rable to that of pulps made from softwoods. Itcould be used as a substitute for softwood pulpand for flexible packaging paper grades, eitheralone or in blends with other fibre sources

Bombuso uulgoris can also be successfully usedas a raw material for particle board manufac-ture

Young shoots are whitish pink in colour andtender, and they have a fair canning qualityThe approximate chemlcal composition ofyoung shoots per 100 g edible portion for green-stem and yellow-stem cultivars respectively is:water 90 g and 88 g, protein 2.6 g and 1.8 g, fat4.1 g and 7.2 g, carbohydrates 0.4 g and O g,fibre 1.1 g and 1.2 g, Ca 22.8 ing and 28.6 ing,P 37 ing and 27.5 ing, Fe 1.1 ing and 1.4 ing,ascorbic acid 3.1 ing and O ingIngestion of large amounts of leaves has beenrecorded to cause neurological disorders inhorses, but this was not related to poisoning byhydrocyanic acid. A chloroform extract of theleaves is active against Mycobocterium tuber-CUIosis. The presence in the plant of taxiphyl-Iin, a cyanogenic glucoside, has been reportedStigmasterolhas been isolated from Arthroboc-ter globiformis fermented shoot.

Description Bamboo with a short, thick rhi-zome and not closely tufted stems; stem (CUIm)erect, sinuous or slightly zig-zag, up to 20 intall, up to 12 cm in diameter, hollow, wall(3-)7-16(-20) min thick, glossy green, yellow, oryellow with green stripes, internodes 20-45 cmlong, with appressed dark hairs and whitewaxy when young, becoming glabrous, smoothand shiny with age, nodes oblique, slightlyswollen, basal ones covered wlth aerial roots;young shoots dark brown to yellow-greenLeaves alternate, simple; stem leaves withsheath more or less broadly triangular, 15-45cm x 17-35 cm, covered with appressed blackor brown hairs, auricles 2, 0.5-2 cm long, withpale brown bristles 3-8 min long along theedges, 11gule 3-8 mm long, slightly toothed,blade broadly triangular, 4-15 cm x 5-10 cm;branch leaves with sheath 6-10 cm long, hairyoutside, auricles rounded, 0.5-1.5 mm long,with a few bristles I-3 min long, Iigule 0.5-1.5min long, blade linear-lanceolate, 6-30 cm x I-5 cm, base rounded, apex acuminate, marginscabrous, glabrous. Inflorescence a large pani-cle 2-3 in long, on leaness branches, bearingsmall groups of spikelets at the nodesSpikelets 12-20(-35) min x 3-6 mm, lateralIyflattened, comprising I-2 glumes, 4-12 floretsand a terminal vestigialfloret, glumes ovate,acute, lemma ovate, 9-11 mm long, 11-15-

I I. \,6

BAMBUSA 101

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21

Bombuso uulgoris - I, young shoot, . 2, stem led/:3, leg/y 670nch, . 4, upper port of Ieof sheerth, . 5,flowering brunchlet, ' 6, spikelet.Source. PROSEA

veined, palea as long as lemma, 2-keeled, 6-veined; norets with 6 exserted stamens, 3 un-equal}odicules, ovary narrowly oblong, stylelong, thin, hairy, terminating in 2-3 stigmasFruit not developing

Other botanical information Bombuso

comprises 120 species indigenous to Asia andthe Americas. Bombuso uu!goris is the mosteasily recognized species of allknown bamboos.In a young stem, the primary branches areprominent; they are borne alternately alongthe stern, together forming a gigantic fan-likestructure which is conspicuous from some dis-tance. The vanability within Bombuso uulgorisis great and a thorough study of the worldwidevariation is necessary. Classification of thedifferent forms should preferably be done atcultivar level (cultivar groups, cultivars). Atleast three groupings of cultivars can be distin-guished:- Plants with green sterns.- Plants with yellow stems ('golden bamboo'),

often with green stripes of different inten-sity. Usually the stems have thicker wallsthan those of the green stern group. Thisgroup is often distinguished as Bombuso

..

if '11

5

4

102 TIMBERSl

strioto Lodd. ex Lindl.

- Plants with stems up to about 3 in tall, I-3cm in diameter, green, with 4-10 cm long in-flated internodes in the lower part ('Bud-dha's belly bamboo'). This group is often dis-tinguished as Bombuso worntn Brandis exEGGamus

Bombuso bombos (L. ) Voss (spiny bamboo;synonym: Bombuso urundinoceo (Retz. ) Wind. )is native to India to southern China, and culti-vated throughout the tropics, including Ghanaand Kenya. It has stems up to 30(-40) in tallwith a diameter up to 18 cm. In Ghana thestems are used for construction. In South-East

Asia they are important raw material for thepaper making industry

Growth and development Bombuso uul-guns clumps expand rapidly during the first 5-6 years (from 0.5 in diameter in the first yearto 4.5 in in the 6th year) and slower thereafter(to 7 in diameter after 10 years). Young shootsgrow rapidly. In 2 weeks they can develop intostems 3-4 in tall, reaching 20 in in length in 3months. Sterns reach maximum diameters af-

ter 9 years. The number of young shoots perclump that develop into full-grown stems in-creases on average from 1.6 in the first year toa maximum of 5.3 in the 4th year and decreasesto 2.5-3.5 from the 9th year onward. On aver-age, a mature clump produces 3-4 new stemsper year and bears 30-90 sterns. In the Niarivalley in Congo (average annual temperature25.5'C, average annual rainfall around 1000mm), 4.5-year-old Bombuso uulgoris planted ata spacing of 6 in x 6 in, with surviving densityof 226 plants/ha, on average had 31 stems perclump and 7000 stems per ha. Flowering isuncommon in Bombuso uulgoris. When a stemflowers, it produces a large number of flowers,but no fruits. Low pollen viahinty due to Ir-regular meiosis seems to be one of the reasonsfor the absence offruiting. Eventually the stemdies, butthe clump usually survives.

Ecology Bombuso uulgoris grows best atlower altitudes; above 1000 in altitude stemsbecome smaller in length and diameter. Itthrives under a wide range of moisture and soilconditions, growing in almost permanentlyhumid conditions along rivers and lakes, butalso in areas with a severe dry season, wherethe plants may become completely detonatedIt is frost hardy down to -3'C.

Propagation and planting Bombuso uu!-guns is easy to propagate vegetativeIy, byclump division, by rhizome, stem and branchcuttings, by layering and by tissue culture

Rhizome cuttings (offsets) give good resultswhen taken from I-2-year-old stems, but cut-ting damages the mother clump and rhizomecuttings are not convenient for large-scaleplantations. The easiest and most practisedpropagation method is by stem or branch cut-tings. In the Philippines best results have beenobtained with one' node cuttings from the lowerparts of 6-month-old stems, planted horizon-tally in moist soil at about 20 cm depth. Treat-ing cuttings with a 100 ppm solution of thegrowth hormone indole butyric acid improvedresults. Planting Is recommended during thelate rainy season and at a spacing of 6-12 in x6-12 in. In C6te d'TVoire 2-node cuttings arerecommended, planted obliquely in the soilwith the upper part emerging. The easy vege-tative propagation of Bombt, so uulgoris ex-plains its often apparently wild occurrence:clumps may establish from pieces of stemsused for poles, fences, props, stakes and postsset on river banks for mooring boatsSuccessful in-vitro regeneration procedures forBombuso uu!guns have been developed usingnodal segments or buds on Murashige andSkoog medium

Management Weeding is recommended inthe first 2-3 years after planting, 2-3 times peryear, preferably during the rainy season. Irri-gation and fertilizer application considerablyimprove growth and yield. Protection fromanimals is Important, because they readilybrowse the young shoots. The plants shouldalso be protected from fire. Where Bombusouulguris is planted at close spacings to coin-pensate for young plants dying, the first thin-rimg is carried out at an age of 5-6 years, andthe final density is established at 12-13 yearsage. In Tanzania management of plantedBombuso uu!goris mainly consists of clearingthe ground around the clumps.

Diseases and pests Diseases of BombusoDMZgoris include leaf blight (Cercosporu sp. ),basal CUIm rot (Fusorium sp. ), CUIm sheath rot(Glomerello cmguloto), leaf rust (Kweilingiodiuino, synonym: Dusture!IQ drumo) and leafspots (Doctylorto sp. and Glomere!IQ cmgu-loto). A serious disease in Bangladesh is barn-boo blight caused by 8070clodium oryzoe, kill-ing affected clumps within 3-4 years, Bambooblight especially occurs on sites with impededdrainage. It can be controlled by cultural prac-tices such as removal of infected sterns and

burning of debris, and by fungicides.Harvested stems are very vulnerable to attackof powder-post beetles (Dinoderus spp. ). Ter-

I,

mite damage can be serious, especially of har-vested stemsin contact with the ground

Harvesting Harvesting normally starts 3years after planting. Fullproduction is reached6-8 years after planting. Selective cutting ofstems 2-year-old or older is recommended. Intropical Africa it has been recommended toselectively harvest one half to two-thirds of theadult stems on a clump every 3-4 years. Youngshoots for consumption should be harvested inthe first week of their emergence.

Yield Yields recorded for tropical Africa are10 t dry weight per ha per year for C6ted'TVoire and 15 t for Congo. In trials in Congo,yields were higher for Bombuso uulgoris thanfor Orytenonther0 o6ysstnico (ARIch. ) MunroFor the Philippines the annual yield per ha isestimated at 2250 sterns or 20 t dry weightThe dry weight ratio for stem, branches andleaves is about 70%, 22% and 8% respectivelyThe ratio of paperpulp/stem production Isabout 1:3

Handling after harvest Post-harvesttreatment of the stern should aim at reduction

of the moisture and starch content to increase

its durability. Traditionany, harvested stems ofBombt, so uu!goris are Immersed in running ormuddy water for about 3 months. In this proc-ess, the starch content is reduced, mainly bybacterial degradation. However, the suscepti-hinty to insects and fungiis normally hardlyreduced. Other traditional practices are paint-ing with slaked lime, which reduces water ab-sorption and makes the surface become alka-line, thus delaying fungal attack, and plaster-ing of bamboo mats with a thick layer of mud,clay or sand mixed with lime or cement, whichkeeps the mats protected against rain and pre-vents the entry of insects. For safe and long-term use, however, traditional methods do notsuffice and chemical preservation is necessaryFor preservation, boron salts dissolved in wa-ter are most commonly used. Often, the stemsare simply treated by placing their base in acontainer with a preservative; complete sub-mersion of stems is also practised.Young shoots should be consumed or preparedimmediately after harvesting because theybecome triedIble within some hours if left un-

treated.

Genetic resources Bombuso uulgoris Isrepresented in most botanical gardens in thetropics. Because of the wide vanability of thisbamboo, extensive, worldwide germplasm col-Iection is recommended

Prospects Because of its easy propagation,

rapid growth, long life, high yield and wideapplicability, Bombuso uulgoris has good pros-pects as an alternative for wood in tropicalAfrica. Drawbacks, however, are the limiteddurability and poor working properties of thestem material. Structural use is not recoin-

mended without preservation to protect thestem material against insects and fungi. Born-buso uu!guns has good prospects for the pro-duction of pulp and biomassfor energy

Major references Burki11, 1994; Chihongoat a1. , 2000; Chilufya & Tengnas, 1996; CTFT,1962a; Dah-Dovonon, 2001; Doat, 1967; Drans-field & Widjaja, 1995a; Groulez, 1966; Liese,2004; Seethalakshmi& Muktesh Kumar, 1998

Other references Bathosa at a1. , 2006; CABInternational, 2005; Clayton at a1. , 1994; deCarvalho Jr, 2001; de in Mensbruge, 1968;Dunyaprapan & Jansen, 1995; Gunb-Fakim ata1. , 1994; Igoli at a1. , 2005; Kalanda & Bo-lamba, 1994; Khristova at a1. , 2006b; Koshy &Jee, 2001; Latham, 2007; Mishra & Thakur,1998; Ndiaye at a1. , 2006; Papadopoulos at al. ,2004; Rainanayake, Meemadumaa & Weer-awardenea, 2006; Rugalema, Okting'atI &Johnsen, 1994; Sarangthem, Singh &Srivastava, 1998; Sarpong, 2000; van der Zori,1992

Sources of illustration Dransfield & Wid-

iaia, 1995a.Authors M. Brink

Based on PROSEA 7: Bamboos.

BAPHiA 103

BAPHiAKiRKllBaker

Protologue 011v. , F1. trop. Mr. a 250(1871).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Vernacular names Gevire (Po). Mkuruti(Sw)

Origin and geographic distribution Bophiohirhiiis restricted to coastal regions of Tanza-nia and to southern Mozambique

Uses The wood (trade names: baphia, cam-wood)is used for furniture (especiaUy table andcounter tops), heavy duty flooring and turneryIn tropical Africa it is used for construction oflocal houses and in boat building, but also fortool handles, pestles, stools and carvings, asfirewood and in charcoal production. Bophiohirkiiis planted as an avenue and ornamentalshade tree. A root decoction is drunk to treat

epilepsy.Production and international trade In the

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104 TIMBERSl

first half of 2004 there was still considerable

export ofBophio hirhtilogs from Tanzania, butin July 2004 a ban was imposed on the exportof round wood.

Properties The heartwood is purplishbrown and contains red gum; the sapwood isyellowish white and up to 2 cm wide. The grainis straight or slightly wavy, texture fine andeven. The wood has a peppery scent. It isheavy, with a density of 1280 kg/ing at 12%moisture content, and hard. Shrinkage ratesfrom green to 12% moisture content are 1.5%radial and 2.4% tangential. Green timber has alow moisture content, and air-drying is rapidwith little degradation. The wood is difficult towork with machine tools, rapidly blunting sawteeth and cutters. It finishes and polishes verywell. It is resilient with good weathering prop-erties. The durability is high, the wood beingonly rarely attacked by marine borers and ter-mites. The sapwood is not susceptible to Lyetusborer attack. The heartwood is impermeable topreservatives

Botany Medium-sized tree up to 27 in tall;hole up to 15 in long, with a diameter up to 0.9in, irregular in form with deep nuting at thebase; outer bark grey-brown; crown rounded,much-branched; twigs pendulous, rusty brownhairy but glabrescent. Leaves alternate, simpleand entire; stipules early caducous; petiole I-3cm long, thickened at base and top; bladebroadly ovate to elliptical, 4-14 cm x 1.5-7 cm,curieate to shallowly cordate at base, obtuse toacuminate at apex, leathery, glabrous, pin-nately veined with 5-8 pairs of lateral veinsInflorescence an axillary raceme 3.5-7.5 cmlong, often grouped into panicle-like leafy inno-rescences at the ends of branches. Flowers bi-

sexual, papilionaceous; pedice1 6.5-18 mmlong, hairy; calyx 9-15 min long, 2-10bed,sparsely brown hairy; corolla with almost or-bicular standard up to 20(-25) mm in diame-ter, white with yellow blotch at base, wingsand keel white with a pocket near the base;stamens 10, free, up to 16 min long; ovary su-perlor, glabrous, style lncurved. Fruit an ellip-sold-oblong pod 7.5-16.5 cm x 2.5-5 cm, flat-tened, woody, pale brown, dehiscing with 2valves, I-2-seeded. Seeds lens-shaped, 1.5-2.5cm x I-2 cm, black or dark brown.Bophio comprises about 45 species and is re-stricted to Africa, including Madagascar where2 species are found. The distribution area ofBophio Aichiiis disjunct, the two sub-areasseparated by about 1500 kin. Plants from Tan-zania are considered to belong to subsp. Airhii,

those from Mozambique to subsp. ouoto (Sim)Soladoye (synonym: Bophio ouoto Sim), differ-ing in slightly smaller flowers and more hairycalyxThe wood of some other Bophio species is usedfor similar purposes as that of Bophio hirhit,but most species are of too small size to be jin-portant. The best known timber species isBophto nitido Lodd. (camwood), but the use ofthe red dye from its wood is more importantBophio copperridifoliu Baker belongs to thesame section as Bophio hirhii. This widespreadspecies, occurring from Guinea to western Tan-zania and Zambia and in Madagascar, is usu-ally a climbing or scrambling shrub of whichthe wood is nevertheless used for walkingsticks and as supports for fishing nets. Itsleaves are used to treat fever and are given topregnant women when the foetus develops tooslowly. The wood of Bophio IOUrifolio Baill. , ashrub or smallto medium-sized tree up to 20 intall occurring from eastern Nigeria to centralDR Congo, is used in Gabon for kitchen imple-merits

Ecology Bophio kithti occurs in coastal for-est, thickets and savanna up to 400(-900) inaltitude

Genetic resources and breeding The dis-tribution area of Bophio hirhii consists of twosmall regions and the species can easily be-come endangered with ongoing logging activi-ties. It has been classified by IUCN as unlner-able. Monitoring of the existing populations isrecommended.

Prospects There is very little informationon wood properties, growth and propagation ofBophio hirkii, although it is a locally favouredsource of timber. Research on these aspects isdesired, as well as on proper managementmethods of naturalstands

Major references Bryce, 1967; Milledge,2005; Soladoye, 1985

Other references Bolza & Keating, 1972;Gillett at a1. , 1971; Lovett & Clarke, 1998a;Lovett at a1. , 2006; Neuwinger, 2000; Raponda-Walker & Sillans, 1961

Authors R. H. M. J. Lemmens

BEILSCHMIEDIA CORBISIERl(R. obyns) Robyns& R. Wilczek

Protologue Bull. Jard. Bot. Etat 19: 468(1949).

Family LauraceaeSynonyms Ty!OStemoit corbisieri Robyns

.

I ,'

(1930), Benschmiedio megaphyllo Pierre exRobyns & R. Wilczek (1950).

Origin and geographic distribution Bet!-schmiedio corbisieriis distributed in Gamer-

o0n, Gabon and DR Congo.Uses The wood is particularly suitable for

carving and turnery, but it can also be used forInterior carpentry, construction, flooring, join-ery, furniture, cabinet work, shipbuilding,railway sleepers, sporting goods, agriculturalimplements, mine props, poles and piles

Properties The heartwood is pale red-brown; it is clearly demarcated from the pink-ish white sapwood. The grain is usuallystraight, texture medium. The wood containssome resin canals. The density is 730-800kg/ms at 12% moisture content. The wood ismoderately fissile. It seasons rather slowly,and severe splitting occurs when drying is ac-celerated. At 12% moisture content, themodulus of rupture is 137 N/min', compressionparallel to grain 52 N/min', cleavage 15.7N/min and Monnin side hardness 3.5. The

wood saws well, but slowly; it has a fairly highsilica content. It nails well and has excellent

nail-holding properties. It finishes and polisheswell, and has satisfactory working and planingcharacteristics. It is resistant to wear and

weathering. The wood is often attacked by bor-ers, but is fairly resistant to attacks by marineborers and termites. The sapwood is liable toattack by Lyetus beetles. The wood is imper-meable to preservatives

Botany Evergreen shrub or small to me-dium-sized tree up to 20 in tall; hole straight,cylindrical, up to 50 cm in diameter, withoutbuttresses; outer bark smooth; branches cylin-drical, with bark blackish grey and rough,young branches flattened and reddish hairy.Leaves alternate to almost opposite, simpleand entire; stipules absent; petiole I-2 cmlong, more or less channeUed, hairy; blade ob-long to oblong-elliptical, 6-26 cm x 3-9 cm,base curieate, apex acuminate, thinly leathery,glabrous above, sparsely hairy below, ohvecoloured above, reddish brown below, pinnatelyveined with 6-10 pairs of lateral veins. Inflo-rescence a lax and slender panicle 6-12 cmlong, axiUary near the top of young branches,reddish hairy, branches thread-like; peduncle1.5-5 cm long; bracts acute, hairy, persistentFlowers bisexual, regular, small, greenish,hairy; pedicelthread-like, 2.5-5 mm long; perl-arith cup-shaped, lobes 6(-8), broadly deltoid toovate, c. I mm long, glandular; fertile stamens9, in 3 whorls, those in outer 2 whorls sessile,

those in inner whorlstalked and with 2 glandseach, staininodes in a fourth whorl; ovary su-perlor, c. I mm long. Fruit an ovoid to oblong-ovoid berry up to 5 cm x 3 cm, glabrous, red-dish and rough, I-seededBenschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Beilschmtedio corbisieri belongs to sub-genus Synthorodenio. Benschmiedio diuersi-fjord Pierre ex Robyns & R. Wilczek belongs tothe same subgenus and is closely related toBenschmiedio corbisieri, but the leaves aresmaller with reddish curly hairs below, and theperianth lobes are shorter;It is sometimes con-sidered a variety of Benschmiedio corbisieri.

Ecology Benschmiedio corbisieri occurs inswamp forest and periodically flooded forest.

Genetic resources and breeding Bet!-schmiedio corbisieri has a limited distributionand it is unclear whether this species is threat-ened by genetic erosion. It is not included inthe 2006 IUCN Red list of threatened species.

Prospects The wood of Benschmiedio cor-histeriis considered suitable for a wide range ofapplications, but it is unclear to what extent itis presently used. Its potential is difficult toassess because too little is known about growthcharacteristics and abundance

Major references Bolza & Keating, 1972;Fouarge, Sacr6 & Mottet, 1950; Robyns &Wilczek, 1951

Other references Foullloy, 1965; Fouilloy,1974

Authors M. Brink

BEiLscHMIEDiA 105

BRILSCHMIEDIADrvERSIFLORA Pierre ex

Robyns & R. Wilczek

Protologue Bull. Jard. Bot. Etat 20: 200(1950)

Family LauraceaeOrigin and geographic distribution Ben-

schmiedio diuersi/10ro is distributed in Garner-o0n, Gabon and Congo

Uses The wood (trade names: kanda, brownkanda) is suitable for construction, flooring,interior trim, joinery, turnery, furniture, cabi-net work, shipbuilding, vehicle bodies, railwaysleepers, poles, piles, mine props, veneer andplywood

Properties The heartwood is red-brown; itis clearly demarcated from the yellowish sap-wood. The grain is straight, texture fine to me-dium. The wood contains resin canals. The

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106 TIMBERSl

density of the wood is about 730 kg/in3 at 12%moisture content. The wood seasons slowly, butwith little degrade. Shrinkage rates are 3.3-4.0% radial and 5.6-6.4% tangential fromgreen to oven dry. Once dry, the wood is stable

At 12% moisture content, theIn service

modulus of rupture is 154 N/mm2, modulus ofelasticity 9700 N/min2, compression parallel tograin 60 N/mm2, shear 8.6 N/min2, cleavage 22N/mm and Chalais-Meudon side hardness 5.2

The wood saws fairly well, but with some dull-ing effect on tools due to the presence of silicaIt nails well and has good nail and screw hold-ing properties; it works and finishes satisfacto-rily. The wood is resistant to fungi, termitesand marine borers, but the sapwood is liable toattack by Lyetus beetles. The heartwood is re-SIStantto impregnation with preservatives

Botany Evergreen medium-sized tree up to27 in tall; bole straight, branchless for up to 15in, cylindrical, up to 60 cm in diameter, with-out buttresses; young branches reddish hairy.Leaves alternate to almost opposite, simpleand entire; stipules absent; petiole c. I cm long,hairy; blade obovate to oblong, 7-14 cm x 3-6cm, leathery, olive-brown and glabrous above,reddish hairy on the veins below, pinnatelyveined with 6-8 pairs of lateral veins. Inflores-cence a terminal or axillary, robust panicle 4-8cm long, reddish hairy; peduncle I-1.5 cm long,flattened; bracts caducous. Flowers bisexual,regular, small, brownish yellow; pedice1 I-2min long, reddish hairy; perianth bell-shapedor cup-shaped, lobes 6, up to I mm long; fertilestamens 9, in 3 whorls, those in outer 2 whorlssessile, those in inner whorlstalked and with 2glands each, staininodes in a fourth whorl;ovary superior, c. I mm long. Fruit a pear-shaped berry 2.5-3 cm x c. 1.5 cm, I-seededBenschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Benschmiedio dirersi/lord belongs tosubgenus Synthorodenio. It is closely related toBetlschmiedio corbisieri (Robyns) Robyns &R. Wilczek, but the leaves of the latter are Iar-ger and sparsely hairy, and the perianth lobesare slightly longer. Benschmiedio diuersiftorois sometimes considered a variety of Betl-schmiedio corbisieri

Ecology Benschmiedio diuersi/1070 occurs inmixed forest, mainly in humid lowland areas.

Genetic resources and breeding Beil-schmiedio diuersi/1070 has a limited distrihu-tion and it is unclear whether this species isthreatened by genetic erosion. It is not in-

cluded in the 2006 IUCN Red list of threatened

species.

Prospects The wood of Benschmiedio diuer-sinoro is considered suitable for a wide rangeof applications, but it is unclear to what extentit is presently used or can be made available.

Major references Bolza & Keating, 1972;Foullloy, 1974; Sallenave, 1964; Takahashi,1978.

Other references CTFT, 1961d; Foullloy,1965; Robyns & Wilczek, 1950

Authors M. Brink

BEILSCHMIEDiAKWEO nulldbr. ) Robyns &R. Wilczek

Protologue Bull. Jard. Bot. Etat 19: 494(1949)

Family LauraceaeSynonyms Tylostemon hweo Mildbr. (1914)Vernacular names Mfimbo (Sw).Origin and geographic distribution Betl-

schmiedio Aweo is endemic to the Eastern Arc

mountains in Tanzania

Uses The wood is used for furniture, floor-ing, panelling, veneer, gun stocks and toolhandles. It is also suitable for light construc-tion, Interior trim, Joinery, shipbuildlrig, sport-ing goods, agricultural implements, toys, novel-ties, turnery, poles, piles and mine props. It isalso used as fuelwood

Production and international trade Appre-ciable volumes of the wood were formerly usedlocally in Tanzania and exported to Germany,where it was used for wagons and shipsNowadays it is scarce.

Properties The heartwood is olive green, ondrying darkening to greenish brown or chest-nut brown;it is distinctly demarcated from thecream coloured and up to 5 cm-wide sapwood.The grain is straight, texture medium. Thewood is oilyThe density of the wood is 540-740 kg/in" at12% moisture content. Shrinkage rates are1.8% radial and 4.1% tangential from green to12% moisture content, and 3.0% radial and6.8% tangential from green to oven dry. Air-drying is slow, with material more than 2.5 cmthick having a tendency to split. Material 2.5cm thick can be kiln-dried satisfactorily. Logsmay develop star shakes a few weeks afterfelling. Dried wood is stable in service.At 12% moisture content, the modulus of rup-ture is 79 N/mm2, modulus of elasticity 11,100N/mm2, compression parallel to grain 45

.

N/min2, shear 10.5 N/min2, cleavage 65 N/mmradial and 76 N/mm tangential, and Janka sidehardness 4320 N.

The wood saws well, but with a tendency tosplit when star shakes are present, and with amarked blunting effect on sawteeth. The woodplanes well, but only at low speed, otherwisetearing may occur. It moulds, bores and inor-tises cleanly, sands and polishes to a cleanfinish, and glues well. The nailing propertiesare poor, and pre-boring is necessary nearedges. Turning is easyThe durability of the wood is moderate to high,and it is moderately resistant to marine borersand termites. The sapwood is liable to attackby Lyetus beetles. The heartwood is imperme-able to preservatives, the sapwood moderatelyresistanttO Impregnation

Botany Evergreen, medium-sized tree up to30 in tall; hole branchless for up to 15(-25) in,straight, up to 100 cm in diameter, withoutbuttresses; outer bark pale grey or pale brown,nearly smooth, flaking in large scales leavingliver-coloured patches, inner bark red; youngbranches finely hairy. Leaves alternate or op-postte, simple and entire; stipules absent; peti-o1e 1.5-2 cm long; blade oblong-ovate to ellipti-cal or oblong, (10-)11.5-18(-22) cm x 4-9(-11)cm, base broadly curieate to rounded, apex nar-rowly acuminate, glabrous, pinnately veinedwith c. 10 pairs of lateral veins. Inflorescencean axillary panicle, lax, 4-6 cm long; pedunclec. 5 cm long; branches up to 3 cm long. Flowersbisexual, regular, dark red; pedicel c. 2 minlong; perianth top-shaped to bell-shaped; tubec. 1.5 min long, lobes 6, ovate, c. 1.5 min long;fertile stamens (6-)9, in (2-)3 whorls of 3, in-ner whorl with glands, staininodes 3; ovarysuperior, at top gradually narrowing into thestyle. Fruit an ellipsoid olive-like berry 3.5-5cm x 1.5-2.5 cm, I'Seeded. Seed with semicir-CUIar, very thick, pale violet-brown cotyledonsOnly 12% of seeds of Beilschmiedio Aweo wererecorded to germinate after 2 months, rising to70% after 3 months, indicating physiologicaldormancy. The seed is recalcitrantBenschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Betlschmiediu hweo is classified in sub-genus Stemonodenio, section HUIe!gridio

Ecology Benschmiedio h"eo occurs in rain-forest at 800-1800 in altitude. In the East

Usambara mountains (Tanzania) the speciesshows good natural regeneration in old Moe-sopsis eminii Engl. plantations

Genetic resources and breeding Betl-schmiediu hweo is classified as vulnerable inthe 2006 TUGN Red list of threatened species,due to its limited distribution area

Prospects In view of its limited distributionand scarcity due to former exploitation there isno scope for increased use of BetlschmiedioAweo as a source of timber, and it should ratherbe protected or managed, for example in formerMoesopsis eminiiplantations.

Major references Bolza & Keating, 1972;Bryce, 1967; Takahashi, 1978; Verdcourt, 1996

Other references Baskin & Baskin, 2005;CAB International, 2005; Lovett & Clarke,1998b; Lovett at a1. , 2006; Robyns & Wilczek,1949; Roe at a1. , 2002; Sautu at a1. , 2006;Viisteensaari at a1. , 2000.

Authors M. Brink

BETLSCHMIEDIA 107

BEILSCHMIEDIALOUISllRobyns & R. Wilczek

Protologue Bull. Jard. Bot. Etat 19: 470(1949).

Family LauraceaeOrigin and geographic distribution Betl-

schmiedio IOUisiiis distributed in Cameroon,Central African Republic and DR Congo

Uses The wood (trade names: kanda, brownkanda) has an attractive appearance and issuitable for furniture, cabinet work, interiortrim, joinery, construction, flooring, shipbuild-ing, vehicle bodies, railway sleepers, mineprops, poles, piles, veneer and plywood. Thefruit is used as a condiment;it has a cinnamon-like taste

Properties The heartwood is brown, turningslightly greenish on exposure; it Is clearly de-marcated from the 5 cm-wide, pale yellow-brown sapwood. The grain Is usually straight,texture fine. At 15% moisture content, the den-sity of the wood is 720-800 kg/in3. The wood isfairly unstable, and Is best quarter-sawn. At15% moisture content, the modulus of ruptureis 128-168 N/min2, compression parallel tograin 38-65 N/min2, cleavage 15.7 N/min andChalais-Meudon side hardness 3.5. The woodsaws easily but slowly. It works wellwith handand machine tools, although there Is a markedblunting effect on tools due to the presence ofsilica. The wood nails easily without splitting,and it holds nails well. It finishes well, polishesto a walnut-like sheen and glues well. Thewood is only rarely attacked by termites andmarine borers, and it is moderately resistant tofungi. The sapwood is susceptible to attack by

.

108 TIMBERSl

Lyetus borers. The heartwood is impermeableto preservatives; the sapwood is moderatelyresistantto Impregnation

Botany Evergreen, large tree up to 50 intall, sometimes shrubby; hole branchless for upto 32 in, straight, cylindrical, with a diameterup to 100 cm, without buttresses; outer barkrough, brown, flaking; inner bark brownishpink, with a strong turpentine-like smell;crown flattened; young branches flattened,more or less hairy. Leaves alternate or almostopposite, simple and entire; stipules absent;petiole I-2.5 cm long, channelled, hairy to gla-brous; blade broadly oblong-lanceolate toobovate, 5-18 cm x 3-8 cm, base curieate, apexrounded to obscurely acuminate, leathery, up-per surface olive-brown and glabrous, lowersurface finely hairy to glabrous, pinnatelyveined with 6-10 pairs of lateral veins. Inflo-rescence a terminal or axillary, robust panicleup to 8 cm long, short-hairy; peduncle I-2 cmlong; bracts c. 3 min long, acuminate, caducousFlowers bisexual, regular, ochre-red to olive,short-hairy; pedicel up to 2 mm long; perianthbell-shaped, lobes ovate, c. I mm long, obtuse,thickened; fertile stamens 9, in 3 whorls, thosein outer 2 whorls sessile, those in inner whorlstalked and with 2 glands each, staininodes ina fourth whorl; ovary superior, c. 0.5 mm longFruit an ellipsoid-oblong to ovoid berry, 2-4.5cm x 1.5-2 cm, smooth, shiny, brown, I-seededBenschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Bet!schmiedio IOUisii belongs to subge-nus Synthorodertio.

Ecology Bet!schmiedio IOUisiioccurs in rain-forest and gallery forest.

Genetic resources and breeding Beil-schmiedio IOUisiihas a limited distribution and

it is unclear whether this species is threatenedby genetic erosion. It is not included in the2006 IUCN Red list of threatened species

Prospects The attractive appearance of thewood and its physical and technical propertiesmake it suitable for a range of purposes, butthe extent of its actual use and potential avail-ability are unclear.

Major references Bolza & Keating, 1972;Fouarge, G6rard & Sacr6, 1953; Robyns &Wilczek, 1951

Other references Fouilloy, 1974; Normand& Paquis, 1976; Robyns & Wilczek, 1949

Authors M. Brink

BEILSCHMIEDIAMANNll(Meisn. ) Benth. &Hook. f.

