky outcrops, or in water-retaining soils of termite mounds

Propagation and planting Usually Dios-pyros o6yssinico is considered a pioneer speciesthat is particularly characteristic for the earlystages of forest succession, but may persist inold forest and may even regenerate there undershady conditions. However, light is reported tohave a positive influence on seed germinationand seedling growth. Seedlings are often coin-mon close to mother trees, but mortality is con-sidered to be high.Diospyros o6yssintco is only propagated byseeds. One kg contains 2500-3000 seeds. Ger-ruination is slow. In a test in Kenya, 12% offresh seeds had germinated after 3 weeks and20% after 7 weeks. Seeds can be stored for sev-

eral years if they are protected from Lnsects.Ash is added to reduce insect damage. Pre-