Protologue Gen. PI. 3(I): 158 (1880)Family LauraceaeChromosome number2n=24

Synonyms Tylostemon monnii Clieisn. ) Stapf(1909)

Vernacular names Spicy cedar (En). C6dreepic6 (Fr)

Origin and geographic distribution Ben-schmiedio monniiis distributed from Guinea

eastward to DR CongoUses The wood (trade names: kanda, pink

kanda) is used for construction, planks, doorframes, interior and exterior joinery, furniture,cabinet work, stairs, flooring, vehicle frames,canoes, carpentry, plywood, and other purposesrequiring an attractive appearance. It can beused as a substitute for mahogany (Swieteniospp. ) and bosse (Guoreo spp. )The pounded bark is eaten with rice as an ap-petizer. The fragrant leaves are pounded inwater, and after pressing through muslin theliquid is drunk. In Liberia the flowers arecommonly used to flavour rice and other food.The fruit is eaten and is an ingredient ofsauces. The seed is a popular food commonlysold in West African markets;it is roasted andground before consumption, and added as acondiment and enrichment to soups, rice andvegetables. The seed yields an edible oil. Betl-schmiedio monniiis planted as a shade tree forcoffee in Liberia.

A decoction of the bark and leaves is used as a

lotion to treat headache. Pounded fruits are

used to treat cough, bronchitis, intercostalpain, rheumatism and dysentery, whereas a

.

\ I^.' A-

,

co

Benschmiedio morinii- wild

\..

o

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..

decoction of the fruit is used in the treatmentof diarrhoea in calves

Properties The wood has an attractive ap-pearance. The heartwood is reddish yellow tored, with a persistent spicy smell, and is dis-tinctly demarcated from the pale yellow orcream-coloured, nearly odourless sapwood. Thegrain is straight, texture moderately fine. Oilcells are present in the wood. The density ofthe wood is 660-720 kg/in' at 12% moisturecontent. The wood dries extremely slowly. At12% moisture content, the modulus of ruptureis 98 N/mm2, modulus of elasticity 12,400N/min2, compression parallel to grain 50N/min2 Janka side hardness 5160 N, andJanka end hardness 5030 N. The wood is easyto work with antools and finishes well. It glueswell. The wood is resistant to fungal and insectattack and durable even in contact with thesoil or with fresh water. The heartwood is re-

SIStant to impregnation with preservativesPer 100 g edible portion the dried seeds con-tain: water 14.6 g, energy 1395 kJ (333 kcal),protein 5.9 g, fat 0.5 g, carbohydrate 75.8 g,fibre 1.6 g, Ca 220 ing, P 100 ing (Leung, Bus-son & Jardin, 1968). The bark contains tracesof alkaloids, the leaves traces offIavones.

Description Evergreen shrub or small tofairly large tree up to 35 in tall; hole branchlessfor up to 20 in, straight and cylindrical orslightly angular and sinuous, with a diameterup to 100 cm, base with heavy root swellings ornarrow buttresses up to I in high; outer barkgrey-brown or brown, often with large scalesand numerous smalllenticels, Inner bark pink-ish red to pinkish brown, turning red-brown onexposure, with a strong cedar-like smell; crownfairly narrow, dense, dark green; youngbranches and buds densely hairy; all plantparts with oil glands. Leaves alternate or al-most opposite, simple and entire; stipules ab-sent; petiole 0.5-1.5 cm long; blade oblong-lanceolate to oblong-elliptical or broadlyoblanceolate, (4-)7-26(-30) cm x (2-)3.5-10cm, base obtuse to curieate, apex acute to acu-inmate, markedIy folded, papery to leathery,glabrous, fragrant when crushed, pinnatelyveined with 6-10 pairs of lateral veins. Inflo-rescence an axillary panicle 4-15 cm long; pe-duncle 0.5-1.5 cm long. Flowers bisexual, regu-Iar, small, greenish; pedice1 I-3 mm long; perl-arith cup-shaped, 2-3 min long, lobes oblong,rounded, c. 1.5 min long; fertile stamens 9, in 3whorls, inner whorl with glands, staininodes 3,between the inner whorls of stamens; ovarysuperior, I-celled. Fruit a spindle-shaped, often

44

BRILSCHMIEDIA 109

,.

5

.

:. ?

4

,, .,

\

I.

,. ,

*I

3

I

. ,

2

Benschmiedio monnii - I, buse of 601e, 2, flow-ering brunch, . 3, flower in longitudinal section,4, 6707tch with fruit, ' 5, seed.Redrou)it und odopted by Achingd Satin Nur-hornon

slightly oblique berry 1.5-5.5 cm x 1.5-2.5 cm,red when ripe, I'Seeded. Seed with thin seed-coat, cotyledons thick, cone-shaped. Seedlingwith hypogealgermination

Other botanical information Bet!schmie-did is a pantropical genus coinprislrig about250 species, with about 80 species in tropicalAfrica and Madagascar. Benschmiedio indrini!is classified in subgenus Bet!schmiedio, sectionBenschmiedio. Benschmiedio msulorum Robyns& R. Wilczek is also included in this section; itis a shrub up to 4 in tall occurring in Camer-o0n, Gabon, Congo and DR Congo, in periodi-cally inundated forest, on riverbanks and is-lands. Its flexible stems are used for makingbows

Bet!schmiedio goboonensis oneisn. ) Benth. &Hook. f. , Bet!schmiedio Iebrunii Robyns &R. Wilczek and Betlschmiedio nitido Engl. areincluded in section HUIelondio of subgenusBenschmiedio. Benschmiedio goboonensis is amedium-sized tree up to 30 in tall with a holediameter up to 60 cm. It is distributed fromNigeria to DR Congo and occurs in wet and

,.. ,**,

,*** ,-,*.~ 1.31*

."

.

110 TIMBERSl

marshy locations in lowland rainforest. Itswood is easy to work and suitable for carpentryand joinery. Like that of Benschmiediu monnii,it is known under the trade names 'kanda' and

'pink kanda'. The ground bark of Benschmiediugoboonensis is a component of a paste rubbedon painful areas. Bet!schmiedio Iebruniiis 10-cally exploited for its wood. It is a smalltree upto 15 in tall with a hole diameter up to 30 cm,occurring in DR Congo in forest at 1450-1700in altitude. It seems rare. Benschmiedio nitido

is a shrub or smalltree up to 8 in tall, distrib-uted in Garnero0n and Congo. The density ofthe wood of Beitschiniedio nitidu is about 650

kg/in' at 12% moisture content. Shrinkagerates are 4.0% radial and 5.6% tangentialfromgreen to oven dry. At 12% moisture contentthemodulus of rupture is 65 N/mm2, modulus ofelasticity 5100 N/mm2, compression parallel tograin 32 N/min2, cleavage 15.3 N/mm andChalais-Meudon side hardness 2.6

Betlschmiedio uoriobilis Robyns & R. Wilczekand Benschmiedio zenheri Engl. (synonym:Benschmiedio leemonsii Robyns & R. Wilczek)are included in section ACrothecon of subgenusBenschmiedio. Benschmiedio Dongbilis is a

shrub or smalltree up to 10 in tall with a holediameter up to 25 cm, occurring rather coin-monly in DR Congo in the understorey offorestin swampy, periodically Inundated or drierlocations. Its hole is used for poles. Betl-schmiedio zenkeriis a shrub or smalltree up to15 in tall, occurring in Garnero0n and DRCongo in swampy and periodically inundatedforest. Its wood is used for planks, and for inak-ing canoes and paddles

Anatomy Wood-anatoimcal 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 F1m); 32vessel-ray pits with much reduced borders toapparently simple: pits horizontal(scalariform,gash-like) to vertical(palisade); 41: mean tan-gential diameter of vessellumina 50-100 pin;(42: mean tangentialdiameter of vessellumina100-200 F1m); 47: 5-20 vessels per square inn-limetre; (48: 20-40 vessels per square minime-tre); 56: tyloses common. Tracheids and fibres61: fibres with simple to minutely borderedpits; 65: septate fibres present; 66: non-septatefibres present; 69: fibres thin- to thick-walledAxial parenchyma: 79: axial parenchyma va-

siCentric; (80: axial parenchyma alitorm); (81axial parenchyma lozenge-anform); (83: axialparenchyma connuent); (89: axial parenchymain marginal or in seemingly marginal bands);91: two cells per parenchyma strand; (92: four(3-4) cells per parenchyma strand). Rays: 97ray width I-3 cells; 106: body ray cells pro-cumbent with one row of upright and/or squaremarginal cells; 107: body ray cells procumbentwith mostly 2-4 rows of upright and/or squaremarginal cells; 1/5: 4-12 rays per min. Secre-tory elements and cambial variants: 126: oiland/or mucilage cells present among fibresMineral inclusions: 159: silica bodies present;160: silica bodies in ray cells; 161: silica bodiesin axial parenchyma cells(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development In Liberia flow-ering is from January-April; ripe fruits arefound in November. In C6te d'Ivoire floweringis from April. Iuly and fruiting from October-December

Ecology Benschmiedio monniiis shade-lovingand usually grows in evergreen primary andsecondary forest. Outside evergreen forest, it isfound mainly in riverme and swamp forest

Propagation and planting Betlschmiediomonitii can be propagated by seed. The 1000-seed weight is about 6 kg. Germination takes21-30 days. The germination rate is about 80%

Management Benschmiedio monniiislocal-Iy being domesticated in C6te d'Ivoire

Genetic resources As Betlschmiedio mon-

nit occurs widespread and is common in manyregions, it is not threatened by genetic erosion

Prospects The wood of Benschmiedio mon-nit is suitable for a wide range of uses. Its sta-hinty and durability render it suitable for themanufacture of exterior wood finishing, win-dows and doors. Its attractive colour makes it

appropriate for floors, interior wood finishingand interior fitting. Its workability favours itsuse in the manufacture of furniture. The main

limitation is its delicate drying. In view of itsfavourable wood properties, there is potentialfor Benschmiedio morinii to become more jin-

portant as a source of timber. In Liberia, forinstance, it is considered to have export poten-tial. It deserves more research on propagationtechniques and SIIviculture for use in forestryplantations or in agroforestry programmes, e. gas shade tree for coffee cultivation.

Major references Abbiw, 1990; Burkill,1995; Foullloy, 1974; Kryn & Fobes, 1959;Robyns & Wilczek, 1951; Takahashi, 1978;Voorhoeve, 1965.

Other references ATIBT, 1986; Aubr6ville,1959c; Bouquet & Debray, 1974; CTRAD For-estry Department, 2003; de in Mensbruge,1966; Dudek, F6rster & missenbauer, 1981;Fouilloy, 1965; Foullloy, 1974; Hawthorne,1995; InsideWood, undated; Irvine, 1961; Jay,1948; Keay, 1954b; Leung, Busson & Jardin,1968; Oxford Forestry Institute, 1997-2004;Robyns & Wilczek, 1949; Tahoux Toua0, 2002;Vivien & Faure, 1988b.

Sources of illustration Voorhoeve, 1965Authors NyunaiNyemb

BEILSCHMIEDIA OBLONGIFOLIA Robyns &R. Wilczek

Protologue Bull. Jard. Bot. Etat 19: 466(1949)

Family LauraceaeOrigin and geographic distribution Bet!-

schmiedio oblongtfo!io is distributed from Ga-bon and Congo to eastern DR Congo.

Uses The wood (trade names: kanda, brownkanda) is locally used for construction and car-pentry. It is also considered suitable for furni-ture and for inside and exteriorjoinery

Properties The heartwood is yellowishbrown, becoming greyish on exposure, andturning yellowish brown again on drying; it isclearly demarcated from the about 4 cm-widebright yellow-pink sapwood. Quarter-sawnsurfaces have an attractive flame figure. At12% moisture content, the density of the woodis 690-815 kg/in". The wood air-dries well, butit is moderately unstable and quarter-sawingand careful drying are recommended to avoidwarping. At 12% moisture content, themodulus of rupture is 1/4-145 N/min2,modulus of elasticity 21,300 N/mm2, coinpres-SIon parallel to grain 43-64 N/mm2, cleavage14.9-19.6 N/min and Chalais-Meudon sidehardness 2.8-6.3. The wood saws well. Work-

ing properties for hand and machine tools aregenerally good, but variable due to occasionalinterlocked grain. The nailing, screwing andmortising properties are good. The wood is du-rable.

Botany Evergreen, medium-sized tree up to25 in tall; hole branchless for up to 16 in, cylin-drical, up to 70(-100) cm in diameter; outerbark bright brown, scaly, inner bark reddishbrown to pinkish yellow, with a sweet spicysmell; young branches flattened, more or lesshairy. Leaves alternate to opposite, simple andentire; stipules absent; petiole 0.5-I cm long,

more or less channeUed, hairy; blade oblong-lanceolate to narrowly oblong-elliptical, 4-15cm x 1.5-5 cm, base curieate to acute, apexrounded to obscurely notched, margin up-curved, leathery, glabrous, dark olive-greenabove, lighter below, pinnately veined with 8-11 pairs of lateral veins; blade of young leaveswine red, with translucent dots. Inflorescencean axillary, lax panicle I-4 cm long, glabres-cent; peduncle 0.5-1.5 cm long; bracts ovoid, c4.5 mm x 2 min, persistent. Flowers bisexual,regular, fleshy, small, pale green, glabrous;pedicel up to I mm long; perianth bell-shaped,c. 2 min long, lobes ovate-triangular, c. I mmlong; fertile stamens 9, in 3 whorls, those inouter 2 whorls sessile, those in inner whorlstalked and with 2 glands each, stainlnodes 3,in a fourth whorl; ovary superior, c. 0.5 min indiameter. Fruit a pear-shaped berry up to 2 cmx I cm, glabrous, I'Seeded.Seedlings are shade tolerant and grow well inthe understorey, but they are vulnerable in fullsun. Initial growth of seedlings is fast. In IQvu(DR Congo) Betlschmiedio oblongtfolio pro-duces fruits abundantly at the end of the yearBenschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Betlschmiedio oblongifolio belongs tosubgenus Synthorodenio. Benschmiedio congo-lurid Robyns & R. Wilczek and Benschmiedioletouzeyt Robyns & R. Wilczek are also Includedin this subgenus and sometimes also knownunder the trade names 'kanda' and 'brownkanda'

Betlschmiedio congolono is a large tree up to50 in tall, with a hole diameter up to 90 cm,and leaves characteristically rounded at bothends. It is a forest tree distributed in Camer-o0n, Central African Republic, Gabon, Congoand DR Congo. Its wood is white, rather softand very porous. At 12% moisture content thedensity of the wood is about 580 kg/in", themodulus of rupture 99 N/min2, modulus ofelas-ticity 11,500 N/mm2, compression parallel tograin 38.6 N/mm2, cleavage 9.8 N/min andChalais-Meudon side hardness 2.7.Betlschmiedio letouzeyiis a medium-sized treeup to 30 in tan, with a hole diameter up to 50cm. It is distributed in Gamero0n, Central Mri-can Republic and Congo. At 12% moisture con-tent the density of its wood is about 650 kg/in',the modulus of rupture 164 N/mm2, coinpres-sion parallel to grain 73.4 N/mm2, cleavage 6.5N/min and Chalais-Meudon side hardness 3.2

Ecology Bet!schmiedio oblongifolio usually

BEILSCHMIEDIA 1/1

.

1/2 TIMBERSl

occurs in small groups in rainforest up to 2200in altitude

Management Bet!schmiedio oblongifolio waslocally abundant in DR Congo in the 1950s,with up to 20 treesA1a with hole diameter over15 cm, and 12 ina of timber per ha, but morerecent information is not available. Growth

ring analysis indicates that the trees are coin-mercially exploitable when they are 50 yearsold.

Genetic resources and breeding Ben-schmiedio oblongtfolio is not very widespreadand recent data on its abundance are lacking,so it is unclear whether this species is threat-ened by genetic erosion. It is not included inthe 2006 IDCN Red list of threatened species

Prospects The wood of Benschmiedi0 o6-longifolio has favourable strength properties,as well as good durability and workability. Toolittle is known about the present distributionand abundance of the tree to assess whether its

Importance will increase beyond local use. As itseems to grow rapidly, it deserves more re-search on its possible use in plantations.

Major references Fouarge, Quoilin &Roosen, 1970; MIChelson, 1952; Robyns &Wilczek, 1951; Takahashi, 1978

Other references ATIBT, 1986; Foullloy,1965; Foullloy, 1974; Normand & Paquis, 1976;Robyns & Wilczek, 1949; Robyns & Wilczek,1950

Authors M. Brink

30(-100) cm in diameter, cylindrical, oftencrooked; outer bark greyish, rough, flaking inroughly circular patches, inner bark red, fi-brous, deeply pitted; crown rounded; youngbranches hairy to glabrous. Leaves alternate oralmost opposite, simple and entire; stipulesabsent; petiole 0.5-1.5 cm long, shallowlychannelled above; blade elliptical-oblong toelliptical or obovate, 2.5-19(-21.5) cm x 1.5-8.5cm, base curieate to rounded, apex acute, acu-inmate, rounded or obtuse, leathery, glabrous,pinnately veined with 8-14 pairs of lateral

Inflorescence a terminal or axillarypanicle 3-9 cm long, hairy, many-flowered;peduncle 0.5-2.5 cm long; bracts ovate, keeled,c. 7 min x 5 mm, deciduous. Flowers bisexual,regular, small, fleshy, brownish yellow to pink-ish brown; pedice1 0.5-3(-4) mm long; perianthcup-shaped to bell-shaped, c. 2 mm long; lobesrounded ovate, c. I mm long; fertile stamens 9,in 3 whorls, inner whorl with glands, staini-nodes 3, in a fourth whorl; ovary superior, c.0.5 mm long, gradually narrowed into the styleFruit an ellipsoid or ovoid berry 2-4 cm x 1.5-2cm, dull purple to black when mature, withdark red-purple juice, I-seededGrowth of Betlschmiedio ugondensis is slow. InDR Congo it flowers in August-October; ina-ture fruits have been recorded for August.Benschmiedio comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. Betlschmiediu 11gondensis belongs tosubgenus Synthorodenio. Two varieties aredistinguished: var. MgOndensis, distributed inDR Congo, Sudan, Uganda and Tanzania, and

kotongenis Robyns & R. Wilczek, withlonger inflorescences and larger flowers, anddistributed in the Central African Republic, DRCongo and Zambia

Ecology Benschmtedio MgOndensis occurs at900-1500 in altitude in gallery forest and lake-side forest, often in swampy localities

Management Benschmiedio ugonde, 1sts ispropagated by seed, sown in pots or directly inthe field. Seed can be obtained by collectingfallen fruits, putting them in a heap and allow-ing the fruit to decompose, after which theseeds can be separated and dried in the sun.Germination is accelerated by cracking thehard seedcoat. Wildlings are also used forpropagation. GOPpicing and pollarding are POS-sthle. Betlschmiedio ugundensis has beengrown in tree planting projects in Masaka(Uganda)

Genetic resources and breeding Ben-

veins

BEILSCHMIEDIAUGANDENSiS Rendle

Protologue Journ. Linn. SOC. , Bot. 37: 203(1905).

Family LauraceaeSynonyms Tylostemon ugondensis (Rendle)

Stapf (1909).Origin and geographic distribution Ben-

schmiedio ugondensis is distributed in theCentral African Republic, DR Congo, Sudan,Uganda, Tanzania and Zambia

Uses The hole is made into dugout canoes;the wood is also used for construction, furni-ture and toolhandles, and it has been used formine props. It is also used as fuelwood and forcharcoal making. The fruits are eaten

Production and international trade The

wood is traded within the countries of produc-tion

Properties The wood is durable.Botany Evergreen shrub or small to me-

dium-sized tree up to 27(-45) in tall; hole up to

var

.

schmiedio ugondensis is classified as vulner-able in the 2006 IUCN Red list of threatened

species. Major threats are overexploitation(commercial exploitation and local use), espe-cially in Uganda, as well as general decline ofits habitat

Prospects The wood of Benschmiedio MgOn-densis is locally used and traded, but its exploi-tation is unsustainable. Increased use shouldnot be promoted, and present use should bemade more sustainable.

Major references Eggeling & Dale, 1951;Katende, Birnie & Tengnds, 1995; 0magor,1999; UNEP-WCMC, 2004; Verdcourt, 1996.

Other references Begu6, 1958; Diniz, 1997;Lovett at a1. , 2006; Makerere University Insti-tute of Environment and Natural Resources,1998; Robyns & Wilczek, 1949; Robyns &Wilczek, 1951

Authors M. Brink

wood borer insects. The wood impregnates wellunder pressurein autoclaves

Botany Medium-sized tree up to 30 in tall;inner bark reddish, brittle; young branchesglabrous to brown-hairy. Leaves alternate,simple and entire; stipules absent; petiole I-2.5(-3) cm long; blade elliptical or ovate, 9-20(-28) cm x 4-11(-20) cm, base obtuse toacute, apex acute or shortly acuminate, papery,upper surface glabrous or somewhat hairy on

lower surface glaucous andmajor veins,sparsely hairy to densely brown-hairy, pin-nately veined with 7-12 pairs of lateral veinsInflorescence an axillary panicle 4-10 cm long,many-flowered. Flowers bisexual, regular,greenish; perianth with 2 series of 3 tepals, upto 4 min long, brown-hairy on both surfaces;stamens 9, in 3 whorls of 3, inner ones with 2glands near the base, staininodes 3; ovary su-perlor, gradually narrowing into the shortstyle. Fruit an ellipsoid to roundish berry c. 3cm in diameter, I-seeded.Benschmiedio Delutino flowers and fruits yearround.

Betlschmiediu comprises about 250 species andis distributed throughout the tropics, withabout 80 species in tropical Africa and Mada-gascar. In Madagascar about 10 species havebeen recorded

Ecology Benschmiedio uelutino occurs inevergreen forest up to 1200 in altitude

Management Logs need to be treated withpreservatives ifthey are not immediately proc-essed after feUing

Genetic resources and breeding Bet!-schmiedio Delutirto is rather uncommon andrestricted to a threatened habitat in Madagas-car, which makes it easily liable to genetic ero-SIon. It is not mentioned in the 2006 IUCN Redlist of threatened species

Prospects The wood of Bet!schmiedio ue-jutino is easy to work, but its low durabilitylimits is usefulness. It is suitable for Interior

carpentry, but outdoors it can only be usedafter treatment wlth preservatives.

Major references Gu6neau, 1971; Gu6-neau, Bedel & Thie1, 1970-1975; van der Werff,2003.

Other references Cainiez & Gu6neau,1972; Styger at a1. , 1999

Authors M. Brink

BEILSCHMIEDiAVELUTiNA (Kosterm. ) Kos-term.

Protologue Journ. Sci. Res. (Jakarta) I: 1/5(1952).

Family LauraceaeSynonyms Benschmiedio grandi/lord (Kos-

term. ) Kosterm. (1952).Origin and geographic distribution Ben-

schmiedio Delutino is endemic to Madagascar,where it occurs in the northern and eastern

part of the countryUses The wood of Benschmiedio Deluttno

(Malagasy trade name: voankoromanga) isused for light construction, Interior Joinery,moulding, panelling and shuttering. It is con-sidered suitable for rotary-peeled veneer. Thefruits are used as a condiment

Properties The heartwood is beige and notsharply demarcated from the somewhat palersapwood. The density of the wood is 500-620kg/in" at 12% moisture content. The wood driesrapidly, with only slight risk of distortion.Shrinkage rates are moderate, from green tooven dry 4.0% radial and 8.5% tangential. At12% moisture content the modulus of ruptureis about 46 N/mm2, modulus of elasticity 8700N/mm2, compression parallel to grain 46N/mm2 and Monnin side hardness 2.1. The

wood is easy to work. Nailing, screwing andgluing do not cause problems, painting andvarnishing give good results and a smooth sur-face is generally produced. The wood is notdurable and susceptible to attack by fungi and

BEILSCHMiEDiA 1/3

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BERRYACORDiFOLiA (Wind. ) Burret

Protologue Notizbl. Bot. Gart. Berlin-Dahlem 9: 606 (1926)

Family Tiliaceae (APG: Malvaceae)Chromosome number2n=40

Synonyms Berryo Qinmonillu Roxb. (1820)Vernacular names Tnncomalee wood, hal-

mina wood (En). Faux teck, teck du pays (Fr).Origin and geographic distribution Berryo

cordifolio occurs naturally in tropical Asia. Ithas been introduced into tropical Africa (SierraLeone, Nigeria, Tanzania, Reunion, Mauritius),Australia, Hawaiiand Fiji

Uses The wood of Berryo cordifolio yields avaluable timber used for high quality rumi-ture, but suitable for a wide range of uses, in-cluding light and heavy construction, flooring,mine props, boat building, vehicle bodies, cart-wheels, tool handles, ladders, sporting goods,agricultural implements, boxes and other pack-ing materials, sleepers and beams, poles, carv-ing, turnery, draining boards, cooperage, oarsand paddles, panelling, and paper making.Uses are mainly recorded for India, but in tropi-calAfrica the wood could be used similarly.The bark yields a fibre of low quality. In WestAfrica Berrya cordtfo!to has been planted inwindbreaks and shelter belts. It is also plantedas an ornamental, e. g. on Mauritius.

Production and international trade In In-

dia and Myanmarthe wood of Berrya cordifoliois commercially traded. In the 1970s annualexports from Myanmar amounted to about 500t, which corresponds to approximately 600 ina.

Properties The heartwood of Berryo cordifo-Jig is dark red-brown, often with darker streaksIt is distinctly demarcated from the whitish topale brown sapwood. The grain is straight orshallowly interlocked, texture fine to mediumThe wood surface is rather dull, with a slightlyoily feel. Freshly sawn wood has a pungentodour.

The wood is very resilient and tough, with goodwearing and weathering characteristics. It hasa density of 960 kg/in3 at 12% moisture con-tent. At 12% moisture content the modulus ofrupture is 92.5-1/7 N/mm2, modulus of elastic-ity 13,105-14,425 N/mm2 and compressionparallel to grain 44-55 N/mm2. The wood sea-sons slowly with no risk of serious splitting,but surface checks may develop. Drying woodstacks must be weighted down and ventilationmust be providedThe wood is difficult to saw, but it works wellwith other tools and has good bending proper-

ties. It finishes and polishes well and splitscleanly, but is difficult to glueThe heartwood is durable and extremely resis-tant to impregnation. The sapwood is suscepti-ble to Lyetus attack.The seeds of Berryor cordifolio contain 11% oil;they have stiff hairs which may penetrate theskin and cause painful itching. All ethanol ex-tract of the seeds proved highly toxic to theEuropean maize borer (OStrinio itubilolis). A hotwater extract of the leaves has shown antifun-

gal activity against Pythium ophonidermotum,whereas a cold water extract had no activity.

Botany Small to medium-sized tree up to20(-30) in tall; hole branchless for up to 11 in,up to 200 cm in diameter, often fluted butfairly straight; bark greyish brown, smooth;crown much-branched; branchlets glabrescentLeaves alternate, simple; stipules subulate, 4-7 mm long, caducous; petiole up to 7 cm long,slender, sparsely stellate-pubescent near apex,glabrescent; blade broadly ovate to broadlyelliptical, 7-24 cm x 5-16 cm, base cordate,apex shortly acuminate, margin entire to wavy-toothed, glabrous above, with tufts of hairs invein axils below, 5-7-veined from the baseInflorescence a terminal or axillary panicle,usually many-flowered; bracts narrowly ovate,up to 8 mm long. Flowers bisexual, regular;pedicel slender, 10-12 mm long, stellate-hairy;calyx campanulate, 4-5.5 mm long, Irregularly3-4-10bed, lobes up to 2 mm long, acute to ob-tuse; petals 5, 0bovate, up to 8 mm x 3.5 min,pinkish white or white, glabrous; stamensmany, up to 5 mm long; ovary superior, 3(-4)-celled, style slender. Fruit a 6(-8)-winged cap-SUIe, surrounded at the base by persistentflower parts, stellate-hairy, wings up to 2.5 cmx 1.3 cm, several-seeded. Seeds up to 4.5 minlong, angular, with caducous bristlesBerrya comprises 4 species distributed in tropi-calAsia. Berryo mollis Wall. ex Kurz has beenplanted in Tanzania. It resembles Berryo cordi-folio, but has pubescentleavesWhile seedling growth is slow, later on Berryocordifo!to grows fairly fast with an averageannual hole diameter increase of I cm for at

least the first 30 yearsEcology Berryo cordifolio requires partial

shade especially in the seedling stage; solitarytrees in full sun grow poorly. It does not growwell on clayey soils; it tolerates poor drainage,butts not resistant to drought. In Thailand firewas found to favour germination of the seeds inthe soil

Management Berryo cordifolio can be propa

gated by seed. The 1000-seed weight is about19 g. About 30% of the seeds germLnate in 14-33 days, but a germination percentage of 20%has also been recorded. In India and Sri Lankaseedlings 8-10 months old are used as bare-rooted planting stock. Stumps prepared from1.5-2-year-old stock with a diameter of 2 cmcan also be used; the shoot is trimmed to 3-4cm and the roots to 20 cm. Planting of stumpsresulted in 70-75% survival. Berryo cordifolioCDPpices welland produces root suckers

Genetic resources and breeding Becauseof the wide geographic distribution of Berryocordifolio there is little risk of genetic erosion

Prospects The fairly fast growth of Berryucordifolio and the high quality of its timbermake it worthwhile to intensify SIIviculturalresearch on this species in tropical Africa

Major references Boer & SOSef, 1998b;Burki11, 2000; Keating & Bolza, 1982; Robyns& Mener, 1991; Whitehouse at a1. , 2001

Other references Bhat, SIvaprakakasam &Jeyarajan, 1994; Bosser, 1987; Freedman at al. ,1979; Hanelt & Institute of Plant Genetics andCrop Plant Research (Editors), 2001; Marod ata1. , 2002

Authors M. Brink

Based on PROSEA 5(3): Timber trees: Lesser-known timbers

Bolusontht, s speciosus Is a beautifulcramps

ornamental and roadside tree, and it is a beeforage

Properties The heartwood is reddish brown,often streaked, and distinctly demarcated fromthe narrow, whitish yellow sapwood, that be-comes grey-brown in the course of time. Thewood is heavy, with a density of about 930kg/ina at 12% moisture content. It is hard, butturns well. It is durable, and resistant to ter-mites and pinhole borersF1avonoids (several pterocarpans and manyISOflavonoids) with antimicrobial activityagainst Bocillus subtilis, Candido mycodermo,ESCherichio coli, SOCchoromyces cereuisioe andStaphylococcus oureus have been isolated fromthe root and stem bark and the root wood. F1a-vonoids from the root wood also have shownantioxidant properties. Saponins with monus-cicidal activity have also been isolated fromBolusortthus speciosus

Botany Small deciduous tree up to 12(-18)in tall, frequently multistemmed; stems usu-ally straight, sometimes branching low, up to40(-100) cm in diameter; bark deeply fissured,pale grey to dark grey or dark brown; crownnarrow, dense, with upwards growing branchesand drooping foliage. Leaves alternate, clus-tered at ends of twigs, pendent, imparipm-nately compound with 3-7 pairs of leaflets, upto 28 cm long; stipules awl-shaped, minute;leaflets opposite, stalked, lanceolate, 4-8 cm xI-2 cm, asymmetrical and slightly curved, ta-permg to a sharp point, margins minutelytoothed, glabrescent. Inflorescence a terminalraceme up to 30 cm long, pendent, many-flowered. Flowers bisexual, papilionaceous,almost scentless; pedicel c. 1.5 cm long; calyxbroadly campanulate, c. 8 min long, 5-lobed;corolla c. 2 cm long, violet-blue with a whitespot inside standard; stamens 10, free; ovarysuperior, oblong-linear, densely hairy, I-celled,style curved. Fruit an oblong-linear pod 7-10cm long, flattened, pale brown, later becominggrey or blackish, tardily dehiscent, 3-8-seeded.Seeds ovoid, flattened, c. 6 min x 4 min, smoothand shiny, yellow-brownBolusonthus comprises a single species andappears related to Dieroeopetolum and Ploty-eelyphiumGrowth rates of up to 80 cm/year have beenrecorded for Bo!usernthus speciosus. The tree Isdeciduous for a short period. Under optimalconditions, young trees start to flower after 5-7years. It flowers from August to January, butthe flowering period of a single tree is short

BOLUSANTHUS 1/5

BOLUSANTHUSSPECIOsuS (Bolus) Harms

Protologue Repert. Spec. Nov. RegnlVeg. 2:15 (1906)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Chromosome number2n = 16.18Vernacular names Elephant wood, tree

wisteria, Rhodesian wisteria tree (En). Glycinearbre (Fr).

Origin and geographic distribution Bolu-sonthus speciosus occurs from Malawi andZambia south to north-eastern South Africaand Swaziland. It has been introduced else-where, e. g. in Kenya, Uganda, India and AUS-tralia.

Uses The wood is popular for the productionof high-quality furniture, household articlesand implements. The stems are used as fenceposts. In Zimbabwe a leaf decoction Is drunk tostop vomiting, whereas a root decoction is ap-plied as emetic. In Malawi a root decoction isdrunk to treat abdominal complaints. In SouthAfrica the roots are used to treat stomach-acheand the dried inner bark to treat abdominal

.

1/6 TIMBERSl

The flowers are pollinated by bees. Wild ani-inals including monkeys, giraffes and antelopeseat the pods and leaves and may disseminatethe seeds. The roots develop nodules containingnitrogen-fixing bacteria.

Ecology Bolusonthus speciosus occurs inwooded grassland up to 1100 in altitude, inregions with an annual rainfall of 450-700mm. It can be found on a variety of soils, buttsmost common on heavy alkaline soils. It cansurvive periods of drought and moderate frost

Management Seeds should be placed in hotwater and soaked for about 12 hours beforesowing. They can be sown in river sand andshould be covered with a thin layer of finersand. Germination starts 4-5 days after sow-Ing, but it may take up to 30 days, and thegermination rate of the seeds may be up to95%. The seedlings can be transplanted into asoil mixture of equal parts of sand, loam andcompost when the second leafhas emerged

Genetic resources and breeding A1-though Bolusonthus speciosus has a fairlylarge distribution area, it rarely occurs gre-gariously. It is protected in Swaziland, where itis considered seriously threatened because ofoverharvesting for building material

Prospects Although the wood is highly val-ued, Bolusonthus speciosus will remain of 11m-ited importance as a timber tree because of itscomparatively small size. It seems to have agood future as ornamental tree, having thesplendour of Joeorondo intinosifolio D. Donfrom tropical regions and Wisterio spp. fromtemperate regions.

Major references Ellis, 2003; mapwijk,2003; Palmer & Pitman, 1972-1974; van Wyk,1972-1974.

Other references Bojase at a1. , 2002;Coates Palgrave, 1983; Erasto, Bojase-Moleta& Maimda, 2004; Gelfand at a1. , 1985; ILDIS,2005; National Academy of Sciences, 1979;Neuwinger, 2000; van Wyk & van Wyk, 1997.

Authors R. H. M. J. Lemmens

.

Broussonetiogreueono- wild undplonted

it occurs throughout the western part of thecountry, and in Mayotte.

Uses The wood (trade names: vory, somely)is excellent for framing and construction. It isused for lightinteriorjoinery and cabinet work,ceilings, panelling, Interior trim, boxes, crates,blockboard, laminated board, peeled and slicedveneer for non-decorative purposes, core stockand particle board. The wood is traditionallyused for construction, boat building, canoes,carts, and bars of xylophones. It is also used asfuelwood. The bark is used for clothing. A de-coction of the bark is considered restorativeand arithelmintic

Production and international trade The

wood of Broussonetio greueono is much soughtafter for local use in western Madagascar. In1987 the log volume equivalent of boards soldon the market of Morondave was estimated at6200 ms. The wood has probably never beenexported.

Properties The heartwood is creamy tobrown; it is clearly demarcated from the 8-12cm wide, very pale sapwood. The wood islightweight, with a density of 400-500 kg/ina at12% moisture content. It dries easily and rap-idly, with low risk of distortion and checkingBoards 25 mm thick take I-2 months to airdry, boards 40 mm thick 5-6 months. The ratesof shrinkage are low to moderate, from green tooven dry 2.9-3.7% radial and 5.8-6.6% tangen-tial. Once dry, the wood is stable in serviceThe wood is soft. At 12% moisture content, themodulus of rupture is 81-104 N/mm', modulusof elasticity 6800-8200 N/min', compressionparallel to grain 34-43 N/min' shear 3-4N/min', cleavage 16-17 N/mm and Chalais-

\..,

o

,A

BROUSSONETIAGREVEANA (Baill. ) C. C. Berg

Protologue Bull. Jard. Bot. Etat 47(3-4)356 (1977).

Family MoraceaeSynonyms Chlorophoro greueono (Baill. )

Leandri(1948), Andeonthus greueonus (Baill. )Capuron (1972).

Origin and geographic distribution Brous-sonetzo greueono occurs in Madagascar, where

,.

.,

Meudon side hardness 1.3-1.9. The wood saws,

peels and works easily with ordinary tools, anddoes not have a marked blunting effect ontools. It does not split on nailing and screwing,and it holds nails and screws well. Gluing

properties are good. The wood polishes, waxes,varnishes and paints well. The heartwood ismoderately resistant to fungal attacks, fairlyresistant to termites, and resistant to dry woodborers. The sapwood has a low durability andshould be removed or be treated with preserva-tives before the wood is used. The heartwood is

only moderately permeable to impregnationDescription Dioecious, deciduous or some-

times evergreen, medium-sized tree up to 25 intall, white latex present in all parts, rapidlyturning pale brown on exposure; bole branch-less for up to 12 in, up to 80(-150) cm in diame-ter, generally straight; bark surface yellowishpink, longitudinalIy fissured, with rows ofbrown lenticels, flaking off in thin layers, Innerbark with yenowish and hard outer part andwhite and very fibrous inner part; crownspreading; branchlets with caducous greyishhairs. Leaves alternate, SLmple; stipules up to16 min x 3.5 min, membranous, more or lesscrumpled, late-caducous with clear scars; peti-

o1e slender, 5-40 mm long, grooved above,hairy; blade elliptical to oblong or lanceolate,2-20 cm x 0.5-8 cm, rounded or slightly cor-date at base, acuminate at apex, papery, lowersurface of young leaves greyish hairy, pin-nately veined with 6-20(-25) pairs of lateralveins. Inflorescence an axillary catkin; maleinflorescence towards the tip of the branch,cylindrical, I-12 cm long, reddish, many-flowered, peduncle 0.5-2 cm long, denselywhite-hairy; female inflorescence globose tooblongoid, c. 0.5 cm in diameter, peduncle 05-1.5 cm long, white-hairy. Flowers unisexual;male flowers sessile, with 4-5 free perianthsegments and 4 stamens; female flowers cov-ered by bracts, perianth 2-4-10bed, ovary supe-nor, c. I min long, stigmas 2, one of themstrongly reduced and the other up to 7 mmlong. Fruit a I'Seeded oblongoid drupe 4-5 minlong, many together arranged in a globose tooblongoid infructescence 1.5-2.5(-3) cm in di-ameter. Seed 2-3 min long.

Other botanical information Broussonetio

comprises 8 species, of which 7 occur in tropi-cal, subtropical and warm temperate regions ofAsia, and I in Madagascar

Anatomy Wood-anatointcal description (IAWAhardwood codes)Growth rings: a growth ring boundaries Iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 25: intervessel pits small(4-7prn); 26: intervesselpits medium (7-10 prn); 30:vessel-ray pits with distinct borders; SImlIar tointervessel pits in size and shape throughoutthe ray cell; 31: vessel-ray pits with much re-duced borders to apparently simple: pitsrounded or angular; 42: mean tangential di-ameter of vessellumina 100-200 F1m; (46: S 5vessels per square minimetre); 47: 5-20 vesselsper square minimetre. Tracheids and fibres:61: fibres with simple to minutely borderedpits; 66: non-septate fibres present; (68: fibresvery thin-walled); 69: fibres thin- to thick-walled. Axial parenchyma: 79: axial paren-chyma vasicentric; 80: axial parenchyma all-form; 81: axial parenchyma lozenge-allform;(83: axial parenchyma confluent); 91: two cellsper parenchyma strand; 92: four (3-4) cells perparenchyma strand. Rays: (97: ray width I-3ceUs); (98: larger rays commonly 4- to 10-senate); (102: ray height > I mm); 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; (110: sheath

BROUSSONETIA 1/7

3

Broussonetia greueo, "0 - I, twig with mole In-norescences, 2, twig with fernole inflorescences,3, twig with inIructescences.Redrown ond adopted by R. H. MJ. Lemmens

2

1/8 TIMBERSl

cells present); 1/5: 4-12 rays per mm. Secre-tory elements and cambial variants: 132: Iati-cifers or tanniferous tubes. Mineral inclusions136: prismatic crystals present; 137: prismaticcrystals in upright and/or square ray cells;(141: prismatic crystals in non-chambered axialparenchyma cells)on. Thiam, P. 06tienne & E. A. Wheeler)

Growth and development The availabilityof abundant water favours regeneration. Treesreach a height of 4-6 in when 7 years old. Ingeneral, flowering is in (June-)September-December, and fruiting in November-January(-February). In the dry, deciduousKirindy forest near Morondava Broussonetiogreueonu bears leaves during 3-4 months, fromNovember to March. Here, flowering is in Oc-toher-November, coinciding with the appear-ance of new leaves, and fruits are ripe in Janu-ary-February. It flowers and fruits regularly,every year or at least once every 2 years' Thefruits are much appreciated by lemurs andparrots, which certainly disperse the seeds

Ecology Broussonetio greueonu occurs up to800 in altitude, in all forest types of westernMadagascar. In deciduous dry forest it is partof the dominant forest layer. It mainly occurson slopes. In the much drier southern part ofMadagascar, it is confined to locations alongwater courses. In Mayotte it grows in allforesttypes, from humid to dry.

Propagation and planting Broussonetiogreueono is now commonly planted in the Mo-rondava region of Madagascar. The fruits areharvested when they are ripe and juicy. Theseeds are extracted from the pulp by repeatedwashing, and they are dried in the shade. The1000-seed weight is about 5 g. The seeds storepoorly and are preferably sown soon after col-Iection. They are sown in the shade, and thegermination rate is about 45%. After germina-tion, the plants are pricked out at a spacing of10 cm x 10 cm. After a year in the nursery theyare 50-100 cm tall and ready to be planted out.Best results are obtained by planting out leaf-less plants during the dry season, which givesmore than 90% success, against less than 80%success for planting out in the rainy season.Seedlings attain a height of up to I in in 2years under shade, compared to only 40 cm inmore exposed conditions

Diseases and pests In forest north-east ofMorondava it has been reported that the Mala-gasy giant rat (Hypogeomys ontimeno), an en-dangered rodent, feeds on young Broz, 880netiogreueono plants, and that these should be pro-

tested

Genetic resources In view of its rather

wide distribution, its plasticity in climatic andedaphic requirements, and the fact that it isone of the more widely planted tree species inwestern Madagascar, Broussonetia greueonodoes not seem Immediately threatened withgenetic erosion. Nevertheless, observationsshowed a rapid decrease in sales of boards ofthis species on the Morondava markets duringthe 1990s, accompanied by a reduction of seed-bearing trees in the forest as well as less abun-dant natural regeneration. Moreover, its habi-tat is under strong pressure from forest clear-ing for agriculture and livestock keeping

Prospects Byoussonetio greueono is highlyappreciated for its wood, which locally resultsin overexploitation, disappearance of its woodfrom markets, and reduced natural regenera-tion. The species deserves attention becausethe wood has interesting technological charac-tenstics, and propagation and planting tech-niques are becoming mastered. Techniques ofpropagation by cuttings deserve to be investi-gated

Major references Berg, 1977; Capuron,1966a; Ganzhorn & Sorg (Editors), 1996; Gu6-neau, 1971; Gu6neau, Bedel & Thie1, 1970-1975; Sallenave, 1971

Other references Berg, 1982; Berg & Hi-jinan, 1989; Cainiez & Gu6neau, 1972;Capuron, 1972; Debray, Jacquemin & Razafin-drainba0, 1971; Ganzhorn, 1995; InsideWood,undated; Pascal at a1. , 2001; Perrier de inBathie & Leandri, 1952; Schatz, 2001

Sources of illustration Berg, 1977Authors D. Louppe

CALODENDRUM CAPENSE (L. f. ) Thunb.

Protologue Nov. gen. PI. 2: 43 (1782)Family RutaceaeChromosome number2n=54

Vernacular names Cape chestnut (En)Chataignier on Cap (Fr)

Origin and geographic distribution CUIo-dendrum copense occurs naturally from north-ern Kenya to eastern South Africa. It is occa-SIonally planted as an ornamental and road-side tree within its area of distribution andelsewhere, e. g. in Australia

Uses The wood is used in house constructionand for poles, furniture, grain mortars, toolhandles, Implements and yokes. It is also suit-able for flooring, joinery, boats, vehicle bodies,

.

sporting goods, toys, novelties, musical instru-merits, boxes, railway sleepers, turnery, ve-neer, plywood and pulpwood. It is considered agood firewood and suitable for charcoal produc-tion. CUIodendrum copense is planted as anornamental and roadside tree, e. g. in Nairobiand Kampala; it is spectacular when in fullbloom. It is also used for shade, and it is

planted as a windbreak. It produces largeamounts of leaves that are useful as mulch. Itis a good source of nectar for honeybees. InSouth Africa the bark, which is sold in localmarkets as 'white umemezi', is used in cosinet-ICs to dye the skin whitish. The seed oil, caned'yangu oil' is popular in Africa as a skin-careproduct, e. g. in Kenya it is used as a skin emol-lient. The oilcan be used in soap making andas biofuel

Properties The heartwood is whitish tostraw-coloured, occasionany with greenish orbrownish markings, and indistinctly demar-cated from the sapwood. The grain is straight,texture fine to medium. Growth rings are dis-tinct.

The wood is medium-weight, with a density of610-740 kg/ms at 12% moisture content. Usu-ally drying causes no problems and little de-formation. The rates of shrinkage from green tooven dry are 4.8% radial and 7.3% tangentialAt 12% moisture content, the modulus of rup-ture is 97-99 N/min2, modulus of elasticity10,400-12,700 N/min2, compression parallel to

49-52 N/min2 shear 12-14 N/mm2,grain

Janka side hardness 6270-6450 N and Jankaend hardness 6180-7910 NThe wood saws and works easily with bothhand and machine tools, but surfaces may be-come slightly woolly. It can be finished to asmooth surface and polishes well. The nailing,boring, gluing, jointing and turning propertiesare allsatisfactory. The bending properties areexcellent. The wood is not durable and is sus-

ceptible to termites and Lyetus attack. Theheartwood is moderately resistant to linpreg-nation with preservativesSeeds contain about 60% oil. The seed oilis

yellowish and bitter. The fatty acid coinposltionof the o111s approximately: palmitic acid 18-24%, stearic acid c. 5%, oleic acid 33-44%, 11-Doleic acid 29-36%, linolenic acid c. I%, ara-chidic acid <1%. Tests showed a high potentialfor use as lubricant and as fuel in diesel en-

gines. The root bark contains calodendrolide, aIimonoid derivative, which has Iarvicidal prop-erties againstthe mosquitoAedes oegypti.

Botany Semi-deciduous medium-sized tree

to 20 in tall; bole branchless for up to 12 in,up

up to 90 cm in diameter, often fluted, old treessometimes buttressed; bark surface greyish,smooth; crown spreading but often dense; twigsdensely short-hairy. Leaves opposite, SLmple;stipules absent; petiole 2-10 mm long; blade

3.5-8(-11) cm,elliptical, 6-14(-18)curieate to rounded or slightly cordate at base,rounded to acute or shortly acuminate at apex,margin wavy, gland-dotted, pinnately veinedwith up to 20 pairs of lateral veins. Trillores-

terminal panicle with oppositebranches. Flowers bisexual, regular, 5-merous,

large and showy; pedicel up to 2 cm long; se-pals nearly free, ovate, 3-5 min long, minutelyhairy; petals free, narrowly oblanceolate, 2-3.5cm long, short-hairy, whitish to pink or palepurple; stamens as long as petals, alternatingwith narrowly oblanceolate, glandular staini-nodes slightly longer than petals; ovary supe-

2-3 min in diameter, 5-10bed, warty-rior,

glandular, 5-celled, on an elongated stipe 4-6mm long, style 1.5-2 cm long. Fruit a globosecapsule c. 3.5 cm in diameter, stiped, 5-10bed,strongly warty, brown, dehiscing with 5 woodyvalves remaining attached to the top, up to 10-seeded. Seeds half-globose to pyramid-shaped,I-1.5 cm in diameter, blackCUIodendrum copense usually grows moderatelyslowly, but under optimal conditions growth inheight may be up to I in/year. The root systemis not aggressive. Trees may start flowering 7-8 years after sowing, but 10 years Is morecommon. Trees grown from cuttings may flowerafter 4 years, The trees often lose most of theirleaves and are then almost bare for severalmonths, but they may be evergreen in coastalforest. Trees may flower throughout the year,but there are peaks, e. g. in DecemberJanuaryin Kenya and in October-December in south-ern Africa. The flowers are pollinated by but-ternies and bees. Fruits ripen about 3 monthsafter flowering. Monkeys and squirrels feed onthe fruits, whereas pigeons and parrots eat theseeds; these animals may disperse the seedsCUIodendrum comprises 2 species. Caloden-drum etchii Engi. is endemic to the westernUsambara Mountains in Tanzania, where itoccurs in montane Juniperus forest. It is asmall tree up to 15 in tall, differing from CUIo-dendrum copense in Its smaller flowers andlarger fruits with long spines. The wood of0010dendrum etchit is used for poles, tool han-dles and grain mortars. It is also used as fuel-wood. The roots are used medicinalIy for treat-

ing hypertension. The tree Is used for amenity

CALODENDRUM 1/9

cence a

cm x

120 TIMBERSl

and shade. In the IUCN Red list, CUIodendrumetchiiis classified as criticaUy endangered dueto habitat loss and degradation, caused by es-tablishment of commercial pine plantationsand expansion of agriculture

Ecology In EastAfrica Golodendrum copenseoccurs In evergreen montane forest and river-me forest at 1200-2300 in altitude, but insouthern Africa it can also be found in scrubvegetation and coastal forest at sea-level. Ittolerates various soil conditions including dryblack cotton soils, but prefers moist forest soils.In cultivation 0010dendrum capense thrives indeep, fertile, well-composted soil with plenty ofmoisture. Young trees are susceptible to frost,but old ones are frost tolerant.

Management Nori-viable seeds can be sepa-rated from viable ones in water because theyfloat. Pre-treatment of seed is not necessarybefore sowing. The germination rate of freshseed is generally high, up to 90%. Germinationtakes 10-40 days. One kg contains 600-1000seeds. Seeds can be stored for up to one yearwhen protected from insects. Cuttings takenfrom new growth and treated with a rootinghormone can also be used for propagation.Grafting has been practised successfully. Wild-lings are sometlmes used for planting. Youngplants can be transplanted easily. Managementof trees by pruning is possible, but only youngtrees can be coppiced.

Genetic resources and breeding A1-though Golodendrum copense is widespreadand not immediately endangered, it is uricom-mon in several regions within its distributionarea, e. g. in Uganda, Malawi, Zimbabwe andMozambique. Some caution is needed to pre-vent it from becoming threatened by geneticerosion, as is the case for Calodendrum eichii

Prospects CUIodendrum capense deservesmore attention for planting. It combines a highornamental value with products such as multi-purpose timber and useful seed oil. Althoughthe little information available indicates thatthe tree is not difficult to grow, more researchis needed on propagation techniques and inari-agement in relation to its uses.

Major references Bolza & Keating, 1972;Maundu & Tengn6. s, 2005; Palmer & Pitman,1972-1974; Takahashi, 1978; World Agrofor-estry Centre, undated

Other references Cassady & Liu, 1972;Grace at a1. , 2002a; Katende, Birnie &Tengnas, 1995; Kiprop, Rajab & Warijala, 2005;Lovett at a1. , 2007; Notten, 2001; Oxford For-estry Institute, 1997-2004; Shitanda, Mutuli &

Oding0, 1997; van Vuuren, Banks & Stohr,1978; van Wyk & Geneke, 2000.

Authors R. H. M. J. Lemmens

CALOPHYLLUMINOPHYi, LUM L.

Protologue Sp. PI. I: 513 (1753).Family Clusiaceae (Guttiferae, HyperICaceae)Chromosome number2n=32

Vernacular names Alexandrian laurel,beauty leaf, bintangor (En). Vintanina, hintan-gor (Fr). Loureiro de Alexandria (Po). Moton-doo, intoridoo, inkanja (Sw).

Origin and geographic distribution CUIo-phyllum mophy!Ium is widespread along thecoasts of eastern Africa (from Kenya to north-ern Mozambique), Madagascar and other In-dian Ocean islands, tropical Asia, northernAustralia and the islands of the Pacific OceanAlthough it is considered wild in most of thisarea, It is often unclear where it is truly wild ora relict of former cultivation. In Reunion andMauritius it has possibly been introduced. It islocally planted outside the natural distributionarea, e. g. in Guinea, Sierra Leone, C6ted'TVoire, Ghana, Nigeria, Cameroon and Ga-bon, where seemingly wild trees and seedlingscan be found near beaches. In Uganda it hasbeen established near Lake Victoria. It is alsoplanted in tropical America

Uses Throughout its area of distribution0010phyllum mophyl!urn is used for the con-struction of canoes and smallboats, but also formasts, keels, knees and pulley blocks. Its woodwas already used around 1600AD by the Span-jards in the Philippines for parts of their ships,especially the keel, ribs and cheek. The wood is

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Golophyllum mophy!Ium - wild

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occasionally also used for construction, carpen-try, flooring, stairs, furniture and cabinet work,cart-wheel hubs, vessels and musical instru-merits.

CUIophyl!urn mophyl!urn is planted as a road-side tree, in hedges and as a wind break, e. g. inGhana and Nigeria. The seed o111s used forillumination while the purified oil can be usedin soap production and as a carrier oil, skinmoisturizer and hair o11 in cosmetics and alsoin aromatherapy. In Mauritius a root decoctionis used to treat ulcers, boils and ophthalmia,the bark to treat orchitis, the latex rubbed onthe skin against rheumatism and psoriasis,and a leaf decoction to treat eye infections. Inthe Seychelles the resin is used to treat woundsand insect bites. In Kenya the seed o111s ap-plied to glandular swellings in the neck andjaws. Numerous uses in traditional medicinehave been recorded from tropical Asia and thePacific: the latex and pounded bark are appliedexternalIy on wounds, ulcers and to treatphthisis, orchitis and lung affections, and in-ternally as a purgative, after childbirth and totreat gonorrhoea; a leaf infusion is used totreat sore eyes, haemorrhoids and dysentery;heated leaves are applied to cuts, sores, ulcers,boils and skin rash; leaves are used in inhala-tions to treat migraine and vertigo; the seed oilis applied externalIy as an analgesic againstrheumatism and sciatica, and as a medicationagainst swellings, ulcers, scabies, ringworm,boils and itch; and the flowers are used as ahearttonic. The seeds are used as a fish poison.The stones of the fruit are used as marbles.The pulp of immature fruits is recorded as edi-ble, but caution is needed as toxic compoundsmaybe present in mature fruits.

Production and international trade Co!0-

phyllum timber ('hintangor') is of considerableimportance on the international market. How-ever, the bulk is produced in Borneo and NewGuinea and consists of other 0010phy!Ium spe-

0010phyllum mophy!min has only localimportance. Timber of CUIophyllum speciesfrom Madagascar is known as 'vintanina'. Theoilis traded internationally as tamanu oil orforaha oil, but quantities Involved are notknown. It is expensive; in 2005 the retail pricewas over Us$ 45011

Properties The heartwood is pinkish to red-dish brown, and clearly demarcated from thepale sapwood. The grain is interlocked, spiralor wavy, texture moderately coarse and un-even. Planed surface lustrous; strlpe figurepresent on radial surface and darker coloured

zigzag markings on tangentialsurface.The wood of 0010phyl!urn mophyllum is me-dium-weight to moderately heavy (560-800kg/ina at 15% moisture content) and moder-ately hard. The rates of shrinkage are medium,from green to oven dry 4.2% radial and 5.3%tangential. Drying is moderately difficult; thewood seasons fairly slowly with moderate de-fects such as end checking, splitting, cuppingand springing. Weighting down of stacks dur-ing air drying is recommended as this reducesthe tendency of the timber to warp and twistIn a test in the Philippines, the wood of Co!0-phyl!urn mophyllum showed the following me-chantcal properties at 15% moisture content:modulus of rupture 48 N/min2, modulus ofelas-ticity 7545 N/mm2, compression parallel tograin 42 N/min2, shear 6 N/min2, cleavage 60N/min radial and 72 N/mm tangential, Jankaside hardness 4820 N and Janka end hardness6045 N

The wood is often rather difficult to saw, andsawn surfaces tend to be woolly. The wood isnot easy to plane because of spiral or Inter-locked grain. It is not recommended to use thistimber for lengths over 3 in because it is oftenbent. Colophyllum mophyllum timber is ratedas moderately durable under exposed condi-tions. It is durable when used under water.The wood is resistant to termites. The heart-wood is resistant to preservative treatment.The wood contains 58% cellulose, 31.5% 11gnin,17% pentosan and no silica. The solubility ls4.4% in alcohol-benzene, 1.0% in cold water,4.5% in hot water and 12.4% in a I% NaOHsolution. The energy value is 19,100 kJA<g.Mature and dried seed contains 50-60(-73)% ofa bluish yellow to dark green viscous oil with adisagreeable flavour, that may contaln up to30% resinous materials. The fatty acid coinpo-sition of the seed o111s: palmitic acid 15%,stearic acid 13%, oleic acid 49%, linoleic acid21%, linolenic acid 0.3%, etcosanoic acid 0.9%and eicosenoic acid 0.7%. The oil also contains

glycolipids (6.4%) and phospholipids (1.6%). Ithas cicatrizing properties, explaining its tradi-tional and modern use in a wide range of skinproblems. The oil has also shown anti-inflammatory, antifungal, antibacterial andinsecticidal activity. It stimulates phagocytosisof cells of the reticulo-endothelial system andhas protective activity on the vascular systemClinical tests showed that the oil may reduceold scars. Refined oil, which is pale yellow, hasstrongly diminished medicinal properties.Triophyllums (4-phenyl-pyranocoumarins) have

CALOPHYLLUM 121

cies

122 TIMBERSl

been isolated from the leaves and seeds of Co-lophyllum mophyllum and these coumarinsproved to be non-nucleoside reverse transcrip-tase inhibitors of HIV type I. Triophyllum Band P are the most active compounds isolatedThey are closely related to the anti-HIV coin-pound (+)-calanolide A originally extractedfrom CUIophyl!urn 10ntgerum Miq. Some of the4-phenyl-coumarins isolated from 0010phyllummophy!Ium might be valuable as cancer che-mopreventive agents. Seed extracts showedsignificant molluscicidal activity; the hydroxy-acid calophyllic acid was isolated as the activecompound. The ether extract of the leavesshowed piscicidalactivity.The bark is astringent and contains 11-19%tannins and is reportedIy antiseptic and disin-fectant. The o1eoresin from the bark, whichcontains benzoic acid, shows cicatrising proper-ties

Description Medium-sized tree up to 25(-35)in tall, with sticky yellowish latex, usually withtwisted or leaning bole up to 150 cm in diame-ter, without buttresses; bark shallowly longi-tudinally fissured, pale grey and fawn, innerbark usually thick, soft, fibrous and laminated,pink to red, darkening to brownish on expo-sure; twigs 4-angled or rounded, terminal bud

plump, 4--9 min long. Leaves decussately OPPo-site, simple and entire; stipules absent; petioleI-2 cm long; blade elliptical, ovate, obovate oroblong, (5.5-)8-20(-23) cm long, rounded tocurieate at base, retuse to obtuse at apex,leathery, with closely parallel secondary vena-tion alternating with latex canals. Inflores-cence an axillary false raceme 7-15 cm long,

branched, 5-15(-30)-flowered.sometimes

Flowers bisexual, regular, sweetly scented;pedice1 1.5-4 cm long; tepals 8(-13), 0bovate, c.ICm long, white; stamens numerous, in 4 bun-dles, slightly connate at base, orange-yellow;ovary superior, globose, I-celled, style long andslender, flexuous, stigma peltate. Fruit a glo-hose to obovoid drupe 2.5-5 cm long, smooth,greyish green, I-seeded. Seed globose to ovoid,1.5-2 cm long, brown, very oily. Seedling withhypogeal germination; cotyledons remainingenclosed in stone

Other botanical information CGIophy!Iumis a very large genus comprising about 190species. Tropical Asia Is richest in species, butMadagascar and the Mascarene islands arealso comparatively rich with together about 20species. 0010phyllum mophyllum is the onlyspecies occurring in mainland Africa. It is themost widespread 0010phyllum species and itshows some variation, especially in fruit size;the fruits of tropical African specimens areusually comparatively small.CUIophyllum toeomohoco Wind. , endemic toReunion and Mauritius, is closely related to0010phyllum mophyllum; it usually occursmore inland than the latter and differs in themore close venation of the leaf. The two specieshave been confused, and their wood and seedoil are used for the same purposes. In Mada-gascar the wood of several other 0010phy!Iumspecies is also used for construction, carpentry,furniture and canoes, e. g. CUIophyllum chope-lien Drake, 0010phyllum drouhordii H. Perrier,0010phyllum fibrosum FF. Stevens, Golophyl-Ium Jingulotum P. F. Stevens, CGIophyllum inil-uum P. F. Stevens, Golophyllum punicu!aturnFF. Stevens, Golophy!Ium recedens Jum. &H. Ferner and CUIophyllttm DerticillotumP. F. Stevens

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; 7: ves-sels in diagonal and/or radial pattern; 9: ves-sels exclusively solitary (90% or more); 13:simple perforation plates; 22: intervessel pits

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alternate; 25: intervessel pits small (4-7 prn);31: vessel-ray pits with much reduced bordersto apparently simple: pits rounded or angular;32: vessel-ray pits with much reduced bordersto apparently simple: pits horizontal(scalari-form, gash-like) to vertical(palisade); 42: meantangential diameter of vessellumina 100-200pin; (45: vessels of two distinct diameterclasses, wood not ring-porous); 46: s 5 vesselsper square minimetre; 47: 5-20 vessels persquare minimetre; 56: tyloses common. Tra-cheids and fibres: 60: vascular/vasicentric tra-cheids present; 61: fibres with simple to inI-nutely bordered pits; (63: fibre pits common inboth radial and tangential walls); 66 non-

septate fibres present; 69: fibres thin- to thick-walled. Axial parenchyma: 76: axial paren-chyma diffuse; (85: axial parenchyma bandsmore than three cells wide); 86: axial paren-chyma in narrow bands or lines up to threecells wide; 93: eight (5-8) cells per parenchymastrand. Rays: 96: rays exclusively uniseriate;104: all ray cells procumbent; 106: body raycells 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. Mineral inclusions: 136: pris-matic crystals present; (141: prismatic crystalsin non-chambered axial parenchyma cells);142: prismatic crystals in chambered axial pa-renchyma cells.co. Ebanyenle, A. A. Oteng-kirioako, P. Baas &P. 06tienne)

Growth and development Under naturalconditions in tropical Asia, seedlings reach aheight of 40-50 cm in 6 months after germina-tion. Growth of young trees is discontinuousand branching is rhythmic. The hole and

orthotropic, although thebranches are

orthotropy of the hole is weaker than in other0010phyllum species, often resulting in a pro-cumbent hole, especially in trees growlrig onbeaches. However, CGIophyllum mophy!Ium issometimes an erect tree of closed forest. Oldertrees show sympodial growth. Young treesstart fruiting after about 10 years. The fruitsare mainly dispersed by water and can often befound washed up on beaches. When the outerlayer of the fruit has rotted away, fruits floateasily and can be dispersed by sea currentsHowever, fruits are also dispersed by fruitbats.

Ecology 0010phyllum mophyllum occurs wildon rocky and sandy sea shores, where it growsjust above the high-tide mark. Temperatures

where it grows are moderated by the proximityof the sea. annual rainfallis 750-5000 mm.The habitat is often pronouncedIy xerophyticdue to the exposed situation, brackish ground-water and salt-laden winds. Calophyllum trio-phy!!urn is sensitive to frost and fire. It issometimes found inland on sandy soils up to200 in altitude, especially on islands. It isplanted inland, up to 1200 in altitude

Propagation and planting Natural regen-eration usually occurs near the mother treeSeedlings grown in nurseries require shadeRemoval of the endocarp significantly reducesthe germination period to about 22 days, in-creases the germination rate and improvesseedling growth and development. Initialgrowth is slow and repeated weeding is neces-sary. There are up to 200 seeds/kg. Seed stor-age behaviour is recalcitrant.

Management Golophyllum mophy!Ium israrely planted for its timber, more commonlyas an ornamental or roadside tree

Diseases and pests In the Seychelles manyCUIophyllum mophyllum trees are affected bya vascular wilt disease caused by the fungusLeptogrophium CUIophy!It, which causes severedie-back and ultimately death of the tree. Thebeetle Crypholus tryponus is the likely vectorof the pathogen

Harvesting For production of timber longerthan 3 in trees with straight holes should beselected. For seed oil extraction, seeds areshelled; initially the kernels contain little ap-parent oil, but after about a month the kernelsturn chocolate brown and become sticky wlthoil; they are chopped, dried, pounded and thenboiled. The o111s skimmed from the top of theboiling water. The seeds may also be crushed toa paste and the o111s then drained off. Indus-trialIy the o111s extracted by cold expression,and is not refined so as to conserve all medici-

nalpropertiesYield CGIophy!!urn mophy!Ium timber is

rarely available in large quantities. A maturetree may yield 50 kg of dry fruits and 18 kg ofseed oil

Genetic resources 0010phy!Ium mophy!Iumis very widespread, both in the wild andplanted, and does not seem to be in danger ofgenetic erosion. However, witd populations arenot common in tropical Africa, and need to bedocumented. Some of the Colophyllum speciesendemic to Madagascar and the Mascareneislands might easily become endangered.

Prospects Types with straight boles inlghtbe interesting for cultivation as timber tree,

CALOPHYLLUM 123

.

124 TIMBERSl

but research on silvicultural aspects is re-quired, as well as more detailed information onworking properties, drying, preservative treat-merit and mechanical properties. Interest inthe oil for cosmetic purposes is likely to in-crease.

Golophyllum mophy!Ium is a source of poten-tial cancer chemopreventive agents and mayalso play a role in combination therapy againstAIDS.

Major references Barnps, Robson & Verd-court, 1978; Burki11, 1994; Gunb-Fakim &Brendler, 2004; Lemmens, 2003; Lim & Lem-mens, 1993; Patil at a1. , 1993; Perrier de inBathie, 1951; Stevens, 1980; Tomlinson, 1986;Wainhouse at a1. , 1998.

Other references Beentje, 1994; Decary,1946; Gunb-Fakim, Gu6ho & Bissoondoyal,1995; InsideWood, undated; Kokwar0, 1993;Robson, 1961; Robson & Stevens, 1980; Wil-narus, 1949

Sources of illustration Lim & Lemmens,1993

Authors R. H. M. J. Lemmens

crackles when burnt.

The leaves and bark contain tannins.

Botany Medium-sized to fairly large tree upto 30(-40) in tall, but usually smaller; holestraight, cylindrical, up to 50(-80) cm in di-ameter, without buttresses or with short but-tresses; bark grey to dark brown; crown dense,rounded. Leaves alternate, bipinnately coin-pound with a single pair of pinnae; stipuleslinear, caducous; petiole short, with large glandat apex on upper side; axes of pinnae 15-20 cmlong; leaflets in 5-6 pairs per pinna, opposite,elliptical, 7-15 cm x 2-6 cm, acuminate, Ieath-ery, glabrous. Inflorescence an axillary or ter-minalpendent spike 6-10 cm long, arranged inpanicles, many-flowered; bracts very small.Flowers bisexual, regular, 5-merous, small,sessile, hairy; calyx campanulate, c. 2 min long,shortly toothed; petals fused at base, c. 4 mmlong, pinkish to brownish orange; stamens 10,fused at base, much longer than petals, withglands at apex; ovary superior, slightly stalked,hairy, I-celled, style long and slender. Fruit anobliquely oblong pod up to 17 cm x 4.5 cm,strongly flattened, long-attenuate at base,rounded at apex, woody, 2-valved, 5-10'seededSeeds angular, flattened. Seedling with epigealgermination; hypocoty1 4-5 cm long, finely red-dish-hairy, epicoty1 3-4 cm long; cotyledonsthick, fleshy, oblong, notched, purplish.Gobyocolyx comprises 11 species and is re-stricted to the more humid forest types of Westand western Central Africa. It is related to

Xylio, which differs in its head-like inflorescen-ces. In western Central Africa 9 species occur,in West Africa 2

Gobyocolyx uubreuillei Penegr. occurs fromSierra Leone to C6te d'Ivoire, and is locallycommon in evergreen forest on moist soils andalong watercourses, especially in Liberia. Itswood Is used for construction, planks and ca-noes, but it is much lighter in weight, not du-rable and liable to warping. Ash from the woodIs used for vegetable salt, the seeds are edibleafter cooking, and a relieving salve is madefrom pounded seeds in palm oil to treatwomen's aching breasts. Golpoco!yx uubreuilleiis a fairly large tree up to 32 in tall, which dif-fors from Calpocolyx bruntbrocteotus in itsusually larger leaflets, which are more con-spicuously veined and shortly hairy, and in itslarger bracts, cylindrical calyx and larger pods.The branches of Gobyocolyx breuibrocteotus arehollow and may be inhabited by ants. Thefruits dehisce explosiveIy during the dry sea-son, shattering the seeds. In Sierra Leone flow-

CALPOCALYXBREVIBRACTEATUS Harms

Protologue Bull. SOC. Bot. france 58, mein. a155 (1912)

Family Mimosaceae (Leguminosae - Mimo-soldeae)

Origin and geographic distribution CUIpo-calyx breuibrocteotus occurs from Sierra Leoneto Cameroon

Uses The wood is locally used in house build-ing, e. g. for posts and planks, for tool handlesand canoes, and as firewood. The bark is usedin traditional medicine; a materation is appliedas a mouthwash to treat sores, and the innerbark Is used against stomach-ache. Ash ofburnt pods is used in soap-making or as saltThe seeds are edible after cooking

Properties The heartwood is dark brown,often variegated with red and black, and dis-tinctly demarcated from the greyish sapwood.The grain is irregular, texture moderatelycoarse. The wood is heavy, with a density ofabout 830 kg/ms at 15% moisture content, andhard. At 12% moisture content, the modulus ofrupture is about 135 N/min2, modulus of elas-ticity 16,650 N/mm2, compression parallel tograin 59 N/min2 and Janka side hardness11,365 N. The wood is difficult to work, butattractive when properly finished. It is durableThe firewood quality is inferior; the wood

ering is in September-December, and fruitingin January-April.

Ecology Gulpocolyx breuibrocteotus usuallyoccurs In evergreen forest, sometimes In semi-deciduous forest and gallery forest. The occur-Tence reaches a maximum in areas with about

2500 mm/year of rainfall. Calpocolyx breuLbroc-teatus Is a fairly common understorey tree,especially in Liberia and Ghana. It can befound on sandy-loamy, sandy-clayey and Iat-eritic soils.

Management Natural regeneration occursoften gregariously in the shade, but in ever-green forest in relatlvely exposed localitiesOne kg contains approximately 3000 seeds.The seed has a high germination rate, andgermination starts after 6-15 days. Freshlyfelled logs sink in water and cannot be trans-ported by floating down a river.

Genetic resources and breeding Golpoco-Iyx breuibrocteotus is widespread and locallycommon, and not under threat of genetic ero-SIOn

Prospects Gobyocolyx breuibrocteotus treesare often of too small slze to be of commercial

importance as a timber, the boles often beingshort and not reaching the minimum diameterlimit allowed for logging. They will remain ofsome importance for local house building be-cause of the durability of the wood

Major references Burki11, 1995; Cooper &Record, 1931; Holingren at a1. , 2004; Viniers,1984; Voorhoeve, 1965

Other references Bouquet & Debray, 1974;Busson, 1965; de to Mensbruge, 1966; Irvine,1961; Keay, 1958c; Kryn & Fobes, 1959; Neu-winger, 2000; Normand, 1960; Saville & Fox,1967; Takahashi, 1978

Authors R. H. M. J. Lemmens

.

CALFocALYx 125

Gobyoculyx heitzii- wild

Ies, furniture, agricultural implements, interiortrim, Joinery, railway sleepers, poles and piles,carving, toys and novelties, turnery and pat-tern making.

Production and international trade Thetimber of Golpocolyx hattzii has been exportedin small quantities from Garnero0n and Gabonto Europe under the trade name 'injama'.Probably it is still traded in mixed consign-merits with other medium-heavy timbers ofsimilar colour

Properties The heartwood is reddish brown,often with irregular brown streaks, usuallydistinctly demarcated from the wide pale pink-ish grey sapwood. It is sometimes traded as asubstitute of African mahogany (from Khoyospp. ), which has a similar figure.The wood is heavy, with a density of about 800kg/ina at 15% moisture content, and hard. Thegrain is often wavy and irregular, texture inod-erately coarse to fine. The rates of shrinkageare high, 4.7-6.5% radial and 8.5-11.4% tan-gential from green to oven dry. The timbershould be seasoned with great care to avoiddistortion, and stock should be quartersawnRecorded mechanical properties show a widerange: at 12% moisture content, the modulus ofrupture is 125-215 N/mm2, modulus of elastic-ity 9220-17,650 N/mm2, compression parallelto grain 54-88 N/mm2, and shear 8.8-17.0N/min2

The working properties are variable, dependingon the amount of irregular grain in the mate-rial. The wood saws moderately well, and plan-ing may result in woolly surfaces due to ten-SIon wood. Pre-boring Is necessary for nailingand screwing. The wood is moderately durable

CALPOCALYXHEITZllPellegr.

Protologue Bull. SOC. Bot. France 84: 643(1937).

Family Mimosaceae (Leguminosae - Mimo-soldeae)

Vernacular names Miama (Fr).Origin and geographic distribution Gobyo-

colyx heitzii has a restricted area of distribu-tion, occurring in southern Gamero0n, Equato-rial Guinea and Gabon, particularly in coastalforest.

Uses The wood is used for indoor as well asoutdoor construction. It is also suitable for

flooring, mine props, shipbuilding, vehicle bod-

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126 TIMBERSl

and liable to termite and Lyetus attacks. Theheartwood is extremely resistant to preserva-tive treatment, the sapwood moderately resis-tant

Description Medium-sized to fairly largetree up to 35 in tall; hole straight, cylindrical,up to 90 cm in diameter, with steep buttresses;bark grey to greyish brown, with irregularridges forming a Teticulate pattern, inner barkfibrous, pinkish-brown; branches often hollowLeaves alternate, hipinnately compound with asingle pair of pinnae; stipules linear, caducous;petiole up to 13 cm long, shortly hairy, withlarge gland at apex on upper side; axes of pin-nae (17-)30-52 cm long; leaflets in 8-15 pairsper pinna, opposite, oblong-lanceolate to nar-rowly elliptical, up to 15 cm x 3.5 cm, acumi-nate, papery, glabrous. Inflorescence an axil-Iary pendent spike 6-15 cm long, many-flowered. Flowers bisexual, regular, 5-merous,small, sessile, slightly hairy; calyx campanu-late, c. 2 mm long, shortly toothed; petals fusedat base, c. 4 min long, whitish yellow to palebrown; stamens 10, free, longer than petals,with glands at apex; ovary superior, slightlystalked, hairy, I-celled, style long and slenderFruit an obliquely obovoid pod up to 20 cm x

4.5 cm, strongly flattened, long-attenuate atbase, obtuse at apex, woody, brown, 2-valved,6-8-seeded. Seeds angular, flattened. Seedlingwith epigealgermination

Other botanical information CGIpocolyxcomprises 11 species and is restricted to themore humid forest types of West and westernCentral Africa. It is related to Xylio, whichdiffers in its head-like inflorescences. Western

Central Africa counts 9 species, West Africa 2Most of the Central African species have smalldistribution areas

Gulpocoly" dinhlogei Harms occurs from Nige-ria to Cabinda (Angola), and is locally commonin evergreen forest, especially in Garnero0n. Itswood resembles that of Gulpocolyx heitzii andis used for carpentry, but Gobyoco!yx dinklogeiIs usually only a smalltree up to 15 in tallwitha hole of up to 25 cm in diameter. It differsadditionally in its glabrous petioles and widerleaflets. In Cameroon pulverized bark of CUI-pocolyx dinh!ageiis applied to wounds and asan anodyne. Golpocolyx dinhlogeiis sometimesconfused with CGIpocolyx Momei Pierre exHarms, also a smalltree, but the latter occursmainly in coastal forest in southern Garnero0n,Equatorial Guinea and Gabon, and differs inhaving shortly hairy petioles and longer fruitsProbably 00400co!yx Mumeiis also occasionallyexploited for its timber. It is rated as vulner-able in the TUGN Red List of threatened spe-cies. Several other Gulpocolyx species are un-doubtedIy also felled occasionally in Gamero0n,Equatorial Guinea and Gabon fortheir timber.

Anatomy Wood-anatoimcal 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); 25: intervessel pits small (4-7 prn);(26: intervessel pits medium (7-10 prn)); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to Intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;(43: mean tangentialdiameter of vessellumina2 200 prn); 46: S 5 vessels per square minime-tre; 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; (65: septate fibres pre-sent); 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: 79:axial parenchyma vasicentric; 80: axial paren-chyma allform; 81: axial parenchyma lozenge-

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allform; 83: axial parenchyma confluent; (84axial parenchyma unilateral paratrachea1); 91:two cells per parenchyma strand; 92: four (3-4)cells per parenchyma strand. Rays: 97: raywidth I-3 cells; 104: all ray cells procumbent;1/5: 4-12 rays per mm. Mineral inclusions:136: prismatic crystals present; 142: prismaticcrystals in chambered axial parenchyma cells(N. P. Moilel, P. 06tienne & E. A. Wheeler)

Growth and development The growth ofGobyocolyx heitziiis quite fast. In the Sthangearboretum near Libreville, Gabon, a 67-year-old planted plotshowed a mean annual growthrate of about 10 ms/}Ia. The stand was verydense, 567 treesA1a, and the mean annualgrowth in diameter was only 5.4 mm/tree (9min for the biggest trees). The mean volumeper tree is 1.4 ina (5.9 in3 for dominant trees)The fruits of CGIpocolyx heit2ii dehisce explo-SIvely during the dry season, shattering theseeds.

Ecology 004poco!yx heitsit occurs in primaryand old secondary rainforest, most abundantlyin littoral forest. In southern Gamero0n Gulpo-CUIyx heitziiis locally common together withSOCoglottis gobonensis (Baill. ) Urb. in lowlandevergreen forest and coastal forest up to 200 inaltitude and with an annual rainfall of 2200-2800 mm

Management Gulpocolyx heitstits characte-rized as a non-pioneer light demander. Trees oflarger sizes usuany occur scattered and in lowdensities. In Gabon the average exploitabletimber volume of Gulpoco!y, c heitzii trees withhole diameter over 35 cm has been recorded asless than 4 mama

Handling after harvest Freshly felled logssink in water and cannot be transported byfloating down a river

Genetic resources CUIpocolyx heitzii has alimited area of distribution and could readilybe threatened by habitat degradation and fell-ing. It has been classified by IUCN as vulner-able. The best means of protection Is by creat-ing forest reserves of large extent within thearea of distribution of the species. Some otherGulpocolyx species would profit similarly fromthese measures.

Prospects Gobyoco!yx heitziiis an Triterest-ing timber tree, of which little Is known. Re-search is needed on Its ecological requirements,regeneration and growth rates before its role innatural forests subject to sustainable timberexploitation can be evaluated. Gulpocolyx ap-pears in need of a taxonomic revision, but thisis hampered by the scanty material of many

species present in herbariaMajor references anonymous, 1978; Bolza

& Keating, 1972; Raponda-Walker & Sillans,1961; Viniers, 1989

Other references de Saint-Aubin, 1963;InsideWood, undated; Normand & Paquls,1976; Takahashi, 1978; Tchout0, 2004; Viniers,1984; Wilks & TSSemb6, 2000.

Sources of illustration Viniers, 1989;Wilks & Issemb6, 2000.

Authors R. H. M. J. Lemmens

CANARiUM 127

CANARIUMMADAGASCARIENSE Engl.

Protologue A. DC. , Monogr. phan. 4: 111(1883)

Family BurseraceaeSynonyms Congriumpulchebrocteotum Gull-

Iaumin (1909)Vernacular nanies Rainy, aramy orr). Mpafu,

inbani(Sw)Origin and geographic distribution Curio-

rium inodogoscoriense occurs in Tanzania,Mozambique andMadagascar

Uses In Madagascar, where Curiorium mudo-goscoriense is known as 'rainy', the wood isused for dugout canoes, boxes, crates, construc-tion, tool handles, matches, everyday furnitureand hidden parts of fine furniture, veneer, ply-wood, hardboard and particle board. It is alsoconsidered suitable for Joinery, Interior trim,turnery, and sometimes for floorLng, poles andpiles. The dry heartwood is used to make torches,and the wood is used for making charcoal.The fruits are eaten, especially by children,and the roasted seeds are eaten like ground-nuts. The resin is used for caulking boats, as

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Guitarium mudogoscortense - wild

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128 TIMBERSl

glue, and for trapping birds and small main-inals. It has been used in perfumes and paints,and it is used in the manufacture of incense

used during religious ceremonies. It is a stimu-Iant and its activity increases when pickled inalcohol. The resin is used medicinalIy for thetreatment of urinary complaints, dental caries,rheumatism, wounds, and as a disinfectant.After heating its vapour is inhaled againstheadache and other pain, and immersion in itsvapour is believed to protect against infectionsThe resin is sometimes used as an insecticideCongrium ingdogoscoriense serves as an or-namentaland shade tree

Production and international trade The

wood is sometimes exported from MadagascarProperties The heartwood is pinkish brown;

it is not distinctly demarcated from the greyishsapwood, which is up to 5 cm wide. Freshly cutwood often has a bluish tinge. The grain maybe straight but is fairly often interlocked orspiral, the texture medium to coarse. The woodis lustrous and contains an o1eoresin.

The wood has a density of 510-690 kg/ina at12% moisture content. It air dries well. Planks

25 min thick can be dried to 30% moisture con-tent in 2 months under humid lowland condi-

tions in Madagascar, whereas it takes about Imonth under higher altitude conditions. Therates of shrinkage are high, from green to ovendry (2.8-)5.1-8.0% radial and (6.4-)7.5-10.8%tangential.At 12% moisture content, the modulus of rup-ture is 105-162 N/min2, modulus of elasticity10,300-14,800 N/mm2, compression parallel tograin 41-56(-65) N/mm2, shear 5-12 N/min2,cleavage (6-)12-23 N/mm and Chalais-Meudonside hardness (1.8-)2.2-4.0.The wood is sometimes difficult to saw due to

high silica content, but normally works easily,with only a slight blunting effect on cuttingedges. It nails well, but has poor nail holdingproperties. It glues easily, takes paint well,stains well and has good veneering properties,butthe use offiller is recommended

The wood is not durable, being liable to attacksby insects, including termites, and fungi. Thesapwood ls susceptible to Lyetus borers. Theheartwood is extremely resistant to impregna-tion, but the sapwood can be treated with pre-servatives. The wood is resistant to sea water

The fruit pulp has a somewhat sweet taste; itcontains up to 60% fat

Description Deciduous, dioecious, medium-sized to fairly large tree up to 37 in tall; holestraight, branchless for up to 27 in, up to 200

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Cartonum ingdogoscoriense - I, Ieob, twig, ' 2,leaf' 3, inflorescence, 4, fruitsRedrown und udopted by AChmod Sotiri Nur-hornon

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cm in diameter, often buttressed; bark surfacesmooth or rough, fissured, brown or greyish,bark containing a turpentine-odoured, white,clear resin turning yellowish on hardening;crown round, with spreading branches; youngbranches, petiole, rachis and inflorescence cov-ered with ferruginous hairs. Leaves alternate,imparipinnately compound with 2-9 pairs ofleaflets, up to 25(-55) cm long; stipules absent;petiole up to 7 cm long; petiolules 0.5-3 cmlong, hairy; leaflets opposite, ovate-oblong tooblong, 4-20 cm x 2.5-10 cm, the lower pairssmaller and often stipule-like, base rounded toslightly cordate, apex bluntly acuminate, mar-gin entire or wavy, glabrous, but rutdveindensely hairy below, pinnately veined with 7-20 pairs of lateral veins. Inflorescence a termi-nal or axiUary panicle, spreading, up to 35 cmlong (female inflorescences smaller than maleones), with flowers in clusters of 6-15. Flowersunisexual, regular; pedicel IC6) mm long; ca-Iyx 2-4(-8) mm long, campanulate, 3-lobed,densely hairy outside; petals 3, free, oblong, c5(-10) min x 3(-8) min, keeled, white, hairyoutside; stamens 6, 2-3 mm long, reduced infemale flowers; ovary superior, 3-celled, absent

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or vestigial in male flowers. Fruit an ovoid-ellipsoid drupe up to 5.5 cm x 3 cm, purplewhen ripe, indehiscent, with yellow, aromatic,fleshy pulp enclosing a trigonous stone 2-3 cmx 1.5-2 cm, with 3 minor ridges on each side,up to 3-seeded. Seeds compressed narrowlyovoid, up to 2.5 cm x I cm, brown. Seedlingwith epigeal germination; cotyledons dividedinto 3 linear-elliptical leaflets, lateral leafletssometimes also deeply divided.

Other botanical information Gorierrium

comprises about 80 species distributed in high-rainfall areas in the Old World tropics; it ismost common in South-East Asia, with only 3-4 species in tropical Africa. However, the genusis currently under revision, and preliminaryestimates indicate that about 30 species arepresent in Madagascar, most of which havestill to be described.

Curiorium ingdogoscoriense is extremely van-able, and it has sometimes been divided intovarious species and wood types (e. g. 'whiterainy' and 'red rainy'). Within Curioriummudogoscoriense several subspecies have ten-tatively been distinguished, with subsp. mado-goscoriense (synonyms: Curiorium lieber-tionum Engl. , Cottonum multi/10rum Engl. )occurring in East Africa and Madagascar, andsubsp. o6tusi/onurn (Scott-Enjot) Leenh. (syno-nyms: Cartonum botu, rin Engl. , Cartonumo6tusifolium Scott-Enjot) and subsp. bullotumLeenh. in Madagascar onlyCuriorium pontcu!drum (Lain. ) Benth. ex Engl.lyernacular name:'bois colophane') is a tree upto 25 in tall, with a hole up to 2 in in diameter.It is endemic to Mauritius, where it occurs oc-casionally in the remnants of native forest inupland regions. Its wood has been used in con-struction. The resin is known as 'demi de

Maurice'. A leaf poultice and the resin are ap-plied to body parts affected by rheumatism; theleaf poultice is also applied on ulcerations. Ex-tracts of the stem, wood and bark have shownantibacterial activity. Collorium puniculotumis now highly threatened and is classified asendangered in the 2007 IUCN Red list.

Anatomy Wood-anatorntcal 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 F1m);27: intervessel pits large (^ 10 prn); 32: vessel-ray pits with much reduced borders to appar-ently simple: pits horizontal(scalariform, gash-

like) to vertical(palisade); 42: mean tangentialdiameter of vessellumina 100-200 pin; 47: 5-20 vessels per square minimetre; 56: tylosescommon. Tracheids and fibres: 61: fibres withsimple to minutely bordered pits; 65: septatefibres present; 68: fibres very thin-walled; 69:fibres thin- to thick-walled. Axial parenchyma:75: axial parenchyma absent or extremely rare;78: axial parenchyma scanty paratrachea1; 92four (3-4) cells per parenchyma strand. Rays97: ray width I-3 cells; 106: body ray cells pro-cumbent with one row of upright and/or squaremarginal cells; 1/5: 4-12 rays per min. Secre-tory elements and cambialvariants: 130: radialcanals. Mineral inclusions: 136: prismatic crys-tals present; 137: prismatic crystals in uprightand/or square ray cellson. Thiam, P. 06tienne & E. A. Wheeler)

Growth and development In MadagascarCuriortum mudogoscoriense flowers mainly inOctober-January. The fruits and seeds areimportant elements in the diets of lemurs andothersmallmammals.

Ecology In Madagascar Congrium mudogos-curiense occurs widespread but scattered inmoist and dry forests, especially along water-courses, from sea-level up to 2000 in altitude.In Tanzania it is rare in forest remnants on

sandy soils in shallow river valleys, from sea-level up to 300 in altitude, in areas with anaverage annual rainfallofabout 1000 min

Management In a forest inventory in north-western Madagascar (altitude 220 in), a one-hectare plot contained 34 Curiorium ingdogos-conense trees with a diameter of 10 cm or

more

Canal{Inni 129

Genetic resources Cottonum mudogoscu-riense seems to be approaching extinction inTanzania. It does not seem to be threatened in

Madagascar.Prospects In Madagascar Canonum mudo-

80scoriense is a useful multipurpose tree, and,despite the poor durability of the wood, it is avalued native species for commercial forestryin Madagascar, due to the large size of the tree.It seems worthwhile to explore the potential forplantations of the species, but no information isavailable on appropriate propagation tech-niques and management practices. Research inthese areas is therefore needed

Major references Bolza & Keating, 1972;Garniez & Gu6neau, 1972; Gillett, 1991; Gu6-neau, Bedel & Thie1, 1970-1975; Leenhouts,1959; Parant, Chichignoud & Rakotova0, 1985;Ferner de in Bathie, 1946; Raharimampionina,2003; Raharimampionona at a1. , 2007; Salle-

130 TIMBERSl

nave, 1971Other references hadriamahery, 1994;

hadriamihaja, 1986; Boiteau, Boiteau & A1-10rge-Boiteau, 1999; Goode, 1979; d'Amico &Gautier, 2000; Gachathi, 1999; Grenfe11, 1999;Gu6neau, 1971; Gunb-Fakim & Brendler,2004; Gunb-Fakim, Marie & Narod, 2005; Hol-loway, 2004; InsideWood, undated; Lovett ata1. , 2007; Raharimampionona, 2006; Raheli-nor0, 1994; Sallenave, 1964; Strahm, 1998;Takahashi, 1978; Vasey, 1997; Wild, 1963a

Sources of illustration Perrier de in

Bathie, 1946Authors M. Brink

CARALLLABRACHIATA (Lour. ) Merr.

Protologue Philipp. Journ. Sci. 15(3): 249(1919).

Family RhizophoraceaeChromosome number2n =48,112Synonyms Gorging mudogoscoriensis (DC. )

Tu1. (1856).Vernacular names Corkwood, corkybark,

maniawiga (En)Origin and geographic distribution Carol-

jig brachioto is widespread from Madagascarto tropical Asia and northern Australia. Twocollections are known from Reunion

Uses The wood is suitable for general con-struction, house building, posts, cabinet work,furniture, parquet flooring, railway sleepers,musical instruments, tool handles, pictureframes, veneer, Interior finish and panelhng,pallets and packing material. Due to its highenergy value the wood yields good-quality fu-elwood and charcoal. The fruits are eaten. In

tropical Asia the leaves and bark are used inlocal medicine to treat septic poisoning anditch. The tree is sometimes planted as an or-namental, especially a cultivar with a narrowcolumnar habit and pendulous branches('Honiara'), e. g. along roads in the SolomonIslands

Production and international trade The

trees usually are too scattered and too smalltobe of great Importance for timber. Smallamounts of timber are exported from Borneoand PapuaNew Guinea

Properties The heartwood is yellowishbrown to reddish brown and indistinctly tomoderately distinctly demarcated from thepaler sapwood. The grain is straight, inter-locked or slightly wavy, texture coarse anduneven. The wood shows a conspicuous silver

grain figure on radialsurfacesCorol!to 670chioto yields a medium-weighthardwood with a density of 710-755 kg/ina at12% moisture content. The rates of shrinkageupon drying are low, c. 0.8% radial and 3.9%tangential for wood of Australian origin, andthe wood seasons well, but end splitting andsurface checking should be prevented by pro-tecting the ends from rapid drying; it takes 2months to air dry 13 min thick boards and 5monthsfor 38 min thick ones.

At 12% moisture content, the modulus of rup-ture is 117.5 N/mm2, modulus of elasticity13,855 N/mm2 and compression parallel tograin 55.5 N/min2 for wood of Indian origin.The wood is strong, easy to saw and plane, andit takes a good finish. Immediately after saw-ing the wood should be treated with anti-stainchemicals. To obtain the attractive silver grain,boards should be quarter-sawn, which limitstheir width to about 20 cm. The wood is dura-

ble under cover, but durability in contact withthe ground or when exposed to the weather ismoderate to poor. It is prone to termite andmarine borer attack, whereas the absorption ofpreservatives is moderate (95-130 kg/ing). Thesapwood is susceptible to Lyetus attack

Botany Shrub or smalltree up to 15 in tall(sometimes up to 50 in in tropical Asia); hole upto 40(-70) cm in diameter, occasionally withsmallbuttresses; bark surface smooth to finelycracking or shallowly to deeply fissured;branches ascending-erect, slightly thickened atnodes. Leaves decussately opposite, simple;stipules lanceolate, up to 2 cm long, early cadu-cous; petiole up to I cm long; blade narrowlyobonate to elliptical, 5-10(-15) cm x 2-5(-Toycm, curieate at base, acute or shortly acumi-nate at apex, margin revolute, entire or some-times slightly toothed, leathery, glabrous, withblack dots below, with many lateral veins. In-norescence an axillary condensed cyme, dis-tinctly resinous. Flowers bisexual, regular, (4-)5(-8)-merous, small, up to 2.5 min in diame-ter, sessile; calyx shortly cup-shaped with tri-angular, thick lobes; petals free, clawed atbase, Iaciniate at margins; disk annular; sta-mens twice the number of petals, free; ovarysemi-inferior, 5(-8)-celled, style thick, stigmaheadlike, obscurely lobed. Fruit a globose,fleshy berry up to 7 mm in diameter, pink tored when ripe, several-seeded. Seedling withepigeal germination; hypocotyl elongated; coty-ledons leafy, greenCording comprises about 10 species, of whichGurul!to 670chioto is the most widespread,

covering the whole distribution of the genusThe other species are confined to tropical AsiaInitial growth is slow and seedlings attain onlyup to 35 cm in height after 2 years and 2.5 inafter 5 years, However, seedlings of columnarornamental cultivars may reach 60 cm tall in10 months. Growth is monopodial and treesflower and fruit abundantly in mast fruitingyears, but individual trees may flower less pro-fusely in other years as well. The fruits, havinga pleasant, sweet-acid flavour, are attractive tobirds, which disperse the seeds.

Ecology In Madagascar Corer11io brachiotooccurs in humid evergreen forest from sea-levelto 1500 in altitude. In Australia it grows wellin open and wet localities, but it can also standquite dry conditions. In China Corolliobrochioto was found to be quite resistant toheavily polluted environments

Management Cordllio brachiotu may bepropagated by seed or cuttings. It seems thatseed soon loses its viahinty. In tropical Asiaseed had 45% to almost 100% germination inI-3.5 months. Seedlings may be kept in thenursery for 2 years before being planted out inthe field. Young plants are sensitive to droughtand tolerate shade; planting in open sites Isdifficult. Trees coppice well and reproducefreely from root suckers. In India techniquesfor rapid multiplication of Coro!lid brachiotoby terminal branch cuttings have been devel-oped; they include treatments with a fungicideand growth hormones

Genetic resources and breeding Carol!tobyochioto is widely distributed but nowhere

CARPODIPTERAAFRICANAMast.

Protologue 011v. , F1. trop. Mr. I: 241(1868)

Family Tiliaceae (APG: Malvaceae)Synonyms Berry0 @1ncono onast. ) Kosterm

(1969)Vernacular names ECalago (Po). Mwanga

maima, inlanga, inkikoma, inkongoro (Sw)Origin and geographic distribution In tropi-

calAfrica Corpodjptero of neono is distributedin Kenya, Tanzania, Mozambique and perhapsComoros. It also occurs in South Africa

Uses The wood of Corpodiptero of neono isused in construction and for poles, bows, toolhandles and spoons. It is also used as firewoodIn times offo0d scarcity in Tanzania the tenderleaves are cooked and eaten as a vegetable,alone or mixed with other vegetables. The barkyields fibre. In Tanzania an infusion of theroots and stem bark is drunk to treat eye prob-Iems and used as a face and eye wash. A rootdecoction is taken as an aphrodisiac

Properties The wood of Goryodiptero ofri-cono bends easily

Botany Dioecious or rarely monoecious shrubor small tree up to 13.5(-20) in tall; bark grey-brown, smooth or slightly rough; branchlets

glabrous.sparsely stellate-pubescent,Leaves alternate, simple; stipules fillform, upto 8 min long, soon deciduous; petiole up to 6cm long, stellate-pubescent; blade ovate to ob-long, up to 20 cm x 15 cm, base rounded tocordate or rarely curieate, apex acute to obtuse,margin entire or repand, steUate-pubescentwhen young but soon becoming glabrous, 3-5-veined from the base. Inflorescence an axillarycompound cyme, many-flowered; peduncle 2.5-7 cm long, stellate-pubescent, branches up to

long. Flowers unisexual, regular,2.5 cm

slightly scented; pedice1 0.5-1.5 min long; calyxcampanulate, divided for c. three-quarters ofits length into 2-3 lobes, lobes broadly triangu-Iar and 3-4 min long with acute or acuminateapex, densely stellate-pubescent; petals 5,obovate, 5-6 mm long, narrowed to the base,white or pink; male flowers with numerousstamens c. 4 mm long, filaments Joined at base;female flowers with numerous short sterile

stamens 2-3 min long and superLor, 2-lobedovary, style short, stigma large and spreading.Fruit an ellipsoid capsule up to 1.5 cm long, 2-lobed, each lobe with 2 wings 2-5 cm x I-2 cm,turning reddish or purplish brown with age,eventually breaking up, each lobe I'SeededSeeds ellipsoid, 5-8 min x 4-5 min, brown, very

CARPODiPTERA 131

common.

Prospects The supply of wood is small dueto the scattered occurrence of the trees andtheir often small size. However, more researchon the SIIvicultural aspects seems worthwhileas the wood is of good quality and utilitarianvalue. Ornamental cultivars are particularlysuited to planting along roads and close tobuildings, and deserve more attention. It maybe useful for restoration of degraded localitiesbecause of its tolerance to pollution

Major references Arenes, 1954; Ding Hou,1998; Schatz, 2001.

Other references Kwan & Whitmore, 1994;Vijaya, Murthy & Srivasuki, 1993

Authors R. H. M. J. Lemmens

Based on PROSEA 5(3): Timber trees: Lesser-known timbers

soon

132 TIMBERSl

finely wrinkled, more or less hairy near apex.Corpodiptero comprises about 5 species, allexcept Carpodjptero of neono occurring intropical America. The genus is closely relatedto Berryo and has even been included in thelatter genus, which differs in its 3-4-10bedovary and long styleIn Kenya Carpodipteru of neono flowers fromDecember to July

Ecology Carpodiptero of neono grows up to550(-900) in altitude in dry evergreen mixedforest, forest margins, bushland and woodedgrassland. It may form pure stands after burn-Ing.

Management Gulpodjptero officono ISIoggedfrom wild stands in Tanzania, and not plantedor locally protected. It coppices well. In Tanza-nia the leaves are collected during the rainyseason

Genetic resources and breeding It isunknown whether Guruodipter0 o17icono isaffected by genetic erosion. Its distributionarea is rather limited, but in Tanzania it iscommon in its habitat

Prospects Very little is known about theproperties and management of Coreodipteroof neono, and it is therefore impossible to makea valid assessment of its prospects

Major references Beentje, 1994; Ruffo,Birnie & TengnAs, 2002; Whitehouse at al. ,2001; Wild, 1963b.

Other references Capuron, 1963; Lovett ata1. , 2006; Neuwinger, 2000; Pakia, 2005; Wild& Gongalves, 1969

Authors M. Brink

7-14 cm x 3-6 cm, base rounded or curieate,apex acuminate, margin toothed to entire,glaucous, pale. Inflorescence an axillary fasci-cle, I-3-flowered, lax. Flowers bisexual, regu-Iar, 5-merous; pedice1 I-3 mm long; calyx cup-shaped, 3-4 mm long, with ovate-lanceolatelobes, glabrous outside, appressed hairy inside;petals deeply fringed, white; stamens 10; ovarysuperior, 3-celled. Fruit an indehiscent capsule,green, few-seeded. Seeds c. 2 min x 2 mm, witharil, testa leatheryGossipoureo comprises about 70 species, wldelydistributed in the tropics. Some other speciesare used for various purposes in West Africa.Gossipoureo firestoneono Hutch. & Dalziel exGP. Cooper & Record is a tree up to 28 in talland endemic to Liberia. Its wood is used for

planks, hut-poles and canoe paddles; it isbrownish yellow, hard, heavy, strong, flexibleand durable. GossipoL, rea ittolotou Aubr6v. &Penegr. occurs in Liberia and C6te d'TVoire. Itis a tree up to 30 in tall and easily recognizedby the characteristically coiled stem base. Itswood is considered suitable for construction

and interior joinery, but the high shrinkagerates and the unfavourable drying propertieslimit its usefulness.

Ecology Gossipoure0 o12elii occurs in wetevergreen forest, gallery forest, and secondaryforest. It occurs especially ln regions with anannual rainfall of over 2500 mm. It tolerates

shade and is usually found in the forest under-storey close to water. Gossipoure0 o12e!it oftenoccurs on sandy and sandy-loamy soils.

Genetic resources and breeding Cossi-poured o121eliiis considered rare

Prospects Gossipoure0 o12elii will remain alocal source of timber in Liberia. It is too un-

common to be exploited in larger amounts andis probably in need of protection.

Major references Burki11, 1997; Holingrenat a1. , 2004; Keay, 1954c

Other references Dudek, F6rster & mis-senbauer, 1981; Liben, 1987; Oliver, 1871.

Authors M. Brink

CAssiPouREAAFZELii(Onv. ) AISton

Protologue Bull. Misc. Inform. Kew 1925265 (1925)

Family RhizophoraceaeVernacular names Elephanttusktree (En)Origin and geographic distribution Cossi-

poured o12eliiis distributed from Guinea toGhana and from Nigeria to Cameroon

Uses The sterns of Gossipoureo Gleeliiare usedin Liberia for hut-poles, pickets, stockades andtraps for animals. In Sierra Leone the bark hasunspecified medicinal use.

Properties The stems of Gossipoure0 o12eliiare strong, tough and durable

Botany Slender-branched shrub or smalltreeup to 15 in tall. Leaves opposite, slinPIe, gla-brous; stipules between the petioles; petiole 4-7 mm long; blade broadly oblong to elliptical,

CASSIPOUREAEURYOIDES inston

Protologue Bull. Misc. Inform. Kew 1925:254 (1925)

Family RhizophoraceaeVernacular names Mwanzangu, mugome,

rigome (Sw).Origin and geographic distribution Cussi-

poured euryoides is distributed from Somalia to

.

Zimbabwe and Mozambique.Uses The wood of Gossipoureo euryoides is

suitable for heavy construction, flooring, shipand boat building, vehicle bodies, furniture,cabinet work, interior trim, joinery, poles andpiles, Implements, toys and novelties, turnery,carving, and veneer and plywood. It is alsoused as firewood and for the production ofcharcoal

Properties The heartwood is creamy whitewith purplish streaks and indistinctly demar-cated from the sapwood. The grain is straight,texture fine and even. The density is about 850kg/ina at 12% moisture content. The wood sea-sons well and rapidly, but protection againstdirect sunlight is necessary to prevent split-ting.The wood is easy to saw, but gum deposits mayclog sawteeth and sawing gives better resultswhen the wood is wet. Fine sawdust can cause

irritation. Although the wood is rather hard, itis easy to work with machine and hand tools,and it planes to a very smooth surface. It fin-ishes well without filler being necessary. Thewood is stable in service and takes glue, nailsand screws well. It is durable and fairly resis-tant to marine borers

Botany Shrub or smalltree up to 15(-18) intall, often with low-branching hole; bark brownto grey, somewhat corky; young branches usu-ally sparsely appressed hairy, later glabrousLeaves opposite, simple; stipules between thepetioles, 3-5 min long, early caducous; petioleI-6 mm long; blade lanceolate to elliptical orobovate, 3-10 cm x 1.5-5 cm, base curieate,apex acuminate to acute, margin entire or witha few teeth in the upper part, papery to Ieath-ery, glabrous above, sparsely hairy to glabrousbelow. Inflorescence an axillary fascicle, I-4-flowered, more or less lax. Flowers bisexual,regular, 4-6-merous; pedice1 I-2 min long,jointed at apex; calyx campanulate, tube I-1.5mm long, lobes 3-5 min long, narrowly triangu-Iar to ovate; petals spatulate, 3-6 mm long,deeply fringed, white to green; stamens 20-24;ovary superior, densely pubescent, 3-4-celled,style 2-2.5 mm long, persistent. Fruit a spheri-cal to ovoid capsule c. 5 min long, pubescent,dehiscent, few-seeded. Seeds black with redanI, testa leatheryGossipoureo comprises about 70 species, widelydistributed in the tropicsDropped seeds of Gossipoureo e!, ryoides, andoccasionally fruits, are carried by unidentifiedsmall ants to their nest; after the ants haveeaten the fatty anI, intact seeds are dispersed

outside the nest.

Ecology Gossipoureo euryoides occurs in dryevergreen or deciduous forest, woodland andsavanna, up to 800 in altitude. In Somalia it isfound on sandy soil.

Genetic resources and breeding Thereare no indications that Gossipoureo euryoidesis endangered by genetic erosion.

Prospects The wood of Gossipoureo euryotdesis of good quality. However, very little is knownabout this species, and it is therefore difficultto assess its prospects as a timber tree

Major references Beentje, 1994; Bolza &Keating, 1972; Friis, 1993; Lewis, 1956; Torre& Gongalves, 1978.

Other references Enge1, 2000; Gonealves &Torre, 1979; Kokwar0, 1993; Lovett, Ruffo &Gereau, 2003; Wimbush, 1957.

Authors M. Brink

CASsiPouREA 133

CASSIPOUREAGUMMIFLUA Tul.

Protologue min. Sci. Nat. , Bot. , s6r. 4, a 123(1856).

Family RhizophoraceaeVernacular names Large-leaved onionwood,

broad-leaved onionwood (En). Msikundazi(Sw).Origin and geographic distribution 0088i-

poureu gummi/Iuo occurs from Sierra Leoneeast to Kenya and south to South Africa, alsoin Madagascar

Uses The wood of Gossipoureo gummiftuo isa useful general purpose timber. It is suitablefor light construction, poles, masts, flooring,vehicle bodies, furniture and cabinet work,handles and ladders, boxes and crates, interiortrim, implements, Joinery, toys and novelties,turnery, veneer and plywood, and for hard-board and particle board. It is also used asfirewood and for the production of charcoal. InMozambique the wood of Gossipoureo gummif-IMO as well as that of Arithocleisto grandi/lordGIIg are known under the trade name'mezambe'Gossipoureo gummiftuo Is also LISed as an or-namental and as a shade tree. The bark is ex-

PIOited for unspecified medicinal uses.Properties The heartwood is greyish or

yellowish white and indistinctly demarcatedfrom the sapwood. The grain is straight, tex-ture moderately fine and even. Freshly-sawnwood has a strong onion-like smell, which per-SISts to some extent in the dry wood. The woodhas a density of 480-720 kg/ina at 15% ino1s-ture content. It is moderately hard and strong.It seasons slowly wlth a tendency to warp, and

134 TIMBERSl

should be dried under cover to avoid excessive

splitting. The wood is liable to blue stain attackand discolours easily.The wood is easy to saw and work with handand machine tools. It finishes to a fine surface

It turns, moulds and glues well, and can bepeeled or sliced satisfactorily. The durability ofthe wood is low. It is susceptible to termite andmarine borer attack. The heartwood is ex-

tremely resistant to impregnation, the sapwoodmoderately resistantThe alkaloid cassipourrine has been isolatedfrom dry twigs and leaves of Cussipoureogummi/jug. The much-cited report that inges-tion of Cussjpoureo gummi/IMO wood shavingsby poultry results in sex change of femalechicks is erroneous. It later appeared that theshavings concerned were from Funtumio ofri-cono (Benth. ) Stapf; the androgenic effect isdue to infection of these shavings with thesteroid-producing saprophytic fungus Fuso-rillm soloni

Botany Evergreen shrub or small to largetree up to 40 in tall; hole straight, up to 40(-60)cm in diameter; bark pale grey or brown,smooth with raised lenticels; young brancheshairy, later glabrous. Leaves opposite, simple,glabrous; stipules between the petioles, 2-4mm long; petiole (6-)8-15(-17) min long; bladeelliptical to oblong or ovate, 5-15(-21) cm x2.5-10.5 cm, base rounded to curieate, apexrounded to acuminate, margin entire to sinuateor toothed. Inflorescence an axillary fascicle,dense and congested, many-flowered. Flowersbisexual, regular, 4-6-merous; pedice1 I-4 mmlong, jointed in upper half; calyx tubular tocampanulate, tube 2-4 min long, lobes 0.5-1.5min long, semi-circular to deltoid; petals linear-spatulate, 4.5-6 min x I mm, deeply fringed,white; stamens 8-12(-14); ovary superior tohalf-inferior, 2(-3)-celled, glabrous to denselyhairy especially above the middle, style 2.5-6min long. Fruit an ellipsoid to spherical or obo-void capsule 6-11 mm x 4.5-7 min, blackish,glabrous or shortly hairy, dehiscent, few-seeded. Seeds ovoid, 3-6.5 min x 1.5 mm, darkred, with orange anI, testa leatheryCussjpoureo comprises about 70 species, widelydistributed in the tropics. Within Cussipoureogummi/Iu0 4 varieties are distinguished: var.gummi/Iuo, restricted to Madagascar; var.monnii(Hook. f. ex 011v. ) J. Lewis (synonym:Cussipoureo globro AISton), distributed fromSierra Leone to Angola; var. ugondensis (Stapf)J. Lewis (synonym: Gossipoureo ugondensis(Stapfj Engl. ), distributed from Gamero0n to

Kenya and Zambia; and var. Derticilloto(N. E. Br. ) J. Lewis (synonym: Cussipoureo uertt-ci!!oto N. E. Br. ), distributed from Cameroon toTanzania and SouthAfrica

The anI of Cussipoureo gummiftz, o is eaten bybirds and monkeys, which may disseminate theseeds.

Ecology Gossipot4reo gummiftuo occurs inrainforest, riverIne forest, swamp forest andmontane forest, up to 2600 in altitude

Genetic resources and breeding In viewof its wide distribution Gossipoureo gummiftt4ois not liable to genetic erosion

Prospects The wood of Gossipoureu gum-milluo will remain a useful local source oftim-her. Information on this species Is too scarce tojudge its potential.

Major references Bolza & Keating, 1972;Burki11, 1997; Liben, 1987; Scott, 1950; Torre &Gonealves, 1978

Other references Beentje, 1994; Bleher ata1. , 2003; Gonealves & Torre, 1979; Lewis,1955; Lovett, Ruffo & Gereau, 2003; Smith &Wells, 1978; Wright & Warren, 1967.

Authors M. Brink

CASSIPOUREAMALOSANA (Baker) inston

Protologue Bull. Misc. Inform. Kew 1925258 (1925)

Family RhizophoraceaeSynonyms Gossipoureo elliottii(Engl. ) inston

(1925), Gossipoureo congoe, ,sis auct. non DCVernacular names Pillarwood, common

onionwood, bastard onionwood (En)Origin and geographic distribution Cussi-

ino1080no occurs from eastern DRpoured

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Congo and Ethiopia south to South AfricaUses The wood of Gossipoureo inglosuno is

traded as pillarwood. It is suitable for construc-tion, flooring (especially heavy duty or indus-trial flooring), vehicle bodies, furniture andcabinet work, tool handles, ladders, sportinggoods, agricultural implements, joinery, sleep-ers, poles and piles, toys and novelties, bee-hives, turnery, and for veneer and plywood. Itslow durability, however, limits its applicabilityoutdoors. Nevertheless, Gossipoureo inglosonois an important timber tree in East Africa. Thewood is also used as firewood

The Loita Maasai people in Kenya eat theouter bark boiled with soup to gain strengthTea is made from the scraped inner bark; it isdrunk by women to help to remove the PIa-centa. In South Africa the bark is used as a

skin lightener, and to treat skin ailments andsunburn. The flowers are a good source of nec-tar for bees. Gossipoureo ing!o80no Is also usedas a shade tree

Properties The heartwood varies in colour,from whitish to brown, often with purplestreaks associated with fungal attack; it is in-distinctly demarcated from the sapwood. Thegrain is usually straight, but with a slight tomarked tendency to spiralling; texture fine andeven. Freshly cut wood smells like onion. Thewood has a density of 600-840 kg/in3 at 12%moisture content. Shrinkage rates are 3.5%radial and 8.0% tangential from green to 12%moisture content. The wood is refractory inseasoning due to high shrinkage and the pres-ence of spiral grain. It dries slowly and Is sub-ject to severe distortion, particularly twist;checking is slight. At 12% moisture content,the modulus of rupture is 93-123 N/mm2,modulus of elasticity 11,500-12,500 N/min2,compression parallel to grain 61-76 N/mm2,shear 9.8-18.5 N/min2, cleavage 16-79 N/mmradial and 30-1/4 N/min tangential, andJanka side hardness 5960-7345 N

The green timber may be difficult to saw as ithas a strong tendency to spring. Dry wood iseasier to work than green wood. The wood isresistant to wear and abrasion. It is moder-

ately easy to work using hand tools and hasexcellent machining properties, especially inmoulding, but it may have an appreciableblunting effect on tool edges. A good finish canbe obtained. The wood tends to split on nailing,and pre-boring is necessary. The wood may besomewhat troublesome to glue and is not suit-able for steam bendingThe durability is low. The wood is susceptible

to attacks by termites and Lyetus borers. Theheartwood is resistant to impregnation, thesapwood moderately resistant.

Description Evergreen shrub or small tolarge tree up to 45 in tall; hole up to 21 in long,straight, cylindrical, up to 60 cm in diameter,without buttresses; bark grey, greyish yellowor brown, smooth, inner layer of outer barkbright red on exposure, inner bark orange withwhite striations, exudate watery; crown small,rounded; young branches hairy, later glabrousLeaves opposite, simple; stipules between thepetioles, 2-5 min long; petiole 2-8 mm long;blade oblanceolate, lanceolate, elliptical, ob-long, ovate or obovate, 2.5-10 cm x I-6 cm,base curieate or obtuse, apex obtuse to acute oracuminate, margin toothed or rarely entire,shiny and glabrous above, dull and sparselypubescent to glabrous below. Inflorescence anaxillary fascicle, lax to congested, I-5(-8)-flowered. Flowers bisexual, regular, 4-5(-6)-merous; pedice1 2-6(-9) mm long, jointed nearapex; calyx with tube 0.5-2 min long, lobes

long,narrowly oblong-triangular, 3-6hairy outside; petals spatulate, 4-8 min long,deeply fringed, white, yellowish or greenish;stamens 15-20(-22); ovary superior, glabrous

CASSiPOUREA 135

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136 TIMBERSl

to apically hairy, 3-4-celled, style 3-5 min longand persistent. Fruit an ellipsoid to ovoid cap-SUIe 6-10 min x 4-6 min, black when dry, hairybut becoming almost glabrous, dehiscent, few-seeded. Seeds with anI, testa leathery.

Other botanical information Cussipoureocomprises about 70 species, widely distributedin the tropics. The wood of Gossipot4reo ridondoJ. Leonard ex F10ret, occurring in DR Congo,Rwanda, Burundi and Zambia in forest at1650-2500 in altitude, is also used for carpen-try.

The holes of many large Gossipoureu ino1080notrees are rotten inside

Anatomy Wood-anatointcal description OAWAhardwood codes):Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(9: vessels exclusively solitary (90% or more));13: simple perforation plates; (14: scalariformperforation plates); (15: scalariform perforationplates with :S 10 bars); 21: intervessel pits op-POSite; 22: intervessel pits alternate; 25: in-tervessel pits small(4-7 prn); 26: intervesselpits medium (7-10 pin); 31: vessel-ray pits withmuch reduced borders to apparently simple:pits rounded or angular; 32: vessel-ray pitswith much reduced borders to apparently SIm-pie: pits horizontal (scalariform, gash-like) tovertical(palisade); 41: mean tangential diame-ter of vessellumina 50-100 pin; 49: 40-100vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 62: fibres with distinctly borderedpits; 63: fibre pits common in both radial andtangential walls; 66: non-septate fibres pre-sent; 69: fibres thin- to thick-walled; (70: fibresvery thick-walled). Axial parenchyma: 76: axialparenchyma diffuse; 77: axial parenchyma dif-fuse-in-aggregates; 78: axial parenchymascanty paratrachea1; 93: eight (5-8) cells perparenchyma strand; 94: over eight cells perparenchyma strand. Rays: (97: ray width I-3cells); (98: larger rays commonly 4- to 10-senate); (102: ray height > I mm); (103: rays oftwo distinct sizes); 108: body ray cells procum-bent with over 4 rows of upright and/or squaremarginal cells; 109: rays with procumbent,square and upright cells mixed throughout theray; 1/5: 4-12 rays per min; 1/6: Z 12 rays permin. Mineral inclusions: 136: prlsinatic crystalspresent; 137: prismatic crystals in uprightand/or square ray cells(L. N. Banak, H. Beeckman & P. E. Gasson)

Ecology Gossipoureo inglosono mostly occursin dry or humid forest at 1000-3100 in alti-

tude. It is locally common, and in montaneforests in Kenya and Tanzania it is sometimesthe dominant species or co-dominant with Po-docorpus lotifo!jus (Thunb. ) Mirb

Propagation and planting Cosstpoureomaiosono is sometimes propagated by wild-lings collected from the forest.

Genetic resources In view of its wide dis-

tribution Gossipoureo inglosonu is not liable togenetic erosion.

Prospects Although the wood is not dura-ble, not easy to work and prone to deformationduring drying, Gossipoureo ino10sono is animportant local source of timber in East Africaand will probably remain so.

Major references Beentje, 1994; Bolza &Keating, 1972; Chudnoff, 1980; Friis, 1995b;Lovett, Ruffo & Gereau, 2003; Lavers, 1969;Lewis, 1956; Oxford Forestry Institute, 1997-2004; Torre & Gonealves, 1978; World Agrofor-estry Centre, undated

Other references Bryce, 1967; Dale &Greenway, 1961; Friis, 1993; Gonealves &Torre, 1979; InsideWood, undated; Liben, 1987;Maundu at a1. , 2001; Neuwinger, 2000; Scott,1950; Sommerlatte & Sommerlatte, 1990; Ta-kahashi, 1978; Tanzania Forest Division, 1967;vanWyk & Gencke, 2000; Wimbush, 1957

Sources of illustration Dale & Greenway,1961.

Authors M. Brink

CASSIPOUREARUWENSORIENSIS (Engl. )AISton

Protologue Bull. Misc. Inform. Kew 1925:263 (1925)

Family RhizophoraceaeOrigin and geographic distribution Cosst-

poureu ruwensortensis is distributed in Centraland East Africa, from the Central African Re-public and DR Congo east to Ethiopia, Kenyaand Tanzania.

Uses The wood of Gossipoure0 711wensoriensisis suitable for light construction, ship and boatbuilding, vehicle bodies, furniture and cabinetwork, musical instruments, boxes and crates,interior trim, toys and novelties, turnery, andfor veneer, plywood, hardboard and particleboard

Properties The heartwood is whitish withmauve markings Tn the core, and indistinctlydemarcated from the sapwood. The grain isstraight, texture fine and even. The density is810-900 kg/ms at 12% moisture content. The

.

wood is refractory in seasoning, but withproper care in stacking, surface checking is notserious. The wood is easy to work by hand andmachine tools, producing a fine surface. Ittends to split on nailing, making pre-boringnecessary. It glues, peels and slices well. Thewood is resistant to wear and abrasion

The durability of the wood is low. Logs are sus-ceptible to stain and insect attack, includingLyetus borers. They should be removed quicklyfrom the forest or be treated with insecticides

or fungicides. The heartwood is extremely re-SIStant to impregnation, the sapwood moder-ately resistant.

Botany Shrub or smallto medium-sized treeup to 25 in tall; bark smooth; young branchesdensely yellow pubescent, darkening and be-coming glabrous. Leaves opposite, simple; stip-u}es between the petioles, 6-7.5(-12) min x 2-3(-4) mm; petiole up to 10 min long; blade el-lipticalto obovate, (4-)7-16 cm x (2-)3-8 cm,base curieate, apex acuminate, margin toothed,glabrous above, shortly hairy to glabrous be-low. Inflorescence an axillary fascicle, (2-)4-15-flowered. Flowers bisexual, regular, 4(-5)-merous; pedice1 I-5 mm long, jointed in upperhalf; calyx with tube c. 1.5 min long, lobes 4-6.5 mm long, triangular, hairy outside; petals4-6 min long, deeply fringed, greenish cream,glabrous; stamens 16-20; ovary half-inferior,hairy, 3-4-celled, style 4-5 min long, persis-tent. Fruit an ovoid capsule 6-8 mm long,densely hairy, dehiscent, usually 2-seeded.Seeds 4-4.5 mm x 3 mm, testa leathery anddark red-brown, anIPink-orange.Gossipoureo comprises about 70 species, widelydistributed in the tropics

Ecology Gossipoureo ruuiensorienszs Is acommon understorey tree occurring in uplandevergreen forest or dense semi-swamp forest,at 500-2500 in altitude.

Prospects Little is known about Gossipoureoruwensoriensis and therefore its conservationstatus and potential as a timber tree of wlderusage are not known.

Major references Bolza & Keating, 1972;Lewis, 1956

Other references Beentje, 1994; Liben,1987; Wimbush, 1957

Authors M. Brink

CATHARIOSTACHYSMADAGASCARIENSIS

(ACamus) S. Dransf

Protologue Kew Bull. 53(2): 394 (1998)Family Poaceae (Gramineae)Synonyms Cepho!OStochyum mudogoscorien-

sisACamus (1925)Vernacular names Giant bamboo (En).

Bambou g6ant(Fr)Origin and geographic distribution Ende-

inIC to Madagascar, where it occurs mostly Inthe interior parts of the country

Uses The stems are used for construction

and they are locally made into musical instru-merits. They are also used as water pipes forirrigation systems; stem parts are used as wa-ter containers. Split and flattened stems arewoven into mats used for the construction offloors and walls of traditional houses

Properties Leaf bases and shoots containcyanide.

Botany Bamboo with elongated, long-neckedrhizomes; stem (CUIm) solitary, up to 15(-22) intall and 8(-12) cm in diameter, woody, hollow,glabrous or hairy, internodes 40-60 cm longwith moderately thin walls; young shoots paleto purplish green, covered with sharp, stiff,brown to black hairs; leafy branches up to I inlong, with up to 50 leaves. Leaves alternate,simple; sheath up to 21 cm long and up to 18cm wide at base, glabrous or brown hairy; auri-cles up to 5 min long with curly bristles, easilyshed; 11gule very short; blade triangular tolanceolate, (447-26 cm x 1.5-6(-11) cm, gla-brous but near the base covered with palebrown hairs. Inflorescence a globose,tracted, head-like panicle, 2.5-3 cm long, 2-4cm in diameter, much branched. Spikelets 20-22 mm long, consisting of (3-)5 glumes and Ifertile floret; lower glumes 3 mm long, 7-veined, upper glumes 7-11 mm long, 11-13-veined, with long pointed tips, lemma 12-13min x c. 4 min, with long-pointed tip, 13-15-veined, palea 17-20 mm x 4-5 mm; floret with6 stamens, an ovary with long style and 3stigmas. Fruit a cylindrical caryopsis (grain),with thin wall, not easily removedBecause of the long-necked rhizomes, Cothor-tostochys inoddgoscoriensis does not formclumps like many other bamboos do, but stemsgrow slrigly and scatteredCuthoriostochys comprises 2 species. The sec-ond species, Cothoriostochys copttoto (Kunth)S. Dransf. , is also endemic to Madagascar,where it occurs along the east coast. It hassterns up to 12 in tall, with a diameter of 4-5

CATHARIOSTACHYS 137

con-

138 TIMBERSl

cm. It is locally used for making flutes.Ecology Cothoriostochys mudogoscoriensis

occurs at 800-1000 in altitude in forest, forestmargins, disturbed forest or on open groundPopulations of Cothoriostochys inoddgos-curiensis sustain the lemur species Hopolemurgriseus, Hopo!emur aureus and Hopolemursimus, of which the latter 2 are criticalIy en-dangered. Despite its high cyanide content,this bamboo constitutes an important part oftheir diets.

Genetic resources and breeding It isunknown to what extent Cothoriostochys ing-dogoscoriensis is threatened by genetic ero-SIon, but it is locally common.

Prospects Little is known about the poten-tial uses, properties and ecological require-merits of Cothoriostochys ingdoguscoriensis.However, It does not seem to have good pros-pects for planting because it does not developclumps.

Major references Bystriakova, Kapos &Lysenk0, 2004; Dransfield, 1998

Other references Birkinshaw, 2005; Clay-ton, Harman & Williamson, 2002- a; Lehman,2006; Tan, 1999; Tan, 2006.

Authors M. Brink

rouge, acajou amer (Fr). Cedro (Po). Mwerezi(Sw)

Origin and geographic distribution Cedre!oodorotu originates from tropical America, butis now much cultivated for its timber and as an

ornamental or wayside tree throughout thetropics. It Is extensively planted in tropicalAfrica, in West as well as East and southernAfrica, and in Madagascar. Timber plantationshave been established in C6te d'Ivoire, Ghana,Uganda, Tanzania, Madagascar and SouthAfrica, and in several other tropical countriesIn Ghana Cedrelo odorato was planted alongroads as early as 1898, and is now one of themost frequently planted species in forest plan-tations. In Tanzania it was introduced in 19/1,in Nigeria in 1929. In C6te d'Ivoire more than9900 ha were planted between 1963 and 1995

Uses The best known use of the wood of

Cedre!o odorato (trade name: cedro)is for cigarboxes, but it is also used for light construction,joinery, mouldings, panelling, louvred doors,boat building, furniture, cabinet work,weatherboards, boxes, household implements,musical instruments, carvings, veneer, ply-wood, turnery and matchboxes. The insect-repellent smell of the wood makes it particu-Iarly suitable for the manufacture of clothingchests and wardrobes. Residue wood is used as

firewood and for charcoal production; in Ghanathe tree is planted in firewood plantations.Cedrelu odorato is commonly planted as anornamental tree, and particularly as a roadsidetree. It is also planted as a shade tree in e. gcocoa and coffee plantations. Flowering treesare a good source of nectar for honey bees. Intraditional medicine in Sao Tome the bark isused to treat malaria. In Tanzania the bark

dipped in hot water is applied to numb footsoles

Production and international trade In

many regions within the natural area of distri-bution of Cedrelo odoroto the timber is highlyprized and has been much overexploited. Ex-PIOitation has continued on a large scale overthe past 200 years in tropical America and inmany areas it is still exploited. In 2004 smallvolumes of Cedrelo odoroto logs were exportedfrom Mexico at an average price of Us$ 2071m3,and small volumes of sawn wood from Colom-

bia at an average price of Us$ 3221m3. Thetimber Is exported from several other tropicalAmerican countries, but then usually in mixedconsignments with other Cedrelo species. Intropical Africa Cedrelo odoruto timber is still oflimited importance, but exploitation of timber

CEDREIAODORATAL.

Protologue Syst. nat. ed. 10, a 940 (1759)Family MenaceaeChromosome number2n = 50.56Synonyms Cedre!o inertcono MRoem. (1846)Vernacular names Spanish cedar, West

Indian cedar, cigarbox cedar (En). C6dre aca-jou, codrela, acajou cede, acajou fernelle, acajou

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Properties The heartwood is pale cream incolour Immediately after sawing, turning pink-ish brown upon exposure, distinctly demar-cated from the narrow creamy yellow or palebrown sapwood. The grain Is usually straight,sometimes interlocked, texture moderatelycoarse. Fresh wood has a distinct cedar-likeodour. Sometimes the wood exudes gumThe wood is light- to medium-weight, with adensity of 260-525 kg/ina at 12% moisture con-tent. In plantations a large variability has beenobserved; wood of 14-year-old trees from C6ted'Ivoire had a density of 260-340 kg/ina at 12%moisture content, wood of 52-year-old trees ofTanzanian origin had a density of 385-480kg/in3 at 12% moisture content. The rates ofshrinkage may be low (e. g. 1.5% radial and2.2% tangential from green to 12% moisturecontent was reported from plantation-grownwood in Samoa), but they may also be fairlyhigh. For the wood of 14-year-old trees fromC6te d'Ivoire, shrinkage rates from green tooven dry were 2.5-3.2% radial and 6.4-6.7%tangential. The wood dries moderately fastwith a slight risk of checking and deformationAir drying to about 30% is recommended priorto kiln drying. Boards 25 mm thick take about2 months to air dry, boards 50 min thick about3.5 months. Once dry, the wood is stable in

results: a yield of 54% with a Kappa number of71; brightness was low.Bark extracts showed moderate In-vitro an-

tiplasmodial activity. In in-vivo assays in miceinfected with Plusmodium berghei the barkextract exhibited significant inhibition of theparasite multiplication at a very high oral doseof 1000 ing/kg per day. Gedunin, a terpenoidwith antimalarial properties present in thebark, may be involved. Wood extracts alsoshowed in-vitro antimalarial activity. In testsin Nigeria crude bark extracts exhibit signifi-cant activity against the maize weevil(Sitophi-Ius 2eomuis), a pest of stored cereals. Geduninisolated from Cedre!o odorato has shown an-tifeedant activity against the rice weevil(Sito-philus oryzoe)

Description Deciduous or evergreen,monoecious, medium-sized to large tree up to40(-50) in tall; hole branchless for up to20G25) in, up to 180(-300) cm in diameter,without buttresses or with low, blunt but-tresses at base; bark surface greyish brown toreddish brown, fissured, inner bark pinkishbrown; crown rounded; young branches withlenticels. Leaves alternate, panpinnately coin-pound with (5-)6-14(-15) pairs of leaflets;

CEDRELA 139

service

The wood is weak. For wood of Tanzanian on-

gin with a density of 450 kg/ina at 12% ino1s-ture content, the modulus of rupture was 54N/min2 modulus of elasticity 8100 N/min2,compression parallel to grain 35 N/mm2, shear8 N/min2 cleavage 49 N/min radial and 56N/min tangential, and Janka side hardness2050 N

The wood is easy to work. It saws well, butgrowth stresses may cause severe end splittingof logs and warping and splitting of the centralcore during saw milling. It bores, turns andsands well and produces a good finish; it iseasy to glue. It nails easily, but the nail-holding power is low. Rotary peeling and SIIc-ing give good results without pre-treatment,producing attractively figured veneer.The heartwood is rated as durable, but onlymoderately resistant to termites; the sapwoodis nori-durable and susceptible to staining andpowder-post beetles. The heartwood is usuallyresistant to impregnation with preservatives.Wood dust may irritate the respiratory organsand skin. Kraft pulping gave only moderate

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140 TIMBERSl

stipules absent; rachis slightly hairy or gla-brous; petiolules up to 2 cm long; leaflets ovateto oblong-lanceolate, 5-17 cm x 2.5-7 cm,asymmetric at base, acute or acuminate atapex, entire, glabrous, pinnately veined. Inno-rescence a terminal, much-branched, pendentpanicle up to 50 cm long, glabrous or slightlyhairy. Flowers unisexual, male and femaleflowers very similar in appearance, regular, 5-merous, fragrant; pedicel up to 2 mm long;calyx cup-shaped, c. 2 mm long; petals free, 7-9min long, creamy white, fused in lower half tocolumnar androgynophore; stamens free, 2-3min long; ovary superior, globose, glabrous, 5-celled, style I-3 mm long, stigma disk-shaped;male flowers with rudimentary ovary, femaleflowers with non-dehiscing, smaller arithers.Fruit an oblong-ellipsoid to obovoid capsule1.5-4(-7) cm long, pendulous, with lenticels,brown, dehiscing with 5 slightly woody valves,many-seeded. Seeds 2-3 cm long, pale brown,winged at apex. Seedling with epigeal germina-tion; cotyledons leaf-like; first leaves opposite,3-foliolate with entire leaflets.

Other botanical information Cedrelo coin-prises about 8 species, with a natural distribu-tion confined to the American tropics. It isclosely related to Toono from tropical Asia andAustralia. Toono enjoto MRoem. is planted asa roadside tree and shade tree in tropical at-Tica, occasionally in timber plantations, andhas often been confused with Cedrelo odoroto.It differs in its flowers without androgynophoreand in its lobed or toothed seedling leaflets

Anatomy Wood-anatomical description (IAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: (4: wood semi-ring-porous); (5wood diffuse-porous); 13: simple perforationplates; 22: intervessel pits alternate; 23?: shapeof alternate pits polygonal; 25: intervessel pitssmall(4-7 pin); 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 pin;46: S 5 vessels per square minimetre; (47: 5-20vessels per square minimetre); 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 65: septate fibres present; 66:non-septate fibres present; (68: fibres verythin-walled); 69: fibres thin- to thick-walled.Axial parenchyma: 76: axial parenchyma dif-fuse; 78: axial parenchyma scanty paratra-chea1; (79: axial parenchyma vasicentric); 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: (97: ray width I-3 cells);(98: larger rays commonly 4- to 10'seriate);106: body ray cells procumbent with one row ofupright and/or square marginal cells; 1/5: 4-12rays per mm. Mineral inclusions: (136: pris-matic crystals present); (137: prismatic crystalsin upright and/or square ray cells); (141: pris-matic crystals in non-chambered axial paren-chyma cells); (142: prismatic crystals in chain-hered axial parenchyma cells)re. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Seedlings growvery quickly and may attain 40-50 cm heightafter 3 months and 130-150 cm after 12

months. Early mean annual growth may be upto 2.3 in in height and 4.8 cm in diameter un-der favourable site conditions and when not

attacked by Hypsjpy!o shoot borers. In a plan-tation in Ghana, mean annual height and di-ameter increments decreased from 48 in and

5.4 cm, respectively, in the 2nd year to 1.4 inand 2.1 cm in the 15th year. In C6te d'Ivoire thebest provenances reach a mean diameter of 23-27 cm after 14 years and of 45-51 cm after 24years. A tree planted in Uganda reached 35 intall after 20 years. In the east Usambaramountains (Tanzania) 50-year-old trees were26-34 in tall, with a hole 14-21 in long and 40-50 cm in diameter. The root system is superfi-cial. First flowering can be expected after10-15 years. Flowering is annual, but goodseed production occurs every I-2 years. Theflowers are pollinated by insects such as beesand moths. Fruits ripen about 3 months afterflowering. The seeds are dispersed by wind.Association with vesicular-arbuscular my-corrhizae has been recorded.

Ecology In its natural area of distributionCedre!o odoroto occurs in primary and secon-dary forest, up to 1200 in altitude. It preferswell-drained sites on a variety of soils, but isusually more common on limestone-derivedsoil, especially in areas with a high annualrainfall (2000-3000 mm). It tolerates somedrought once the tree is well established. It isbest planted in fertile, well-drained soil, pro-viding good aeration required by the root sys-tern. In Uganda Cedrelo odorotu grows well inthe warm and moist climate near Lake Victo-

ria. Cedrelo odoroto is a light demander, andbehaves as a long-lived pioneer.

Propagation and planting Seeds are oftenproduced in great number, and can be collected

.

from the soil or from a canvas mat placed be-low a fruiting tree, but seed collected from ripefruits stillon the tree shows better germinationresults. The 1000-seed weight is 15-30 g. Thegermination rate offresh seeds is usually high,and seeds germinate 14-28 days after sowing.Well-dried seeds can be stored for more than a

year with fair retention of viabillty. Germina-tion of seeds kept in closed glass bottles at4-6'C was 82% after 2 months and 78% after14 months. Seeds are broadcast or sown inlines in level nursery beds and lightly coveredwith soil, sand, sawdust or charcoal. Wherethere is adequate moisture, shade is not neces-sary; it increases the risk of damping-off. Seed-lings are planted out when they are 30-40 cmtall. Direct seeding is feasible, as seedlingsdevelop very quicklySuccessful vegetative propagation of Cedre!oodoroto by air layering and cuttings is knownfrom West Africa. Stumps, striplings and con-tamer-grown seedhngs are used for plantingStumps 20 cm tall and with a diameter of I-2cm planted 10 cm deep showed nearly 100%survival in Indonesia. Wildlings are also usedfor propagation, and often show high survivalrates. Spacing is usually (3-)4-6 in x (3-)4-6

Management Trial timber plantations havebeen established in C6te d'TVoire, Ghana,Congo, Uganda, Tanzania, Madagascar andSouth Africa. Tests in Ghana showed that ap-PIication of 200 inI of 15:15:15 NPK fertilizersolution to seedlings in pots filled with sandyloam once in every I-2 weeks increased stemheight and diameter growth significantly; theoptimum concentration was 1.2-1.6 g/I. Addingcompost to the pots also had a positive effect onseedling growth. Weeding during the first yearis necessary. In enrichment planting it is jin-portant to ensure sufficient overhead lightCedrelo odoroto cannot be managed by CDPpic-ing. For plantations in C6te d'Ivoire with aninitial density of 1/11 treesA1a, it is recoin-mended that about 50% of the trees are

thinned out when they are about 10 in tall,having a basal area of 14 mayha. The secondthinning orrom 600 to 400 trees/ha) should bedone when the basal area is 15 in', the thirdone errom 400 to 200-250 treesA1a) when thebasal area is 16 in' and the last one (to thefinal density of 120-150 treesA1a) when thebasal area is 18 in2

Diseases and pests The moths Hypsipy!ograndello, occurring in America, and Hypsjpylorobusto, occurring in Africa and Asia, severely

damage plantations of Cedrelo odorato by at-tacking young shoots and seedlings. There Issome evidence that attacks are reduced byplanting under shade. Planting Cedrelo odo-roto in mixed plantationsis also recommendedCedrelo odorato grafted on Toono ciliato Isresistant to Hypsipy!o grondello. Heart rot isfrequent in trees with a large diameter

Yield A 40-year-old plantation in Nigeriayielded a timber volume of 455 matha. Insouthern C6te d'Ivoire the productivity Is 7-16matha/year

Genetic resources Cedrelo odorato is much

sought after for its timber in Its natural distri-bution area, and has become scarce in manyregions. Large trees of the form and size de-sired for exploitation are now rare. Cedreloodoroto is included in the TUGN Red list ofthreatened species as vulnerable. The popula-tions in Colombia and Peru are listed in CITES

Appendix 111. This means that trade in logs,sawn wood and veneer from these countries Isallowed only on presentation of appropriatepermits or certificates, but there are no jin-pediments to trading of wood from other coun-tries and from plantations. The planting ofCedre!a odoroto throughout the tropics coin-pensates to some degree for the high pressureon wild populations. In international prove-nance trials, more than 15 provenances arebeing tested in Africa, and breeding pro-grammes have started in C6te d'Ivoire, Ghana,Nigeria and Congo

Prospects Cedre!O Odoroto Is a promisingtimber plantation species because it grows rap-idly and yields a multipurpose timber. Thesuperficial root system is a drawback for plant-ing in agroforestry systems and its susceptibil-ity to Hypsipylu grondello should be consideredwhen using this species in timber plantationsin tropical America, where planting Cedre!oodoroto in mixed plantations, together withnon-Menaceous species, Is recommended.

Major references Bolza & Keating, 1972;Burki11, 1997; CAB International, 2005; Chungat a1. , 1995; CIRAD Forestry Department,2003; Katende, Birnie & Tengnas, 1995; Lamb,1968; Nwoboshi, 1997; Takahashi, 1978; WorldAgroforestry Centre, undated.

Other references Bena & Behaghe1, 1994;Bryce, 1967; Cobbinah, 2004; Corbineau, De-fresne & C6me, 1985; CTFT, 1962c; Do C6u deMadureira at a1. , 2002; Dupuy, 1988; DLLpuy ata1. , 1988; Durand & Kouassi, 1979; Ewete &mainu, 1999; Forest Products Research Labo-ratory, 1971; InsideWood, undated; Jouve,

in

GEDREiA 141

142 TIMBERSl

1984; Lamprecht, 1989; Neuwinger, 2000;N'Guessan, 1988; 0mar at a1. , 2007; 0moyiola,1972; Pennington, 1981; Styles & White, 1991;Tanzania Forest Division, 1963a

Sources of illustration Chung at a1. , 1995Authors R. H. M. J. Lemmens

Based on PROSEA 5(2): Timber trees: Minorcommercial timbers

CEDRELOPSIS GREVEIBaill

Protologue Grandid. , Hist. phys. Madagascar34(4), atlas 2, tab. 257 (1893).

Family RutaceaeVernacular names White panssander (En)

Katrafay, acajou blanc de Madagascar (Fr).Origin and geographic distribution Cedre-

lopsis greueiis endemic to western and south-ern Madagascar.

Uses The wood is locally used for heavy con-struction, carving, cabinet work, tool handles,Interior Joinery, interior trim, heavy parquetflooring, sliced veneer, plywood, ship and boatbuilding, railway sleepers, vehicle bodies, elec-tricity and construction poles and cattle enclo-sures. Because of its hardness and resistance

to fungal and insect attack, the wood is consid-ered imperishable and it is traditionally usedfor making royal Sakalava tombs. The wood isalso used as fuelwood and for making charcoalIn Madagascar Cedre!opsis greueiis one of themost important forest trees known for its me-dicinal uses. in essential oil from the bark is

commonly used in massaging to treat generalbody pain, toothache, broken bones, muscularpain, arthritis and rheumatism, and a massageof the back is given to treat tiredness and fe-

ver. It is also used in baths for these purposesIts tonic effects as well as its aphrodisiac ef-fects are well appreciated, as it is considered toimprove physical and mental fitness. A stembark extract is traditionally taken againstcough, asthma, tuberculosis, pneumonia, dia-betes, diarrhoea, abdominal pain, rheumatism,intestinal worms, headache, tiredness and as apost-partum tonic. It Is also used as a vaginalshower after childbirth for its tonic effects, andis externalIy applied to wounds and skin infoc-tions. Sometimes a root bark decoction is taken

to treat diarrhoea or asthma. A vapour bath ofthe leaves is taken to treat weakness of the

blood vessels, headache and a sore throat. Theseeds are chewed as an arithelmintic and totreat stomach-ache

The bitter and aromatic stem bark is used to

aromatise local ruin, and is also an ingredientof bitter, non-alcoholic drinks

Production and international trade In Ma-

dagascar and on Internet the essential oilfromthe bark is commonly traded at Us$ 8-17 per10 inI;'floral water' is traded for about Us$ 15per 250 inI. The main clients are the foreignand local pharmaceutical industry. The wood ofCedrelopsis greuei comprises 15-20% of thevolume of timber harvested from forests in

Madagascar, and is classified as the 4th mostimportant wood, equal to Daibergio spp. It iscalled 'white panisander' on the local marketSome half-finished products are exported toReunion

Properties The heartwood is pale yellow topale brown, somewhat inottled and slightlydarker than the 2.5 cm wide whitish sapwood.The grain is usually straight, texture fine. Thewood is scented and contains resin cells

The wood is very heavy, with a density of 900-1/10 kg/in' at 12% moisture content. It airdries slowly but well if well stacked; end-splitting sometimes occurs. Boards 25 minthick take 3-4 months to air dry, boards 40min thick 10-11 months. Shrinkage rates aremoderate: from green to oven-dry 3.6-5.6%radial and 5.3-8.8% tangential. Once dry, thewood is stable in service

The wood is flexible and very hard. At 10%moisture content, the modulus of rupture is122-255 N/min2, modulus of elasticity 12,400-17,500 N/min2, compression parallel to grain64-88 N/min2, shear 12 N/min2, Janka sidehardness 10,280 N and Janka end hardness10,280 NThe wood works fairly well with hand and ina-chine tools, but it has a marked blunting effect

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and stellite-tipped sawteeth are needed. Split-ting on nailing and screwlrig is common, andpre-boring is recommended. The wood glues,polishes, waxes, varnishes and paints well. It isreputed for its resistance to wood rot and insectattack. The sapwood is susceptible to Lyetusborers. The heartwood is very resistant to Im-pregnation.The chemical constituents of the essential o110f

Cedrelopsis greuei were found to be extremelyvariable depending on the location of collection.Analysis of five commercial samples exhibiteda wide variation in the contents of the maincomponents: ishwarane (1.0-17.4%), (E)-^-caryophyllene (1.3-12.5%), u-copaene (4.9-11.0%), p-elemene (0.2-9.6%) and u-sellnene(1.1-9.4%). The stem bark oil also containedthe dihydroagarofuran 4-epi-CIS-dihydroagaro-furan and the sesqulterpene Ishwarol B. Astudy analyzing the essential oilfrom the barkof trees from 6 locations found 4 chemotax-

onomic patterns, characterlzed by an eudes-mane skeleton, orpinene and copaborneol, co-paene and ishwarane, and cadinane skeletons,respectively. Variation between origins wasmuch greater than variation within originsThe major constituents of commercial samplesof oil from the stem bark were (E)-p-caryophyllene (9.3%), orcopaene (7.7%), orsennene (5.8%), 6-cadinene (4.9%), p-sellnene(4.5%), orhumulene (3.3%) and p-bisabolene(2.8%). The oils from bark and leaf were foundto have a similar composition, but the relativepercentages of some compounds notably dif-fored. The bark essential oilcontained ^-pinene(17.1%), cis-sesquisabinene hydrate (12.8%)and caryophyllene oxide (7%) as the main coin-ponents, whereas the leaf essential oil waslargely dominated by trans-^-fumesene(35.6%), p-pinene (12.8%), CIS-sesquisabinenehydrate (9.8%) and ar-curcumene (8.6%).Numerous coumarins were isolated from thestem bark. One of these, cedrecoumarln A,showed agonistIC activity on both or and ^-oestrogemc receptors as well as superoxidescavenging activity. The hexane extract of thestem bark furthermore yielded triterpenoids,11monoid derivatives, pentanortriterpenoids, ahexanortriterpenoid and quasslnoids. From theheartwood several chromones have been ISO-lated

The seed of Cedrelopsis greuei yielded UVango-letin, 5.7-dimethylpinocembrin, cardamonin,flavokawin B, 2'-methoxyhelikrausichalconeand the prenylated chalcones cedreprenoneand cedrediprenone

Treatment of rats with 80 ing/kg bark extractfor 4 weeks induced a progressive decrease inblood pressure, which was partly due to thepresence of coumarins. A crude stem bark ex-tract showed significant cicatrizing effect onskin ulcers of mice, as well as antibacterialactivity (e. g. against Stophy!ococcus o16iconsand Pseudomo, ,OS aeruginosa) and antifungalactivity (against Condid0 o16icons). It alsoshowed increased muscle contraction of mouseintestine, aorta and trachea.

Description Deciduous, monoecious or dio-ecious, medium-sized tree up to 28 in tall; holestraight, branchless for up to 9 in, up to 60(-120) cm in diameter; bark surface pale greyishto yenowish, rough; young twigs short-hairy.Leaves alternate, 12-20 cm long, panpinnatelycompound with up to 10 leaflets; stipules ab-sent; petiole 3-4.5 cm long; petiolules 1.5-5min long; leaflets alternate or opposite, ellipti-cal-oblong, 3-5(-8) cm x I-1.5(-3) cm, baseslightly asymmetrical, curieate, apex shghtlynotched, margins slightly wavy, densely gland-dotted, hairy, pinnately veined with 12-18pairs of lateral veins. Inflorescence an axiUarypanicle, short-hairy. Flowers unisexual or bi-sexual, regular, 5-meTous, aromatic; pedicel I-3 min long; calyx with triangular lobes 4-5 minlong, thick, densely short-hairy; petals free,

CEDRELOPSIS 143

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144 TIMBERSl

elliptical-oblong, 8-10 min long, apex pointedand curled inwards, pink to yellowish, outsideshort-hairy; male flowers with 5 free stamensshorter than petals, disk lobed, c. I mm long,ovary rudimentary; female flowers with 5 ru-dimentary stamens, disk small, ovary superior,ovoid, 3-4 min long, slightly 5-lobed, sparselyshort-hairy, 5-celled, style c. I mm long, thick,stigma 5-lobed; bisexual flowers with slightlyreduced stamens and ovary, and biologicallynon-functional. Fruit an ellipsoid capsule up to3 cm long, dehiscing with 5 woody valves,short-hairy to glabrous, brownish to black atmaturity, up to 12-seeded. Seeds ellipsoid, Iat-erally flattened, c. 2 cm long, with a thin apicalwing.

Other botanical information Cedre!opsiscomprises 8 species, all endemic to Madagas-car. It has been placed in Mengceoe and Ruto-cede. In the 1970s it was excluded from Ruto-

ceoe into a separate family Ptoeroxy!oceoe,together with Ptoeroxylon from East andSouthern Africa and later also Bottegoo fromEast Africa, but a more recent phylogeneticanalysis of molecular data indicated that it isbetter included in an enlarged Rutoceoe.The wood of several Cedrelopsis spp. is usedCedrelopsis gracilis J. -F. Leroy occurs in drywoodland of western Madagascar and is rela-tively rare. The wood is not very durable and isonly occasionally used for construction. A barkextract is traditionally taken to treat feverCedrelopsis microfo!to!oto J. -F. Leroy occurs indry woodland of northern, western and south-ern Madagascar. The wood is heavy andstrong, but only occasionally used for construc-tion purposes. A leaf decoction is taken byyoung mothers as a tonic. Cedre!opsis triooluisJ. -F. Leroy occurs in dry woodland of northernand western Madagascar. The wood is used forlocal construction, planks and poles.

Anatomy Wood-anatomical description OAWAhardwood 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;13: simple perforation plates; 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; 40: mean tangentialdiameter of vessellumina s 50 pin; 41: meantangential diameter of vessellumina 50-100pin; 49: 40-100 vessels per square minimetre;50: ;: 100 vessels per square milllimetre; 58gums 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. Axialparenchyma: (75: axial parenchyma absent orextremely rare); 89: axial parenchyma in mar-ginal or in seemingly marginal bands. Rays:96: rays exclusively uniseriate; 104: all raycells procumbent; 1/5: 4-12 rays per mm. Min-eralinclusions: 136: prismatic crystals present;142: prismatic crystals in chambered axial pa-renchyma cells; 143: prismatic crystals in fi-bres

(N. P. Monel, P. 06tienne & E. A. Wheeler)Growth and development Cedrelopsis

greuei grows slowly, up to 50 cm per year, toreach a height of 0.5-3 in when 7 years old. Itis estimated to need over 40 years to produce asinaU pole. In forests Morondovanear

Cedrelopsts greuei bears leaves for 130-210days per year between November and Septem-ber. It flowers annually, in September-November, but fruits irregularly, in Novem-her~January

Ecology Cedrelopsis greuei occurs in openwoodland, scrubland, secondary forest andseasonalIy dry forest, from sea-level up to500(-900) in altitude. It grows on a wide van-ety of soiltypes, often on red or yellow sandysoils, but grows taller in river valleys than onplateau soils

Propagation and planting Cedrelopsisgreueican be propagated by seed and by stem cut-tings. Fresh seeds have a high germinationrate when direct sown in the field.

Management The wood, stem bark andleaves of Cedrelopsis greuei extensivelyexploited on a local scale. As there is an in-creasing demand for the essential oil and forthe timber on the international market, thespecies is currently being overexploited. InAmbararata forest near Belo alllarge treeshave been felled for ship building, butthere areon average 17 seedlings and 10 saplings per hafrom natural regeneration. Cedrelopsis greueiis one of the tree species which are protectedduring land clearing operations. No informa-tion is available concerning its cultivation

Harvesting The stem bark and leaves areharvested from standing trees or from treesfelled for their timber

Genetic resources Cedrelopsis greueiiswidespread and still relatively common, andnot in danger of genetic erosion, although theincreasing pressure on the species for timberand medicinal purposes has led to local overex-PIOitation. It could become threatened in the

are

near future.

Prospects The current overexploitation ofCedre!opsis greuei for timber and medicinalpurposes calls for sustainable forest manage-merit practices to be put in place as well as thedevelopment of plantations. Domesticationprotocols need to be developed. More researchis warranted concerning the pharmacology ofthe isolated compounds from the wood, sternbark and leaves.

Major references Bolteau, Bolteau & A1-10rge-Boiteau, 1999; Ganzhorn & Sorg (Edi-tors), 1996; Gauvin, Ravaomanarivo & Sinadja,2004; Gu6neau, Bedel & Thie1, 1970-1975;Gunb-Fakim & Brendler, 2004; Leroy &Lescot, 1991; Parant, Chichignoud & Rako-tova0, 1985; Rietveld & Farazanamalala, 2008;Schulte, Rticker & Newe, 2006; Takahashi,1978.

Other references Bayala at a1. , 2003; Bolza& Keating, 1972; Capuron, 1967b; CavaUi ata1. , 2003; InsideWood, undated; Koorbanally ata1. , 2002; Koorbanally at a1. , 2003; Mulhollandat a1. , 2002; Mulholland at a1. , 1999; Mulhol-land at a1. , 2004; Mulholland at a1. , 2003; Mul-holland, Parel & Coombes, 2000; Rahelinoro,1994; Raivoarisoa, 1999; Rakotoarison at al. ,2003; Rakotobe at a1. , 2008; Rakotomalala,2004; Ramaromanana, 2001; Ranaivo at al. ,2004; Rasoanaivo & de in Gorce, 1998; Samisoa,1998

Sources of illustration Leroy & Lescot,1991.

Authors D. Dongock Nguemo

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CEIBA 145

A

CEIBAPENTANDRA (L. ) Gaertn.

Catbopentondro- wild gridp!onted

central Africa. The dispersal mechanism bywhich the tree may have crossed the AtlanticOcean is uncertain, but the fruits float andmight have been transported by ocean cur-rents. It has been argued that the tree wasdomesticated in West Africa, from where itspread to East Africa and Asia. It is now culti-vated all over the tropics, but mainly in South-East Asia, especially in Indonesia and Thai-land. In tropical Africa the tree appears to benative from Cape Verde eastwards to Chad andsouthwards to Angola. In addition, there arerecords of the species in 13 other countries inEast and southern Africa (including South of-rica) and the Indian Ocean islands, butthe treehas probably been planted in all other tropicalAfrican countries as well

Uses Getbo pentondro has two main uses,being an important source of fibre and of tim-her. Historically it has been most important asa source of kapok fibre, the floss derived fromthe inner fruit wall. Kapok fibre is used forstuffing cushions, pillows and mattresses, andfor insulation, absorbent material and tinder.The use of kapok fibre declined in the late 20thcentury after the introduction of synthetic sub-stitutes. However, there is a renewed interestin the potential of kapok, using new processingtechniques, especially in textile applicationsThe fibre may also have potential as a biode-gradable alternative to synthetic oil-sorbentmaterials, due to its hydrophobic-o1eophilicproperties.Currently, the main use of Getbu pento, Idro isas a source of timber. The wood (trade names:'fuma','ceiba')is mostly used in plywood manu-facturing, but also for making boxes and crates,

Protologue Fruct. sem. PI. 2: 244 (1791)Family Bombacaceae (APG: Malvaceae)Chromosome number2n= 72-88

Synonyms Eriodendron onfroctuosL, in DC(1824).

Vernacular names Kapok tree, silk-cottontree (En). Fromager, arbre a kapok, ouatier(Fr). Mafumeira, poi15.0, inufuma, arvore dasumaiima, kapoc (Po). Msufi(Sw)

Origin and geographic distribution Getbopentondro originated in the American tropics.Its natural distribution has been obscured byits widespread introduction after about 1500Although it has been described as introducedby humans in tropical Africa, there is no his-toncal evidence of such introduction, and thereis strong ecological, botanical and cytologicalevidence that the tree is native to western and

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and for lightweight joinery. Traditionally, en-tire trunks are hollowed out as dugout canoes,and the wood is used for lightweight furniture,utensils, containers, musical instruments, inor-tars, carvings and similar items. It is suitablefor insulation, wooden sandals, heels, rafts,floats, lifeboats, models, insulation and particleboard. The buttresses are made into doors,table tops, plates and traysThe wood is suitable for papermaking. A1-though it has been recorded to be used as fuel-wood in DR Congo, it is not considered verysuitable as such, because it only smouldersThe wood can be used for fumigating huts orclothes. Wood ash is used as kitchen salt and

for soap making. The bark is used for makinghut walls and doors and yields a gum and areddish brown dye. The leaves and shoots arefodder for goats, sheep and cattle. The leaves,flowers and young fruits are eaten cooked intosauces. The flowers are visited by bees, produc-ing an amber-coloured honey with a character-istic taste. Ash from the fruits is locally madeinto snuff.

A use that has attracted commercial interest is

as a source of seed oil, which has been used insoap, and pharmaceutical manufacturing. Theoil can also be used for Illumination, paintmanufacturing and lubrication. It has beenused for culinary purposes, but this is not ad-visable for health reasons. The seed cake is

good as feed or as fertilizer. The seeds areeaten roasted or they are pounded and groundinto mealor cooked in soup. They are, however,said to upsetthe digestionGetbopentondro finds wide application in Mri-can traditional medicine. The root forms part ofpreparations to treat leprosy. Pulverized rootsand root decoctions are taken against diar-rhoea and dysentery. Root decoctions are oxy-tocic. Macerations of the root bark are drunk

against dysmenorrhoea and hypertension. Theroot and stem barks are credited with emetic

and antispasmodic properties. Stem bark de-coctions are used in mouth washes for treatingtoothache and mouth problems, and are takento treat stomach problems, diarrhoea, hernia,gonorrhoea, heart trouble, oedema, fever,asthma and rickets; they are also applied onswollen fingers, wounds, sores, furuncles andIeprous macules. Bark extracts are consideredemetic; they are drunk or applied as an enemaMacerations of the bark are a cure for heart

trouble and hypertension, and are creditedwith stimulant and arithelminthic propertiesThe powdered bark is applied on wounds. Gum

from the bark is an astringent and is used totreat diarrhoea and as an abortifacient. The

leaves are credited with emollient and sedative

properties. They are used against scabies, diar-rhoea, fatigue and lumbago, and as an alter-atIve, laxative and abortifacient. Young leavesare warmed and mixed with palm oil to beeaten against heart problems. Pounded leavesare applied as a dressing on sores, tumours,abscesses and whitlows. Leaf sap is applied onskin infections, and drunk to treat mental ill-ness. Leaf macerations are drunk or used in

baths against general fatigue, stiffness of thelimbs, headache and bleeding of pregnantwomen. Leaf preparations are used as an eye-bath to remove foreign bodies from the eye. Adecoction of the leaves is applied to treat con-junctivitis and wounds in the eye, and is usedfor bathing and massage to treat fever. In vet-erinary medicine a decoction of the leaves isgiven to treat trypanosomiasis. The flowers aretaken to treat constipation, and flowers andfruits are credited with emollient properties.The powdered fruitistaken with water againstintestinal parasites and stomach-ache. Kapokfibre is used for cleaning wounds. The seed oilis rubbed in for treatment of rheumatism and

applied to healwounds.Getbo pentondro is planted as a wayside andshade tree. The tree is sometimes left by farm-ers after clearing the forest for agriculturalland, to serve as a shade tree for cultivation ofcrops such as coffee and tea. It is increasinglyplanted as an ornamental in subtropical areasThe kapok tree has sacred significance for localpeoples in many parts of the world, includingtropical Africa, where it often serves as a treeunder which meetings are held

Production and international trade Kapokfibre was an important commercial fibre beforethe Second World War, with Indonesia beingthe most important producer. After the SecondWorld War, Indonesian production decreasedand Thailand became the main producer, butlater Indonesian kapok production increasedagain from less than 30,000 t per year in the1960s to about 80,000 t per year in 2000-2004,whereas the annual kapok production in Thai-land remained stable at 40,000-45,000 t, ac-cording to FAO statistics. Most of the kapokproduced is not exported.In tropical Africa production and trade of Getbopentondro historically has centred on the ka-pok fibre. In the early 20th century, colonialforesters planted the tree widely in West at-Tica, as well as in East and southern Africa

Plantings were generally made along roads,providing the benefit of shade in addition tofibre production, and only a few orchards wereplanted for fibre production, e. g. in C6ted'TVoire, Burkina Faso and Ghana. During theinId-1900s, kapok was a valuable export prod-uct for many West African countries, but itsimportance declined from the 1960s onwardsCurrently, production and trade centres on thetimber, mainly for plywood production. Ply-wood manufacturing in Africa expanded fromthe 1960s as several countries pursued importsubstitution, and as world demand for plywoodincreased rapidly. Commercial use of Getbopentondro timber has risen with the decline ofmore valuable species. Exploitation relies al-most entirely on naturally occurring trees, al-though recently efforts have been made to es-tablish plantations in Ghana. In both Ghanaand Nigeria Getbo pentondro is considered a'lesser-known' or 'lesser-used' species, and itsfuture use is expected to increase. The woodhas a relatively low value in international mar-kets, but it is an important export product forseveral African countries. According to recentITTO statistics Ghana exported 61,000 inaGetbo pentondro plywood in 2003, at an aver-age price of Us$ 2931m3, 54,000 ms in 2004, atan average price of Us$ 2921m3, and 37,000 inain 2005, at an average price of Us$ 3011m3.Ghana exported coinparable volumes of Getbopentondro veneer during these years: 62,000ina at an average price of Us$ 2421m3 in 2003,52,000 ing at an average price of Us$ 2711m3in2004, and 48,000 in3 at an average price of Us$2741m3 in 2005. In 2005 the average price ofCeibo pentondro logs in international marketswas Us$ 1561m3, while the export price of sawnwood from Ghana was Us$ 2461m3. 'Ceiba' wasby far the least expensive veneer for plywoodexterior and core stock in Ghana in 2005. In2006 it accounted for 19.6% of Ghana's timber

exports, the largest percentage of any of the 87species exported that year. Local and nationalmarkets for Getbo pentondro timber are weak;the average log price in Ghana's domestic mar-ket was Us$ 31-551m3 in 2005, while the aver-age sawn wood price was Us$ 531m3

Properties The wood is creamy white,streaked with yellow, brown, pink, and/or grey,but discoloration due to fungal attack is ex-tremely common; the heartwood is not clearlydemarcated from the sapwood. The grain Isinterlocked, sometimes irregular, the texturecoarse. Growth rings are prominent. The silicacontentis low

The wood is lightweight, with a density of(200-)240-380(-450) kg/ina at 12% moisturecontent. The density of the wood increasesstrongly from pith to bark, and varies betweentrees from different sites. The wood seasons

rapidly, with only slight risks of cupping,springing, twisting or bending. The rates ofshrinkage from green to oven dry are 2.5-3.5%radial and 3.9-6.7(-9.1)% tangential. The driedwood is stable in service, making it ideal forplywood core stockThe wood is particularly weak and soft. At 12%moisture content, the modulus of rupture is26-61 N/min2 modulus of elasticity 2300-5600N/mm2, compression parallel to grain 14-26N/mm2 shear 2-4 N/mm2, cleavage 4-13N/mm, Janka side hardness 1060-1/10 N,Janka end hardness 1820-1960 N and Chalais-Meudon side hardness 0.4-1.0. The wood isbrittle when dryThe wood is difficult to saw cleanly and finishsmoothly. Sawn surfaces tend to be woolly, andsharp cutters are necessary to achieve asmooth, finished surface. The blunting effect oncutting edges is low. Shaping, boring, turningand mortising often tear the grain, but planingand sanding produce fine results. The woodholds nails and screws poorly, and must beglued for fastening. Fresh logs, iffree of insectsand fungal decay, are easy to peelinto sheetsfor plywood manufacturing. The wood can beglued satisfactorily and takes paint very wellIt has been recorded to cause itching in work-ers handling it.The wood has very low durability, being liableto attacks by insects and fungi. It is amongstthe most vulnerable African timbers to ter-mites, Lyetus beetles, and other boring insectsIt is susceptible to white-rot and blue-stainfungi, but resistant to very reslstant againstbrown rot. The wood is extremely vulnerable todecay when in contact with the soil. It readilyabsorbs preservatives: both vacuum-pressureand open tank systems give good penetrationand absorptionThe wood contains cellulose 39%, pentosans18%, 11gnin 24% and ash 4%. The solubility inalcohol-benzene is 2.7%, in hot water 8.8%. Thewood fibre cells are 2 min long and 30-35 pinwide. Pulp yields are low.The total dry matter of ripe fruits, by weight, Lscomposed of 48% shell, 21% fibre, 25% seedand 6% placenta. Kapok fibre cells are (8-)19-22(-35) min long and (10-)19-20(-30) prn wide,smooth, transparent, cylindrical, with a widelumen and thin walls. Kapok fibre contains

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about 43% u-cellulose, 32% hemicellulose, 13-15% Iignin and I% ash. It is resilient, elastic,light (8 times lighter than cotton), water-repellent and buoyant (5 times more thancork). In an uricompressed state, it can support20-30 times its own weight in water, becausewhen the fibre is immersed in water, the In-men only partly fills with water and containsmany air bubbles. Kapok fibre has a low ther-inal conductivity and very good sound-absorbing properties. A disadvantage is itshigh inflammability, but techniques have beendeveloped to make it non-flammable. In itsnatural form, kapok fibre cannot be spun, be-cause of the smoothness of the outer surface

However, techniques have been developed tomake spinning possible. The fibre is long-lasting and is not attacked by fungi or pestsWhite-coloured fibre is preferred. Kapok fibreIs Irritating to the eyes, nose and throat, andworkers exposed to kapok dust for long periodsmay develop chronic bronchitis. People in-volved in processing of kapok fibre are advisedto wear protective masks. Because of the ITri-tant fibres and the spreading root system, thetree is less suitable as shade or wayside tree.The seeds yield 11-28% oil, with as main fattyacids: palmitic acid (10-16%), stearic acid (2-9%), oleic acid (49-53%) and linoleic acid (26-29%). The oil resembles cotton-seed oil, butdoes not contain gossypol. It does contain cyclo-propenoid fatty acids such as inalvalic acid (7-8%) and sterculic acid (3-4%), which causeabnormal physiological reactions in animalsTherefore the consumption of kapok seeds orseed oil should be discouraged, unless thecyclopropenoid acids have been chemically re-moved. Per 100 g the presscake contains ap-proximateIy: moisture 14 g, protein 26 g, fat 8g, carbohydrates 23 g, fibre 23 g and ash 6 gThe K content of the shellis about 3%, andthat of the ash 20-23%.

The composition of leaves from C6te d'Ivoireper 100 g dry matter was: energy 486-631 kJ(1/6-151 kcal), protein 19.1-26.5 g, fat 2.2-2.7g, carbohydrate 8.0-8.6 g, fibre 55.1-62.8 g, Ca649-1200 ing, Mg 440-452 ing, Fe 7.0-8.3 ing,carotene 22.7-25.5 ing, riboflavin 0.60-0.73 ingand ascorbic acid 148-250 ingA root bark extract has shown in-vivo hypogly-caemic activity in rats. Bark extracts haveshown in-vivo and In-vitro anti-inflammatoryactivity, In-vitro arithelmintic activity, andcurare-like activity anaesthetized caton

nerves. Bark decoctions and sesquiterpeneIactones isolated from the root bark have

shown antimicrobial activity. Leaf extractshave shown in-vitro activity against sickle cellanaemia

Adulterations and substitutes Getbo pen-tondro wood is sometimes sold in mixed con-

signments with that of Bombox buonopozenseP. Beauv. , but the wood of the latter is darkerand heavier. Ceiba wood is heavier than balsa

(Ochromopyromidule (Cav. ex Lain. ) Drb. ), butlighter and weaker than obeche (Triplochitonscleroxylon K. Schum. ). It is not as lustrous asbalsa or obeche

Kapok fibre is obtained from several other Af-rican trees, especially Bombox buonopozenseP. Beauv. and Bomberx costotum Fellegr. &Vuillet. Fibre obtained from the red silk-cotton

tree (Bomberx ceibo L. , synonym: Bomberxmoldboricum DC. ) has many of the qualities ofthat of Ceibo perilondro, but is less resilient,has a more brown or yellow colour, and cansupport only 10-15 times its own weight inwater. For industrial purposes kapok fibre haslargely been replaced by synthetic products, forinstance for the production of life-jackets

Description Very large, deciduous tree upto 60 in tall, with roots spreading quite hori-zoritally, 10 in or longer, in the upper 40-80 cmof the soil; hole branchless for up to 35 in,straight, usually cylindrical, up to 200(-240)

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Getbo pentondro - I, tree habit, 2, floweringtwig, 3, fruit.Redrown und udopted bylshuh Syomsudin

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cm in diameter, usually with large plank-likebuttresses up to 3(-8) in high extending I-2 infrom the hole; bark surface smooth, pale grey,usually with scattered conical spines I-1.5 cmlong, inner bark pale to pinkish red with verti-cal stripes of white or yenowish tissue; crownof emergent trees in the forest spreading and

lateral branchesumbrella-shaped withlower along the hole, crown of trees in moreopen vegetation ovoid with lateral branchesalong much of the hole; branches whorled, di-inorphic, branches usually in whorls of 3, hori-zoritalor ascending, older branches often spinyLeaves alternate, digitately compound, with 5-9(-15) leaflets; stipules linear to narrowly tri-angular, up to I cm long, caducous, leavingscars; petiole (3.5-)5-27 cm long, thickened atbase, apex expanded Into an almost circulardisk; petiolules up to 3 cm long; leaflets sessile,obovate to elliptical, (3.5-)5-21 cm x (I-)2-6.5cm, curieate at base, acuminate at apex, entireor sometimes slightly toothed at margin, gla-brous, pinnately veined with lateral veins in 7-18(-20) pairs. Inflorescence an axillary fascicle,

leaness branches, I-15-floweredoften

Flowers bisexual, 5-merous, regular, stronglybut unpleasantly scented; pedice1 2-5(-8) cmlong; calyx campanulate, 9-15 mm x 11-14mm, lobes rounded to nearly acute, glabrousoutside, hairy inside; petals united at base,oblong-spatulate, 22-46 mm x 6-13 mm, pink,creamy yellow or white, hairy outside, glabrousinside; stamens 5-15, united at base in a sta-minal column, dividing into 5(-6) branches2.5-5 cm long; ovary semi-inferior, 3-6 min indiameter, 5-celled, style 2.5-4 cm long. Fruit apendulous, oblong-erripsoid capsule 7.5-30(-60)cm x 3-15 cm, debtscing with 5 valves ('shells')or indehiscent, smooth-valved, brown whenripe, many-seeded. Seeds almost globose, 4-6min in diameter, glabrous, dark brown orblack, embedded in copious white or greyishfloss. Seedling with epigealgermination

Other botanical information Ceibo coin-

prises about 17 species, most of them occurringin tropical America. Getbo pentondro is van-able. Sometimes 3 varieties are distinguished.

curtboeo (DC. ) Bakh. , occurring in theforests of the American tropics and in WestAfrica; very large tree up to 70 in tall, withunforked, buttressed and spiny hole andhorizontal branches, leaves narrow, flower-ing irregular, fruits rather short and broad,dehiscent, fibre grey to white; 2n = 80, 88.

gumeensis (Schumach. & Thonn. )_ var.

H. GBaker, occurring in savanna woodlands

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of West Africa; small to medium-sized treeup to 18 in tall, bole spineless without but-tresses, often forked, branches strongly as-cending, leaves broad, flowering annually,fruits elongated and narrow at both ends,dehiscent, fibre grey; 2n = 72

- var. pentondro, the cultivated kapok of WestAfrica and Asia; medium-sized tree up to 30in tall, hole unbranched, usually spineless,buttresses small or absent, branches hori-zorital or ascending, leaves intermediate be-tween the 2 other varieties, flowering annu-ally after leaf-shedding, fruits short or long,narrowed at both ends or banana-shaped,usually indehiscent, fibre usually white; 2n =72-84. It would be more appropriate to clas-SIfy this variety as a cultivar groupAnatomy Wood-anatomical description (IAWA

hardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 pin); 27:intervessel pits large G 10 prn); 31: vessel-raypits with much reduced borders to apparentlysimple: pits rounded or angular; 35: vessel-raypits restricted to marginal rows; 43: mean tan-gentialdiameter of vessellumina ;a 200 pin; 46S 5 vessels per square minimetre; (47: 5-20vessels per square minimetre). Tracheids andfibres: 61: fibres with simple to minutely bor-dered pits; 66: non-septate fibres present; (68fibres very thin-walled); 69: fibres thin- tothick-walled. Axial parenchyma: 77: axial pa-Tenchyma diffuse-in-aggregates; 79: axial pa-renchyma vasicentric; (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: 98: larger rays commonly4- to 10-senate; (99: larger rays commonly >10-senate); 102: ray height > I mm; 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; 110: sheathcells present; 1/4: S 4 rays per min. Stonedstructure: 120: axial parenchyma and/or vesselelements stoned. Mineral inclusions: 136

prismatic crystals present; 137: prismatlc crys-tals in upright and/or square ray cells; (138:prismatic crystals in procumbent ray cells);141: prismatic crystals in non-chambered axial

on

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150 TIMBERSl

parenchyma cells.or. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Withoutanypre-treatment seeds germinate slowly (less than10% one month after sowing) and germinationmay continue for 3-4 months. Bush fire maycause simultaneous germination of seedsGrowth is relatively fast. Seedlings planted inGhana were 29 cm tall 6 weeks after germina-tion, and 63 cm after 51 weeks. The annualincreases in height and diameter during thefirst 10 years are about 1.2 in and 3-4 cm, re-spectively. In forest gaps height growth may be2 in/year. The tree is obligateIy deciduous, 10s-ing its leaves for 10-14 weeks in the dry sea-son, and it usually flowers annually in the leaf-less period. Leafing and flowering periods aremore regular in drier parts of the distributionarea; in moister areas, leafing and floweringperiods are highly irregular. The flowers openat night and are seriescent by midday; they arepollinated by bats, but are also visited bymoths and bees. The fruits ripen 80-100 daysafter flowering, the dehiscent types releasingkapok with loosely embedded seeds that arewind-dispersed

Ecology Getbopentondro requires abundantrainfall during the vegetative period and adrier period for flowering and fruiting. Itthrives at elevations below 500 in, but it growsup to altitudes as high as 4000 in. Night tern-peratures below 17'C retard germination of thepollen grains, which limits the area in whichthe trees can be grown for the fruit fibre tolatitudes between about 20'N and 20's, andaltitudes below 1500 in. Rainfall should be

about 1500 mm per year. In its area of naturaldistribution, the average annual rainfall is750-3000 mm. The dry period should not havemore than 4 months, and in this period a well-distributed rainfall of 150-300 min is required.In drier areas, some of the water demand maybe met by groundwater. For best results thetree should be planted on deep, permeable soilswithout waterlogging. The tree is easily darn-aged by strong winds, and it does not survivefire

Getbo pentondro occurs in rainforest, and ingallery forest in drier areas. It Is a pioneer insecondary forest and riverme forest, and israrely found in primary forest. It grows fast incanopy openings caused by disturbance, becoin-Ing an emergent in mature stands. In drierparts of its range, the tree is increasingly rarein natural vegetation, but may be abundant insettlements and fields in humid to semi-and

zones. In these settings, it may be planted, orself-sown seedlings may benefit from humanprotection against livestock and fire. Through-out West Africa, Getbo pentondro is associatedwith occupied and abandoned settlements

Propagation and planting Ceibo penton-dru is usually propagated by seed, although itcan also be grown from cuttings. The 1000-seedweight is 20-100 g. The seeds are easily sepa-rated from the fruit floss, in which they areloosely held, by shaking dehiscent fruits in abag. Seeds may be stored up to one year inglass or plastic containers at 4"C and 60% rela-tive humidity. Long-term seed storage has notproven successful, because the seed oil goesrancid. Fresh seeds have germination rates of90-100% within 3-5 days after sowing, whenpre-treated by scoring lightly and soaking inwater for 24 hours or by soaking in boiling wa-ter for 5 minutes. Germination is good in sandysoil with temperatures of 20-30'C. When theyoung plants are 12-15 cm tall, they can beexposed to full sunlight. Ceibo pentondro islight-demanding, and growth is spindly andpoor and mortality is high for seedlings andsaplings in shaded locations, including smallcanopy gaps that close relatively quicklyYoung plantscan be grown in a nursery and betransplanted into the field when they are 4-10months old. It is recommended, however, tosow directly on land which has been properlycleared for planting. Ceibo pentondro is easilypropagated from cuttings, which should betaken from orthotropic branches. Trees raisedfrom seeds root deeper than those raised fromcuttings, but develop slowerRecommended planting distances are 4.5-5 inx 4.5-5 in, with elimination of every secondrow after 6 years, to arrive at a spacing of 10 inbetween rows. Seeds may also be sown at about7 in spacing between trees, with an understo-rey of crops until the canopy closes, about 5years after planting. In the first 2 years afterplanting, vegetation must be cleared periodi-cally around saplings. General tending may benecessary in the following years, by cuttingclimbers and lianas, and removing dead anddiseased trees. Plantations need not be thinned

ifplanted with 7 in spacing unless intercroppedwith smaller tree crops.

Management Getbupentondro requires littleattention, but the soil must be occasionallyweeded and loosened. Fertilizers are not usu-

ally applied and there is no information onfertilizer experiments. Pruning is not requiredThe tree performs best in locations where the

,

vegetation is actively managed to maintainsunny conditions and to reduce climber burdenon the trees. Coppicing, pollarding and loppingis possibleIn forests in Ghana the number of trees with adiameter above 90 cm is about 35 per kin2. Innatural forests in C6te d'TVoire and Cameroon

average standing volumes of 6-8 in3ftia havebeen measured, in Gabon standing volumeswere only 0.2-0.6 inalba.

Diseases and pests High seedling and sap-ling mortality may occur in humid climates asa result of leaf spot, dieback, damping off andarithracnose. These infections are caused byvarious fungal pathogens. Tests have shownthat leaf spot and arithracnose are caused byConetotrichum copsici, whereas FMSorzum so-10ni and Lostodiplodio theo6romoe were asso-ciated with dieback of stem. The most effective

fungicides for preventing these infections areKocide (Cu-hydroxide, at 6.6 gA) and inlette(Aluminum tris(ethyl phosphite), at 5 g/I), al-though these fungicides retard the growth ofseedlings. Catbo pentondro is a host tree ofCacao Swollen Shoot Virus (CSSV) causingswollen shoot disease in cocoa, a disease whichhas had a devastating effect on cocoa produc-tion in Ghana and neighbouring countriesGetbo pentondro itself shows considerable re-SIStanCe.

Harvesting The tree is usually felled aboveits buttresses, which may necessitate the con-struction of platforms. To obtain kapok fibre,the fruits are harvested when fully ripe and, indehiscent types, before they open. Ripeness isindicated by the fruit colour changing fromgreen to brown and the surface possibly becoin-ing wrinkled. The fruits are normally bar-vested by knocking them off the tree before.Trees normally start to bear fruit when theyare 3-8 years old.

Yield A tree 70 cm in diameter above the

buttresses yields on average 4 ina of timber,and trees 100 and 150 cm in diameter abovethe buttresses 9.3 in3 and 23 in3, respectively.Under optimum conditions a full-grown planta-tion tree may yield 330-400 fruits per year,giving 15-18 kg fibre and about 30 kg seed. Asatisfactory average annual fibre yield is about450 kg/ha, whereas about 700 kgftia is consid-ered very good.

Handling after harvest After the trees havebeen felled, the wood is liable to attacks byinsects and fungi, and logs must be treatedwith preservatives, extracted, sawn and driedas soon as possible.

To obtain kapok fibre, fruits are hulled as soonas possible after harvesting. Drying is carriedout in dry air or with fans in cage-like struc-tures. The seeds lie loose in the floss, andtherefore de-seeding is easy. They are usuallyseparated from the floss by beating. The qual-ity of kapok fibre is judged by fibre length,freedom from seeds and foreign matter, ino1s-ture content, colour, smell and lustre. Exces-SIve pressure should not be used when kapok isbaled for export as it is likely to destroy theelasticity and diminish the quality of the fibre

Genetic resources In view of its pantropi-cal distribution, Getbo pentondru does notseem to be threatened with genetic erosionThe tree is traditionally protected in manyparts of Africa, and is legally protected in sev-eral countries, especially in drier parts of itsrange. However, commercial timber harvestingof Ceibopentondro is considered unsustainablein some areas, including C6te d'Ivolre andGhana. The largest kapok germplasm conec-tion is kept at the Indonesian Tobacco and Fi-bre Crops Research Institute in Malang (Indo-nesia) and contains 180 accessions. Some ac-cessions are held at CNSF (Centre National deSemences Forestfores) in Ouagadougou (Burk-Ina Faso).

Breeding Some work has been done on theselection and propagation of trees with whitefloss, indehiscent pods and lacking spines. Mi-crosatellite markers have been developed andcharacterized, providing a tool for rapidly gen-erating information on patterns of geneticvariation, gene flow and mating systems.

Prospects Timber industries in Africa havebecome increasingly Tenant upon Ceibo pen-tondro in recent decades, and use of the treewill probably intensify in the near future asmany African nations have shifted toward lessvaluable, historically underutilized species.While Getbo pentondro has clear value }n ply-wood manufacture, Increased rates of harvest-ing should not be accepted uncritically for twomain reasons. First, although the use of kapokfibre has declined in recent decades, this re-source has high potential as new processingtechniques are being developed. Use of the treeas timber should not preclude development ofimproved fibre industries. Second, in the longterm natural regeneration may be insufficientto sustain increased use as a source of timber,while minimal efforts have been undertaken to

develop plantations in tropical MrlcaMajor references Baker, 1965; Bolza &

Keating, 1972; Burki11, 1985; Chudnoff, 1980;

CEIBA 151

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CIRAD Forestry Department, 2003; CTFT,1975; Gibbs & Semir, 2003; Sabid & Zeven,2003; Takahashi, 1978; Voorhoeve, 1979

Other references A1varado, A1varado &Mendoza, 2002; Apetorgbor, SIaw & Gyimah,2003; Attah, 2005; Beentje, 1989; Beentje &Smith, 2001; Brondani at a1. , 2003; Fortin &Pollquin, 1976; Herzog, Farah & Amado, 1993;InsideWood, undated; ITT0, 2001; Lamprecht,1989; Lim & Huang, 2007; Mpiana at a1. , 2007;Neuwinger, 2000; 0T Africa Line, 2007;Poorter at a1. , 2004; Rendle, 1969; Roussel,1995; Toledo-Aceves & Swame, 2007; Wiemann& Williamson, 1989; World Agroforestry Cen-tre, undated

Sources of illustration Beentje, 1989; Sabid& Zeven, 2003

Authors CS. Duvall

lets; stipules lanceolate, I-5(-11) cm long,folded along inIdrib, reddish pubescent, ratherlong persistent; petiole 10-80 cm long, grooved,glabrous; petiolules up to 5.5 cm long; leafletselliptical to oblong-elliptical, up to 60 cm x 20cm, entire, base curieate, apex acuminate, gla-brous, reddish brown but pink when young.Inflorescence a fascicle old branches,on

wrapped in ovate bracts c. 8 mm x 6 minFlowers unisexual or bisexual, regular, sessileor almost sessile; calyx campanulate, with c.1.5 cm long tube, 5-10bed, greyish outside, red-dish or purplish inside; corolla absent; maleflowers with c. 2 cm long calyx, androphore3.5-6 mm long carrying a whorl of 15-25 ses-SIIe arithers surrounding rudimentary carpels;female and bisexual flowers with c. 2.5 cm longcalyx, ovary superior, consisting of 7-14 car-pels surrounded by a whorl of rudimentary orfunctional arithers c. 3 mm long. Fruit consist-ing of 9-12 erect fomcles arranged in a whorlaround a disk-shaped receptacle; follicles ob-long-ellipsoid, curved, 6-15 cm x 3-6 cm, witha short and recurved beak, orange or red,densely shortly hairy, filled with a viscous Iiq-uid, indehiscent, 7-25(-30)-seeded. Seedsovoid, lateralIy compressed, 2-3 cm x I-1.5 cm,scarlet, albuminous. Seedling with epigealgermination; hypocoty1 8-10 cm long, epicotyl1.5-3 cm long; cotyledons ovoid, c. 7.5 cm x 5cm; first leaf entireOhiomydocolo comprises 2 species, one onlyknownfrom Gabon

In C6te d'TVoire and Ghana Chiomydoco!ochiomydontho flowers from September-November, and fruiting is from December-February

Ecology Chiomydocolo chiomydontho occursin the understorey of deciduous and evergreenforest, including secondary forest, often in rela-tirely humid locations such as along water-courses. In Central Africa it is also recorded

from swamp forestManagement Ohiomydocolo chiomydontho

coppices well. It is sometimes considered aweed in silvicultural operations. Chinmydocolochiomyduntho is a host tree of Cacao SwollenShoot Virus (CSSV) causing swollen shoot dis-ease in cocoa, a disease which has had a devas-tating effect on cocoa production in Ghana andneighbouring countries. Control of this diseasein cocoa includes elimination of Chiomydocolochlomydontho from the vicinity

Genetic resources and breeding In viewof its wide distribution, Ohiomydoco!a chlomy-dontho is notthreatened by genetic erosion.

CHLAMYDOCOLACHIAMYDANTHA(K. Schum. )MBOdard

Protologue Journ. Agr. Trop. Bot. Appl. I313 (1954)

Family Sterculiaceae (APG: Malvaceae)Synonyms Cold chinmydontho K. Schum

(1900)Origin and geographic distribution Chiu-

mydoco!o chlomydontho occurs from Guineaand Sierra Leone to the Central African Repub-Iic and DR Congo.

Uses The wood of Chiomydocolo chiomydon-tho is used for pestles, house construction, andas firewood. The sour-sweet fruit pulp is edible.The seed cotyledons are chewed as an inferiorsubstitute of those of the true kola (mainlyGolo ocuminoto (P. Beauv. ) Schott & Endl. andColo nitido ryent. ) Schott & Endl. ). Mucilageprepared from young branches or fruits is usedin sauces. Decoctions of the bark are taken to

calm intestinal pain in Cote d'Ivoire, andagainst cough in Central Africa. The colourlessliquid from the fruit cavity is used to treat eyeinflammation in Central Africa.

Properties The heartwood is yellowishbrown; the sapwood greyish. The texture iscoarse. The wood is rather hard and heavy. Theliquid from the fruit cavity has been shown toinhibit seed germination in allspecies tested

Botany Smallto medium-sized tree up to 20in tall; hole straight, up to 30 cm in diameter;outer bark brown, smooth, flaking, inner barkc. 0.5 cm thick, yellow when cut, fibrous.Leaves alternate, grouped at the ends ofbranches, digitately compound with 5-10 leaf-

,

Prospects In view of the relatively smallsize of the stem, the role of Ohiomydocolochiomyduntho as a source of timber will re-main limited. Moreover, as a host of the devas-tating Cacao Swollen Shoot Virus it is unlikelyto become a populartree in West atrLca.

Major references Burki11, 2000; Germain &Bamps, 1963; Han6, 1961; Irvine, 1961; Kryn &Fobes, 1959

Other references Ak6 Assi at a1. , 1985;Bodard, 1954; Bouquet & Debray, 1974; Cheek,2002b; Keay, 1958e; Miege & Miege, 1970;Neuwinger, 2000; Schroth at a1. , 2000; Vivien& Eaure, 1989-

Authors M. Brink

CHLOROXYLONFAHO Capuron

Protologue Adansonia, sat. 2, 7(4): 479(1967).

Family RutaceaeVernacular names Satinwood (En). Faho

(Fr)Origin and geographic distribution Chlo-

roxy10, I 10ho is endemic to north-eastern Mada-gascar

Uses The wood of Chloro, cy!on lofto is usedfor local construction and carpentry. It is suit-able for the same purposes as Chioroxylo, ,swietenio (Roxb. ) DC. ('East Indian satinwood'),such as furniture, panelling, interior Joinery,parquet flooring, cabinet work, carvings, lux-ury goods and veneer.

Production and international trade Thewood is only used locally and not traded on theinternational market.

Properties The heartwood is pale yellow to

golden yellow and is difficult to distinguishfrom the 4-6 cm wide sapwood. The grain Isstraight, sometimes interlocked, texture veryfine. The wood is inottled and lustrousThe wood is heavy, with a density of 915-1020kg/in3 at 12% moisture content. It air driesrather slowly, but without much risk of distor-tion. The rates of shrinkage from green to ovendry are moderate: 3.4-5.6% radial and 6.8-8.4% tangential. Once dried, the wood is un-stable in service

The wood is hard and flexible. At 12% moisturecontent, the modulus of rupture is 147-220N/mm2 modulus of elasticity 10,700-18,650N/mm2 compression parallel to grain 60-86N/mm2 and Chalais-Meudon side hardness 8.2-12.8

The wood works well with hand and machinetools, but with a rather high blunting effect oncutting edges; stentte-tipped sawteeth areneeded. The wood takes a nlce polish. In gluingthere is a risk of discoloration of the wood. Thewood has an excellent resistance to fungi, drywood borers and termites. The heartwood isimpermeable to preservatives

Description Deciduous small tree up to15(-25) in tall; hole straight, branchless for upto 7(-15) in, up to 45(-70) cm in diameter; barksurface pale grey to yellowish, slightly rough tosmooth. Leaves alternate, panpinnately coin-pound with up to 16 leaflets, hairy whenyoung; stipules absent; petiole up to 3 cm long;petiolules short; leaflets alternate, erriptical-ovate, 2-5 cm x I-3 cm, base very asymmetri-cal, one side curieate, other side cordate, apexrounded to acute, margins entire, gland-dotted,pinnately veined with up to 7 pairs of lateralveins. Inflorescence a terminal pyramid-shapedpanicle up to 30 cm long, short-hairy. Flowersbisexual, regular, 5-merous, small; pedicelshort; calyx with small, ovate teeth; petals free,narrowly lanceolate, 2-3.5 min long, pointed,whitish, reflexed, inside slightly short-hairy;stamens 10, free, as long as petals; disk obconi-cal; ovary superior, ovoid, short-hairy, 3-celled,style minute, stigma head-shaped, small. Fruitan erripsoid capsule c. 3 cm long, dehiscingwith 3 woody valves, up to 6-seeded. Seedsellipsoid, lateralIy flattened, c. I cm long, witha thin apicalwing up to 1.5 cm long.

Other botanical information Ohioroxyloncomprises 3 species; 2 are endemic to Mada-gascar and the third, Ohioroxylon swietenio,originates from central and southern India andSriLanka

Chloroxylon foieotum Capuron is a smalltree

CHLOROXYLON 153

I.

Chioro, cylon faho - wild

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154 TIMBERSl

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parenchyma in marginal or in seemingly mar-ginal 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 min. Stoned structure: 1/8: all raysstoned; 120: axial parenchyma and/or vesselelements stoned. Mineral inclusions: 136:

prismatic crystals present; 137: prismatic crys-tals in upright and/or square ray cells; 140:prismatic crystals in chambered upright and/orsquare ray cells; 142: prismatic crystals inchambered axial parenchyma cells; 143: pris-matic crystals in fibres.(N. P. Monel, P. Derienne & E. A. Wheeler)

Growth and development Chior0;10.10n loftois leafless from January-March. It flowersmainly in April

Ecology Chioroxylon faho grows in low-alti-tude humid evergreen forest in north-easternMadagascar, from sea-levelup to 500 in altitude

Genetic resources Although Chioroxy!onfaho is not very widespread or common, thereare no Indications that it is threatened by ge-netic erosion.

Prospects As the wood properties of Ohio-rotylon 10ho are similar to those of Chioroxylonswietenio, a wider use of the timber seems POS-sthle. However, in view of its restricted distri-bution and uncommon occurrence, harvestingthe timber from wild stands should be discour-

aged. Planting experiments may clarify thepossibilities as a timber plantation tree.

Major references Capuron, 1967a; Gu6-neau, 1971; Gu6neau & Gu6neau, 1969; Gu6-neau, Bedel & Thie1, 1970-1975; Parant,Chichignoud & Rakotova0, 1985.

Other references Bedolla, 1997; Inside-Wood, undated; Schatz, 2001

Sources of illustration Capuron, 1967aAuthors G. H. Schmelzer

\

^;b

Ohioroxylon lofto - I, flowering twig, . 2, flower, .3, fruits, . 4, seedRedrown und odopted by Ishoh Syomsudin

up to 15 in tall, endemic to western Madagas-car. The heavy and hard yellowish wood ismainly used for local construction and is suit-able for parquet flooring and carpentry. Thewood is considered of less aesthetic value thanthat of Chloroxylonfuho

Anatomy Wood-anatorntcal description OAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); 24: intervessel pits minute (S 4 prn); 25:intervessel pits small(4-7 prn); 30: vessel-raypits with distinct borders; similar to intervesselpits in size and shape throughout the ray cell;41: mean tangential diameter of vessellumina50-100 prn; 48: 20-40 vessels per square inil-limetre; (58: gums and other deposits in heart-wood vessels). Tracheids and fibres: 61: fibreswith simple to minutely bordered pits; 66: non-septate fibres present; 69: fibres thin- to thick-walled; (70: fibres very thick-walled). Axialparenchyma: 78: axial parenchyma scantyparatrachea1; 86: axial parenchyma in narrowbands or lines up to three cells wide; 89: axial

3 .\

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CHLOROXYLONSWIETENIA (Roxb. ) DC.

Protologue Prodr. I: 625 (1824)Family RutaceaeVernacular names East Indian satinwood,

Ceylon satinwood (En)Origin and geographic distribution Chio-

roty!on suietenio originates from central andsouthern India and Sri Lanka. In Africa culti-

vation has been tried in e. g. Nigeria and

MadagascarUses Chioroxylon swietenio provides a deco-

rative timber, used for furniture, paneUing,pattern making, interior trim, cabinet work,flooring, boxes, crates, Interior joinery, carv-ings, toys, musical instruments and luxurygoods. It is made into decorative veneer, whichis, however, unsuitable for plywood manufac-ture because of its weight. Because of itsstrength it is also used for heavy construction,railway sleepers, boat building and agricul-tural equipment. The wood is also used as fu-elwood

In India most plant parts are medicinalIy usedThe crushed leaves are applied to treatwounds, snakebites and rheumatism. A barkextract is considered astringent and taken totreat fever, chest pain and in a mixture withother plants to treat asthma. In friction it isused to treat bruises and painful joints. A pasteof the leaves and roots Is taken internaUy totreat headache and is applied to the foreheadas a balm for the same purpose. In Sri Lankathe root bark in milk is drunk to treat impo-tence

Production and international trade Ohio-

roxylon swietenio wood is traded internatlon-ally, e. g. on Internet, primarily for interiordesign and architectural inlay. It is expensiveand supplies are limited.

Properties The heartwoodcoloured to golden yellow, darkenLng to brownwith age; it is not clearly demarcated from theslightly paler sapwood. The grain Is usuallyinterlocked or wavy, texture fine and even. Thewood is often striped or inottled, and remarka-bly lustrous. Dark gum veins are a commondefect, as they are liable to develop Into splits.The wood has a slight pleasant fragrance whenfreshly cutThe wood is heavy, with a density of 900-980kg/in" at 12% moisture content. Air drying re-quires care to avoid cracking, twisting or warp-ing. Kiln drying may produce good results withlittle degrade, but is rather slow. The rates ofshrinkage from green to oven dry are moder-ately high: 5.5-5.9% radial and 7.1-9.0% tan-gential.The wood is strong and hard. At 12% moisturecontent, the modulus of rupture is about 174N/mm2 modulus of elasticity 14,900 N/mm2,compression parallel to grain 60 N/mm2, cleav-age 20 N/min and Chalais-Meudon side hard-ness 9.2

The wood is difficult to work with hand toolsand moderately hard to saw and machine, with

a moderate to severe blunting effect on cuttingtools. In planing a cutting angle of 15* is re-quired to avoid tearing. A neat finishing makesit possible to obtain a perfectly polished surfaceand a beautiful glazed effect. Boring operationsare reported to be rather difficult, and thewood requires to be held firmly in boring op-erations to prevent chattering. Preboring Isrecommended in nailing and screwing, butnail-holding and screw-holding properties arerated as good. The wood turns and stains well,but is difficult to glueThe heartwood is rated as highly resistant toattack by all types of fungi, but in India it isreported to possess little resistance to attack bytermites. It is also susceptible to attack by ina-rine borers, and to a lesser extent by pinholeborers and longhorn beetles. The sapwood Isnot susceptible to Lyetus borers. The heart-wood is extremely resistant to preservativetreatment. The wood is reported to cause skinIrritatlOn

From the stern bark coumarins (swietenone,swietenocoumarins A-F, 8-prenylnodakenetinand demethylluvangetin) and quinolinone al-kaloids (swietenidins A and B) were isolatedFrom the heartwood 2.4-dihydroxy-5-prenycin-namic acid was isolated, as well as an alkaloid(skimmianine) and coumarins (swietenol, xan-thyletin, xanthoxyletin, 7-demethylsuberosinand alloxanthoxyletin). The essential oil fromthe leaves contains as main compounds theterpenes Iimonene, germacrene D, generene,pregeijerene, trans-^-ocimene and methyleugenol. Both the leaf and the stem oils exhib-ited moderate to strong activities agalnst apanel of bacteria and fungi. The seeds contain16% non-drying oilEssential oil from the leaves and stems andseveral isolated compounds displayed signifi-cant mosquitocidal activity by fumigationagainst Aedes oregypti, Anophe!es gombioe andCulex quinquefosciotus, as well as activltyagainst tobacco cutworm, Spodoptero lituro.Methanol extracts of dried leaves exhibitedgood analgesic activity in mice.

Botany DeciduoussmaU to medium-sized treeup to 18(-25) in tall; hole straight, cylindrical,branchless for up to 4.5 in, up to 45(-90) cm indiameter; bark surface yellowish or pale grey-ish brown, rough and corky; crown spreading;branchlets greyish hairy. Leaves alternate,partpinnately compound, up to 24 cm long wLth20-40 leaflets, hairy when young; stipules ab-sent; petiole up to 3 cm long; petiolules short;leaflets alternate, oblong, 1.5-3 cm x 0.5-1.5

CHLOROXYLON 155

Is Cream-

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156 TIMBERSl

cm, base very asymmetrical, one side curieate,other side slightly cordate, apex acute, marginsentire, gland-dotted, pinnately veined with upto 5 pairs of lateral veins. Inflorescence a ter-minal or axillary pyramid-shaped panicle up to15 cm long, short-hairy. Flowers bisexual, regu-Iar, 5-meTous, small; pedicel c. 5 min long; ca-Iyx with deltoid lobes, c. 3 mm long; petals free,elliptical, up to 3.5 mm long, slightly clawed,fringed, white, spreading, short-hairy on bothsides; stamens 10, free, c. 2 mm long; disk 10-lobed, thick, densely short-hairy; ovary supe-nor, ovoid, slightly 3-10bed, sparsely to denselyshort-hairy, 3-celled, immersed in disk, styleshort, slender, stigma head-shaped, small.Fruit an ellipsoid capsule 2.5-3 cm x c. 1.5 cm,dehiscing with 3 woody valves, up to 12-seededSeeds ellipsoid, lateralIy flattened, c. I cm long,with oblongwing on one side, up to 1.5 cm long.Chioroxylon comprises 3 species; 2 are endemicto Madagascar. Chloroxylon suitetentu flowerswhen it is leafless, in the dry season, and canbe found fruiting 2-3 monthslater

Ecology In its natural habitat Chioroxylonscoretenio occurs in dry deciduous forest onpoor, well-drained sandy or rocky soils, at lowto medium altitudes. It occurs in regions withan annual rainfallof 750-1500 min

Management Chloroxylon suietenio hasshown good growth rates in Nigeria. It coppiceswell. In India Chioroxylon $00ietenio is an al-ternative food-plant for the caterpillars ofPupilio demoleus, a pest of Citrus

Genetic resources and breeding Chio-Foxyloit 800ietenio has become scarce in severalregions in India and Sri Lanka because oftim-her exploitation, and it is listed as vulnerablein the IUCN Red list of threatened species

Prospects The decorative wood of Ch!oroxy-10n slutetenio Is of excellent quality and willcontinue to be in demand for luxury goods,pattern making, interior turn, interior joinery,carvings, toys and musical instruments. Itwould be worthwhile to explore the possibilitiesof small-scale plantations in Africa

Major references Bolza & Keating, 1972;Burki11, 1997; Kiran & Devi, 2007; Stone, 1985;Takahashi, 1978

Other references Asian Regional Work-shop, 1998a; Bhide, Mujumdar & Ra0, 1977;Capuron, 1967a; Farmer, 1972; 1<1ran, Devi &Reddy, 2008; Kiran at a1. , 2006; Mujumdar,Rathi & Ra0, 1977; Parrotta, 2001; Sentil &Ramkumar, 2003; Upadhyay & Chauhan,2001

Authors G. H. Schmelzer

CHRISTIANAAFRiCANA DC.

Protologue Prodr. I: 516 (1824)Family Tiliaceae (APG: Malvaceae)Origin and geographic distribution Chris-

tmn0 o17icono is widely distributed in tropicalAfrica, from Senegal to Kenya, Tanzania andAngola, and in Madagascar. It also occurs intropical America

Uses The wood of Christtuner of neono ismade into axe-handles, stools and basins. Theleaves are used for wrapping cola nuts andfoods to be sold in the market. In Ghana theyserve as bait in fish- and prawn-traps. In theCentral African Republic a maceration of theleafis used in washings against diarrhoea andpalpitations, and an infusion of young leaves isdrunk hot to treat dysentery. Bark prepara-tions are used in West and Central Africa totreatstomach problems and gonorrhoea.

Properties The sapwood of Christiono ofri-curio is whitish yellow to pale brown, theheartwood darker with purplish blue ripplemarks. The wood is heavy, hard, resilient anddurable

Three compounds that have cardiotonic activityhave been Isolated from the bark and wood: thepolyphenol SCOPoletol, the triterpene taraxeroland the peptide alkaloid frangulanine

Botany Dioecious shrub or smalltree up to14 in tall with stellate hairs; hole short, with adiameter up to 60 cm; bark brown, rough, moreor less fissured, fibrous; crown spreading;branchlets shortly brown-hairy. Leaves alter-nate, simple; stipules fillform, c. 6 min long;petiole up to 26 cm long, shortly hairy; bladeovate to oblong, 9-32 cm x 5-28 cm, base cor-date, apex acuminate, margin entire or slightlywavy, glabrous or slightly stellate-pubescentabove, stellate-pubescent below, palmatelyveined with 5-9 pairs of lateral veins of which2-3 pairs basal. Inflorescence an umbel-likecyme, axillary or terminal, up to 22 cm long,many-flowered, shortly hairy; peduncle 5-13cm long; bracts fillform, 4-6 mm long. Flowersfunctionally unisexual, regular; pedice1 2-5mm long; calyx campanulate, 3-5-10bed, lobesovate and c. 3 min long; petals 5-9, oblong-spatulate, 4-7 mm x 2-3.5 mm, white, gla-brous; stamens numerous, fused at base, sterilein female flowers; ovary consisting of (2-)5-7free carpels, styles 5-7, fused for most of theirlength. Fruit consisting of (2-)5-7 ellipsoidfontcles 8.5-13 mm x c. 8 mm, shortly brownhairy, each splitting into 2 boat-shaped valvesand I'Seeded. Seeds almost globose, c. 6 min x

5.5 mm, marbled. Seedling with epigeal germi-nation; cotyledons thin, rounded, 3-veined fromthe base; first leaves simple, alternate, toothed.Christiano comprises 5 species, of which onlyChristtoner of neono occurs in tropical AfricaThe seeds of Christiono of neono usually ger-inmate 15-20 days after sowing. In C6ted'TVoire Christion0 @1ncuno flowers in Juneand fruits in October-February; in Kenya itflowers in July

Ecology Christiono ofricono occurs in hu-inid evergreen to dry semL-deciduous forest,often on the banks of streams, up to 400 inaltitude.

Genetic resources and breeding In viewof its extremely wide distribution Chr, stionoof neono is not threatened by genetic erosion.Research on the genetic variability may shedlight on the origin and spread in areas as farapart as tropical America, continental Africaand Madagascar

Prospects Very little is known about theproperties and management of Christtung ofri-curio, but there are no indications that its rolewill become more importantthan that of a localsource of wood, wrapping material and tradi-tional medicines. There are many alternativesources of the constituents isolated.

Major references Beentje, 1994; Burkill,2000; Whitehouse at a1. , 2001.

Other references Ak6 Assi at a1. , 1985;Bouquet & Debray, 1974; de in Mensbruge,1966; Keay, 1958f; Kubitzki, 1995; Normand,1955; Normand & Paquis, 1976; Payen-Fattaccioli, 2004.

Authors M. Brink

suitable for construction, carpentry, stairs,light flooring, mine props, ships, vehicle bodies,agricultural implements, sporting goods, Trite-nor trim, toys and novelties, railway sleepers,particle board and pulp for paper production.The edible, pleasantly acidulous fruit is oftensold on markets. The latex from the bark andyoung fruit is used as birdlime. A bark infusionis used in traditional medicine as a stomachicand carminative. A maceration of the stembark is taken to promote lactation. Powderedbark is applied to sores. The fruit is used totreat diarrhoea and vomiting. The seed oilisedible, and the seeds are also used to makesoap

Production and international trade Thetimber of Chrysophy!Ium of neonum is tradedinternationally, but data on volumes are notavailable due to mixing with other Sopotoceoespecies. In C6te d'Ivoire it is rated as one of thehigh-value timbers for export, but it is un-doubtedIy exported from other countries aswell.

Properties The heartwood is whitish, oftenwith a pinkish tinge when freshly cut, butdarkening to yellowish or greyish brown, oftenwith irregular dark stripes, and indistinctlydemarcated from the slightly paler sapwoodThe grain is straight, occasionally wavy or in-terlocked, texture fine to moderately fine.The wood is medium-weight, with a density of560-810 kg/in3 at 12% moisture content. Itdries slowly. The rates of shrinkage are fairlyhigh: 5.8-6.2% radial and 8.1-8.9% tangentialfrom green to oven dry. However, with somecare, the wood air dries well with little de-grade.Variation between results of different analysesof the mechanical properties of the wood arelarge: at 12% moisture content, the modulus ofrupture is 78-162 N/min2, modulus of elasticity12,550-15,885 N/mm2, compression parallel tograin 39-74 N/min2, cleavage 11-21 N/mm andChalais-Meudon hardness 2.4-5.0. The wood isbrittle and not very shock-resistantThe wood is easy to saw, works well with handand machine tools, and it can be planed to anexcellent finish. It does not easily spitt whennailed, but pre-boring is recommended, and itholds nails and screws well. It has good gluingproperties, and peels and slices satisfactorilyThe wood is only moderately durable and liableto fungal discoloration and attack by termitesand marine borers. It can be treated with pre-servatives using pressure processes

The yield of oil from dried seeds is about 25%

CHRYSOPHYl, LUM 157

CHRYSOPHYLLUMAFRiCANUMA. DC.

Protologue Prodr. a 163 (1844).Family SapotaceaeChromosome number2n= 26

Synonyms Gombey0 @1ncuno (ADC. ) Pierre(1891), Chrysophy!!urn edu!e Hoyle (1932),Chrysophy!Ium deleuoyi auct. non De Wild

Vernacular names African star apple (En)Longhi, longhirouge, longuirouge (Fr). Cola doob6 (Po)

Origin and geographic distribution Chryso-phyllum of neonum is widespread from SierraLeone east to Uganda and south to DR Congoand Cabinda (Angola)

Uses The wood (trade name: longhi, longui,akatio) is used for carving, moulding, turnery,cabinet making, veneer and plywood. It is also

I

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158 TIMBERSl

The oilis deep red and it has low saponificationand iodine values

Botany Medium-sized tree up to 25(-35) intall; bole up to 60(-80) cm in diameter,straight, often fluted or angular, reaching up to12 in to the first branches, with steep but-tresses at base; bark up to 2.5 cm thick, surfacegreyish brown, fissured, inner bark fibrous,brown, exuding a white sticky latex; crownspreading; young branches greyish brown toreddish brown appressed hairy. Leaves ar-ranged spiralIy in tufts at the ends orbranches,simple and entire; stipules absent; petiole 1.5-3.5 cm long, appressed hairy; blade elliptical toobovate-oblong, 15-35 cm x 5-13 cm, curieateat base, shortly acuminate at apex, greyishbrown to reddish brown appressed hairy below,pinnately veined with 15-30 pairs of straightlateral veins. Flowers in axillary fascicles, bi-sexual, regular, 5-merous; pedicel up to 5 minlong; sepals free, broadly ovate, c. 4 min long,pubescent outside; corolla with c. 3 min longtube and rounded lobes c. 2 mm long, hairy atmargins, creamy white; stamens inserted incorolla tube, opposite corolla lobes; ovary supe-nor, conical, long-hairy, 5-celled, style short,tapering. Fruit a large, ovoid to globose berryup to 7 cm long, becoming yellow-orange andglabrous when ripe, slightly pointed at apex,up to 5-seeded. Seeds ellipsoid, flattened, up to3.5 cm x 2 cm, shiny brown. Seedling withepigealgerminationIn Ghana Chrysophy!Ium o17icunum flowers inApril-May and fruits are mature in December-March.

Chrysophyllum comprises about 70 species andoccurs throughout the tropics. Tropical Amer-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Aneuchrysophyllumand sect. Done!IQ) contain African species.Chrysophy!Ium o17iconum belongs to sect.Anewchrysophyllum.Chrysophyllumgorungosonum Engl. (synonym:Chrysophyllum Iuluum S. Moore) is related toChrysophyllum of neonum, but differs in itssmaller leaves and fruits. The wood of Chryso-phyl!urn gorungosonum is used for construc-tion in Uganda, Tanzania and Malawi, but theoften strongly fluted hole is a drawback. It isalso used for firewood and the production ofcharcoal. The fruits are edible. In Rwanda anextract of the leaves and roots is drunk and isapplied externalIy as an anodyne

Ecology Chrysophy!Ium o17iconum occurs inlowland rainforest, often near rivers, up to1400 in altitude

Management In Gabon Chrysophyl!urnof neonum occurs reportedIy in low densities(on average 0.11 in3ftia). This is probably alsothe case elsewhere, and this low density, to-gether with its preference for river banks,makes it difficult to specify managementmeasures to promote this species. The 1000-seed weight is 1.2-1.4 kg.The wood has to be treated immediately afterfelling and cutting. Fresh logs do not float inwater and have to be transported by truck

Genetic resources and breeding Chryso-phyllum of neonum is widespread and does notseem to be in danger of genetic erosion. How-ever, in some countries, e. g. Ghana, it is un-common and there protection measures may beneeded

Prospects The role of Chrysophyllum o17i-corium in timber trade is not clear. The timber

is harvested and traded together with that ofother Sopotoceoe species that have timber ofcoinparable colour and density, butthe share ofeach species should be investigated. It seemslikely that the amount of Chrysophyllum ofri-curium timber in international trade is limited

because of the often comparatively smallsize ofthe bole

Major references Aubr6ville, 1959d; Bolza& Keating, 1972; Burki11, 2000; de Saint-Aubin, 1963; Takahashi, 1978.

Other references Ajiwe at a1. , 1997; Au-br6ville, 1961; Chudnoff, 1980; Hemsley, 1968;Katende, Birnie & Tengnas, 1995; Kupicha,1983; Liben, 1989; Neuwinger, 2000; Raponda-Walker & Sillans, 1961; Wilks & Issemb6,2000

Authors R. H. M. J. Lemmens

CHRYSOPHYLLUMBornNIANUM (Pierre)Baehni

Protologue Boissiera 11: 76 (1965).Family SapotaceaeSynonyms Gumbeyo botuiniono Pierre (1891),

Gombeyu ingdogoscoriensis Lecomte (1920).Vernacular names Famelona a grandes

foullles (Fr)Origin and geographic distribution Chryso-

phyllum botuiniont, in occurs in the Comorosand eastern Madagascar

Uses In Madagascar the wood is commonlyused for interior joinery and carpentry, furni-

:

ture, moulding, panelling, light flooring andladders. It is also used in shipbuilding becauseof its elasticity.The fruits are edible. Crushed leaves are ap-plied as a dressing to treat scorpion stings. InMadagascar Chrysophyllum botuinionum formspart of plant mixtures used to treat poisoningand to relieve symptoms of malaria, tirednessand muscular pains.

Properties The heartwood is cream-colouredto brownish yellow or pinkish brown, some-times with irregular dark stripes, and iridis-tinctly demarcated from the sapwood, which is5-6 cm wide. The grain is straight, occasionaUyslightly wavy, texture fine to moderately fineand even.

The wood is medium-weight, with a density of630-710 kg/ing at 12% moisture content. Therates of shrinkage are moderate to high, fromgreen to oven dry 3.7-4.7% radial and 8.6-9.9(-12.5%) tangential. However, the wood airdries well with little degrade, although occa-SIonally with a tendency to warpAt 12% moisture content, the modulus of rup-ture is 128-151 N/mm2, modulus of elasticity11,470-12,940 N/mm2, compresslon paralleltograin 43-53 N/mm2, shear 8.1 N/mm2, cleavage17-23 N/mm and Chalais-Meudon hardness2.5-3.1

The wood is easy to saw, works wellwith handand machine tools, and it can be planed to anexcellent finish. It does not easily split whennailed, but it holds nails and screws only inod-erately well. It has good gluing and paintingproperties. It turns well, and it has good steambending properties. The wood is only moder-ately durable; it is liable to attacks by fungiand insects. The heartwood is resistant totreatment with preservatives, the sapwoodmoderately resistant

Botany Medium-sized tree up to 25 in tall;hole up to 60 cm in diameter, straight, oftenfluted, reaching up to 18 in to the firstbranches, often slightly buttressed at base;bark surface smooth, inner bark exuding asticky white latex; young branches angular,hairy, older branches with leaf scars. Leavesarranged spiralIy, simple and entire; stipulesabsent; petiole 1.5-2 cm long, channelled,hairy; blade elliptical to obovate, 7-12(-40) cmx 2.5-3.5(-12) cm, curieate at base, shortlyacuminate at apex, densely reddish brown ap-pressed hairy below, pinnately veined with 12-30 pairs of straight lateral veins. Flowers inaxillary fascicles, bisexual, regular, 5-merous,sessile or with short pedicel; sepals free, 2.5-3

min long, pubescent; corolla with c. 2.5 mmlong tube and rounded lobes c. 1.5 min long,hairy at margins, creamy white; stamens In-serted in corolla tube, opposite corolla lobes;ovary superior, long-hairy, 5-celled, style short,tapering, glabrous. Fruit a globose berry up to4.5 cm in diameter, up to 5-seeded. Seeds ellip-sold, flattened, up to 3 cm x 1.5 cm.Chrysophyl!urn comprises about 70 species andoccurs throughout the tropics. Tropical Amer-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Aneuchrysophyl!urnand sect. Done!!0) contain African speciesChrysophyl!urn botuinionum belongs to sect.Aneuchrysophy!IumThe fruits of Chrysophyllum botu!itionum arecommonly eaten by lemur species, which maydisperse the seeds.

Ecology Chrysophyl!urn botuinionum occursin humid evergreen forest from sea-level up to1750 in altitude. It is particularly characteris-tic of inId-elevation forest along the escarpmentin eastern Madagascar. It occurs in littoral andsub-littoral forests on sandy soils, but it preferslatentic soils where it may be abundant. It isalso found in forest remnants on high plateaus

Management After felling, logs should beremoved from the forest rapidly as they maydevelop heart shakes and end splits. They areliable to blue stain attack and therefore dip-

ping in anti-sapstain preservatives Ls recoin-mended before stacking

Genetic resources and breeding Chryso-phyllum botuinionum is widespread in easternMadagascar and locally common, and it doesnot seem to be liable to genetic erosion at pre-sent. However, there is some concern about thedecrease of the species ln recent years, and itwas included in a TRAFFIC (the wildlife trademonitoring network, a joint programme ofWWF and IUCN) list of species that are usedin traditional medicine and are in need of con-servation, management and/or research ineastern and southern Africa

Prospects As long as stands of sufficientproportions remain, the timber of Chrysophyl-Ium botuinionum will be important for localutilization. It is recommended that the propa-

gation and management of this species be stud-led as a basis for possibilities for Its use in tim-her plantations of indigenous trees in Mada-gascar

Major references Anbr6viUe, 1974; Bolza &

CHRYSOPHYLLUM 159

160 TIMBERSl

Keating, 1972; Gu6neau, Bedel & Thie1, 1970-1975; Takahashi, 1978.

Other references Capuron, 1966b; Randri-ariarivelojosia at a1. , 2003; Schatz, 2001; Stygerat a1. , 1999

Authors R. H. M. J. Lemmens

CHRYSOPHYLLUMGIGANTEUMA. Chev.

Protologue Bull. SOC. Bot. France 61, Mein.a 267 (1917).

Family SapotaceaeChromosome number2n=26

Synonyms Gumbeyo gigonteo (Achev. )Aubr6v. & Fellegr. (1960), Gombeyobotrys gi-gunteo (Achev. ) Aubr6v. (1972)

Vernacular names Koanandio, abam g6ant(Fr).

Origin and geographic distribution Chryso-phyllum gigonteum occurs from Sierra Leoneto Gamero0n, Gabon and Congo.

Uses The wood of Chrysophyllum gigonteumis similar to that of Pouterio spp. and is used inthe same way, e. g. for interior construction,joTnery and furniture. The fruits are edible;they are sold on village markets in Ghana.

Production and international trade Thewood of Chrysophyllum gigonteum enters tradetogether with that of Pouterio spp. as 'aningr6','ani6gr6' or 'aningu6ri'. In 1972, 1973 and 1983,C6te d'TVoire exported about 135,000 ms of logsof anlegr6' per year, mainly to Italy, Spain andGreece.

Properties The heartwood is yellowish white,darkening to creamy pink, and indistinctlydemarcated from the sapwood. The grain isstraight, occasionally interlocked, texture fineto moderately fineThe wood is medium-weight, with a density of540-570 kg/ina at 12% moisture content. Therates of shrinkage are moderate, from green tooven dry 3.2-4.9% radial and 6.8-8.0% tangen-tial. With some care, the wood air dries wellwith little degrade. Once dry, it is stable in

At 12% moisture content, the modulus of rup-ture is 1/4-123 N/min2, modulus of elasticity9410-10,100 N/min2, compression parallel tograin 44-47 N/min2, shear 5.5-7.0 N/mm2cleavage 13-14 N/min and Chalais-Meudonhardness 2.3-2.6.

The wood is easy to saw, works well with handand machine tools, and it can be planed to anexcellent finish. Unlike Pouterio wood, it doesnot blunt sawteeth because it hardly contains

silica. It does not easily split when nailed, andit holds nails and screws well. It has good glu-ing and painting properties, and peels andslices satisfactorily. The wood is not durableand liable to attack by fungi and termites.However, It is not very susceptible to Lyetusattack. It can be treated with preservativesusing pressure processes

Botany Medium-sized to fairly large tree upto 35 in tall; hole up to 90 cm in diameter,straight, sometimes fluted, reaching up to 20 into the first branches, with steep buttresses atbase; bark thick, surface greyish white, longi-tudinally fissured, exuding a sticky latex whencut; crown dense; young branches greyishbrown to reddish brown appressed hairyLeaves arranged spiralIy in tufts at the ends ofbranches, simple and entire; stipules absent;petiole c. I cm long; blade obovate-oblong, 8-20cm x 3-7 cm, curieate at base, shortly acumi-nate at apex, leathery, silvery to reddish brownappressed hairy below, with small translucentdots above, pinnately veined with 7-10 pairs oflateral veins. Inflorescence an axillary falseraceme c. 3 cm long, densely hairy. Flowersbisexual, regular, 5-merous; pedice1 3-4 mmlong; sepals free, c. 1.5 mm long, pubescentoutside; corolla c. 2 min long, with short lobes,glabrous, creamy white; stamens inserted incorolla tube, opposite corolla lobes; ovary supe-nor, conical, long-hairy, 5-celled, style shortFruit a globose berry up to 5.5 cm in diameter,becoming yellow-orange and glabrous whenripe, up to 5-seeded. Seeds ellipsoid, flattened,up to 3 cm x 2 cm x I cm, brown, with largescar. Seedling with epigealgerminationFruits are produced in the drier months of theyear; the seeds are dispersed by monkeys andother animals that relish the fruits

Chrysophyllum comprises about 70 species andoccurs throughout the tropics. Tropical Amer-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Aneuchrysophyllumand sect. Donello) contain African species.Chrysophyllum gigonteum belongs to sect.Aneuchrysophyllum. It resembles Chrysophyl-Ium o16idum G. Don, which has similar woodbutts more important as a fruittree and whichdiffers in its fascicled flowers

Ecology In 06te d'TVoire and Ghana Chryso-phyllum gigonteum occurs in semi-deciduousforest, where it is common. In Gamero0n andGabon it seems to be much less common.

service

Management The 1000-seed weightis aboutIkg

Genetic resources and breeding In WestAfrica Chrysophyl!urn gigonteum is common insemi-deciduous forest, and although it is coin-monly exploited for its timber, it does not yetseem to be under threat of genetic erosion

Prospects The exploitation of Chrysophyl-Ium gtgonteum seems to take place on a largerscale; it is probably logged indiscriminatelytogether with Pouterio spp. Research is neededon growth and natural propagation to developmethods of sustainable management of forestswhere it occurs.

Major references Aubr6viUe, 1959d; Au-br6ville, 1972; Normand, 1970; Takahashi,1978.

Other references ATIBT, 1986; Aubr6ville,1964; Burki11, 2000; de to Mensbruge, 1966;Durand, 1983a; Hawthorne, 1990; Hawthorne,1995; Saville & Fox, 1967; Siepel, Poorter &Hawthorne, 2004

Authors R. H. M. J. Lemmens

CongoUses The wood (trade names: abam, longhi)

is suitable for construction, light flooring, inte-nor trim, mine props, railway sleepers, ships,vehicle bodies, furniture, cabinet work, imple-merits, sporting goods, carvings, turnery, ve-neer, plywood, hardboard, particle board andpulp for paper production.The sweet acidulous pulp of very ripe fruits iscommonly eaten fresh. In Gabon the Bakotaand Bakw616 people eat the salted pulp withstaple foods such as cassava and plantainDried pulp Is occasionally used in sauces. InCongo a bark decoction is applied vaginalIy totreat inflammations of uterus and vagina, andit is used in vapour baths to treat rheumatismand kidney pain. Bark powder is applied towounds. In DR Congo a maceration of the rootbark is applied topicalIy to treat earache.

Production and international trade Thetimber ofChrysophyl!urn IOCourtionum is tradedinternationally in small volumes, but data arenot available due to mixing with other Sopoto-ceoe species. The fruits are commonly found onlocal markets in Central Africa

Properties The heartwood is yellowish whiteto yellow-brown, sometimes with ribbon-like orinottled figure, and indistinctly demarcatedfrom the sapwood, which is up to 5 cm wideThe grain is usually straight, texture fine tomoderately fineThe wood is medium-weight, with a density of685-730 kg/ing at 12% moisture content. Withsome care, the wood air dries well with littledegrade, but it should be quartersawn anddried slowly.At 12% moisture content, the modulus of rup-ture is 133-150 N/min2, modulus of elasticity15,790 N/min2, compression parallel to grain55-70 N/mm2 cleavage 15.7-20.6 N/mm andChalais-Meudon hardness 3.6-4.2. The wood isbrittle and not very shock-resistantThe wood is fairly easy to saw, works wellwithhand and machine tools, and it can be planedto a smooth finish. It does not easily split onnailing, and it holds nails and screws well. Thewood is only moderately durable and liable toattacks by blue stain, termites and marineborers. The sapwood is fairly permeable to pre-servatives, the heartwood resistantThe fresh fruit contains per 100 g: water 71 g,energy 376 kJ (90 kcal), protein 1.4 g, fat 6.6 g,carbohydrate 6.2 g, Ca 17 ing, P 14 ing.

Description Large tree up to 40 in tall; holebranchless for up to 24 in, straight and cylin-drical, sometimes fluted, up to 100 cm in dia-

CHRYSOPHYl, LUMLACOURTIANUM De Wild.

Protologue Miss. tin. Laurent L 425 (1907)Family SapotaceaeChromosome number2n=28

Synonyms Chrysophyllum outranio, turnAchev. (1917), Gombeyo IOCourtiono (DeWild. ) Aubr6v. & Fellegr. (1960).

Vernacular names Aham, longhi, longhirouge (Fr)

Origin and geographic distribution Chryso-phyllum IOCourtionum occurs from Cameroonto the Central African Republic, Gabon and DR

CHRYSOPHYLLUM 161

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Other botanical information Chrysophyl-Ium comprises about 70 species and occursthroughout the tropics. Tropical America isrichest in species (about 45), followed by conti-nental Africa (about 15), Madagascar (about10) and tropical Asia and Australia (together2). The genus has been subdivided into 6 sec-tions, 2 of which (sect. Aneuchrysophyllum andsect. Donel!a) contain African species. Chryso-phyllum IOCourttonum belongs to sect. Aneu-chrysophyllum.

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; (25: intervessel pits small (4-7F1m)); 26: intervessel pits medium (7-10 prn);30: vessel-ray pits with distinct borders; SImi-Iar to intervessel pits in size and shapethroughout the ray cell; (31: vessel-ray pitswith much reduced borders to apparently sim-PIe: pits rounded or angular); 32: vessel-raypits with much reduced borders to apparentlysimple: pits horizontal(scalariform, gash-like)to vertical(palisade); (33: vessel-ray pits of twodistinct sizes or types in the same ray cell); 42:mean tangential diameter of vessellumina100-200 pin; 46: , 5 vessels per square milli-metre; 56: tyloses common. Tracheids and fi-bres: 61: fibres with simple to minutely bor-dered pits; 66: non-septate fibres present; 69fibres thin- to thick-walled. Axial parenchyma:86: axial parenchyma in narrow bands or linesup to three cells wide; (87: axial parenchymareticulate); (88: axial parenchyma scalariform);93: eight (5-8) cells per parenchyma strandRays: (97: ray width I-3 cells); 98: larger rayscommonly 4- to 10'seriate; 107: body ray cellsprocumbent with mostly 2-4 rows of uprightand/or square marginal cells; 108: body raycells procumbent with over 4 rows of uprightand/or square marginal cells; 1/5: 4-12 raysper mm. Mineral inclusions: 136: prismaticcrystals present; 138: prismatic crystals in pro-cumbent ray cells; 142: prismatic crystals inchambered analparenchyma cells(N. P. Monel, P. DCtienne & P. E. Gasson)

Growth and development Growth ofChry-sophyllum IOCourtionum seedlings is slow, 6-10 cm/year during the first three years. In Ga-bon Chrysophy!Ium IOCourtionum flowers inJuly-August, during the dry season. In thisperiod the trees are often leaness for a shortperiod. They fruit regularly, often every year

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Chrysophyllum IOCot, rtionum - I, base of 601e, '2, leafy bronch, 3, fruit, ' 4, fruit in cross section.Redrown ond oddpted by Achingd Satin Nur-human

meter, with steep buttresses at base; bark upto 2.5 cm thick, surface greyish brown, fis-sured, inner bark fibrous, brown, exuding asticky latex; crown spreading; young branchesgreyish hairy. Leaves arranged spiralIy in tuftsat the ends of branches, simple and entire;stipules absent; petiole 2-3 cm long, groovedabove; blade erriptical to obovate-oblong, 11-36cm x 4.5-12.5 cm, curieate at base, shortlyacuminate at apex, glabrous, pinnately veinedwith 10-17 pairs of lateral veins. Flowers inaxillary fascicles, bisexual, regular, 5-merous;pedicelc. 3 mm long; sepals free, broadly ovate,c. 4 min long, pubescent outside; corolla with c.2.5 mm long tube and rounded lobes c. I minlong, hairy at margins, creamy white; stamensinserted near base of corolla tube, oppositecorolla lobes; ovary superior, conical, long-hairy, 5-celled, style short. Fruit a large, ovoidto globose berry 6-TICm long, becoming orangeto red and glabrous when ripe, up to 5-seededSeeds ellipsoid, flattened, up to 3.5 cm x 2 cm,shiny brown. Seedling with epigeal germina-

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but sometimes once every two years, and fruitstake about 12 months to ripen. In Gabon ripefruits are found in May-September. The seedsare dispersed by primates (including gorillasand chimpanzees) and elephants. Germinatingseedlings have been observed in elephant dung.Seedlings around mother trees are rare. Thishas been attributed to intensive predation ofthe fruits by rodents and wild pigs, which de-stroy the seeds

Ecology Chrysophy!!urn IOCourtionum oc-curs particularly in dense semi-deciduous for-est, where it often occurs scattered and is gen-erally uncommon. In DR Congo it is locallycommon together with Geltis spp. on sandy-loamy soils in gallery forest in valleys, wherethe average annual rainfallis 1400-1500 mmand the mean annual temperature 24'C.

Propagation and planting Seeds startgerminating after 50 days and germinationmay take over 100 days. Seedlings need shade

Management Chrysophy!!urn IOCourtionumis rarely planted, but trees are commonlymaintained when clearing the forest for agri-culture

Harvesting The fruits are collected from theground

Handling after harvest Logs should not beleft in the forest for too long after felling be-cause they are liable to blue stain; they shouldbe converted as soon as possible

Genetic resources Although ChrysophyllumIOCourtionum is relatively uncommon In manyregions within its distribution area, It Is notimmediately liable to genetic erosion because itis quite widespread in regions where forest isstillcommon

Prospects In spite of the fact that Chryso-phyllum IOCourtionum is an Interesting timberand fruit tree, very little research has beendone on its propagation and possibilities ofdomestication. However, its apparently slowgrowth is a serious drawback

Major references Bolza & Keating, 1972;Bourobou-BOUTobou, 1994; Latham, 2004; Ta-kahashi, 1978; Wilks & Issemb6, 2000

Other references Aubr6ville, 1961; Au-br6ville, 1964; CTFT, 1961h; Glouchkoff, un-dated; InsideWood, undated; Lubini & Kusehu-Iuka, 1991; Neuwinger, 2000; 0bama Ondo,2002; Raponda-Walker & Sillans, 1961

Sources of illustration de Wildeman,1905-1907; Wilks & Issemb6, 2000

Authors R. H. M. J. Lemmens

CHRYSOPHYLLUMPERPULCHRUMMildbr. exHutch. & Dalziel

Protologue F1. W. trop. atr. 2: 10 (1931)Family SapotaceaeChromosome number2n=28

Synonyms Gumbeyo perpulchr0 0011dbr. exHutch. & Dalziel) Aubr6v. & Fellegr. (1960)

Vernacular names Red asanfena, monkeystar apple, bird-lime tree (En). hamgu6rirouge(Fr).

Origin and geographic distribution Chryso-phy!!urn peruulchrum Is widely distributedfrom Guinea east to Uganda and Tanzania

Uses The wood (trade name: inululu)is suit-able for construction, flooring, furniture, cabi-net work, joinery, toys and novelties, musicalinstruments, turnery, boxes and crates, vats,carvings, veneer and plywood. In Tanzania thewood is also used for grain mortars and bee-hivesIn C6te d'TVoire a bark decoction is taken as atonic, anodyne, galactagogue and aphrodisiac,and to treat jaundice, asthma and other respi-ratory complaints, and madness. The fruit issometimes eaten. The latex is used as a rubbersubstitute and as birdlime

Production and international trade Thetimber of Chrysophyllum peruulchrum is tradedinternationally in small amounts, but data arenot available due to mixing with other Sopoto-cede species.

Properties The heartwood is yenowish orpinkish white to brownish yellow and iridis-tinctly demarcated from the sapwood. Thegrain is usually straight, texture fine and even.The wood is lustrous. In Sierra Leone the woodis considered too gummy to be useful.The wood is fairly heavy, with a density of660-820 kg/ina at 12% moisture content. Therates of shrinkage from green to oven dry arehigh: 5.9-6.3% radial and 9.7-10.7% tangen-tial; shrinkage from green to 12% moisturecontent is 2.0-3.0% radial and 7.2-8.5% tan-

gential. In spite of the high shrinkage, dryingis rapid, with little splitting or checking. Afterdrying the wood is not stableAt 12% moisture content, the modulus of rup-ture is 143-189 N/mm2, modulus of elasticity14,120 N/mm2, compression parallel to grain69-72 N/mm2, cleavage 15.5 N/min andChalais-Meudon hardness 5.9. The wood is

moderately shock-resistantThe wood is fairly easy to saw, works wellwithhand and machine tools, and can be planed to asmooth finish. Pre-boring is necessary before

CHRYSOPHYLLUM 163

.

164 TIMBERSl

nailing, and the wood holds nails and screwswell. The gluing and steam bending propertiesare good, as are the slicing propertiesThe wood is not durable and is liable to attacks

by Insects. The sapwood is susceptible toLyetus borers. The sapwood is fairly permeableto preservatives, the heartwood moderatelyresistant.

The kernel of the seed contains 4% oil, witholeic acid 39%, linoleic acid 31%, palmitic acid20%, stearic acid 9%, linolenic acid I% andtraces of lauric, myristic and arachidic acidThe alkaloid cardiochrysine has been recordedfor the bark; this compound showed depressantactivity in the thalamus and hypothalamus,and hypotensive and cardiotonic activity

Botany Large tree up to 40 in tall; hole up to100 cm in diameter, straight and cylindrical,slightly fluted to buttressed at base; bark sur-face greyish brown, smooth with longitudinalfissures, Inner bark fibrous, pale brown, exud-ing a white sticky latex; young branchesdensely reddish brown hairy. Leaves arrangedspiralIy, simple and entire; stipules absent;petiole I-3.5 cm long, grooved above; bladeelliptical to obovate-oblong, 8-25(-40) cm x 3-9(-14) cm, slightly curieate to rounded at base,acute at apex, leathery, densely reddish brownvelvety hairy beneath, pinnately veined with(11-)14-21(-25) pairs of lateral veins. Flowersin axillary fascicles, bisexual or occasionallyfunctionally unisexual, regular, 5-meTous;pedicel up to 1.5 min long; sepals free, broadlyovate, c. 4 mm long, pubescent outside; corollawith c. 2.5 mm long tube and rounded lobes c.1.5 mm long, hairy at margins, creamy white;stamens inserted near base of corolla tube,opposite corolla lobes; ovary superior, conical toglobose, long-hairy, 5-celled, style up to 1.5 minlong. Fruit a globose berry 2-4 cm long,densely reddish brown hairy, up to 5-seededSeeds ellipsoid, flattened, up to 2 cm x 1.5 cm,shiny brown. Seedling with epigeal germina-tion; hypocoty1 6-7 cm long, epicotyl c. 2 minlong; cotyledons broadly ovate, 2-5 cm long,leafyChrysophyllum comprises about 70 species andoccurs throughout the tropics. Tropical Airier-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Anet, chrysophyllumand sect. Donello) contain African speciesChrysophyl!urn perpulchrum belongs to sect.Aneuchrysophyllum

Chrysophyllum perpulchrum trees producefruits in abundance when the bole is 60 cm in

diameter, but individual trees with hole diame-ters of 20 cm may already start fruiting. InSierra Leone flowering is at the beginning ofthe dry season, and mature fruits can be foundin February-March. In Ghana flowering is inMarch-April and fruits are ripe in September-December.

Ecology In West Africa Chrysophyllum per-pulchrum occurs in semi-deciduous forest, mostabundantly in dry forest types, particularly onslopes. Locally it can be dominant. In East A1-Tica it is found in rainforest at 800-1200 in

altitude. Seedlings need shade for proper de-velopment. Regeneration is not abundant indry forest which is regularly burned, and seed-lings are more common in less disturbed forest.

Management Recorded 1000-seed weightsare 0.6-1.2 kg. Seeds germinate in 12-30 daysStripped plants transplant well. HarvestingChrysophyllum perpulchrum trees with a bolediameter of less than 60 cm distinctly reducesregeneration because many trees only producefruits abundantly above that diameter. Treesin a 12-year-old experimental plot in Ghanawere 3-6 in tall

Genetic resources and breeding Chryso-phy!Ium perpulchrum is not liable to geneticerosion because it is widespread and in manyregions common.

Prospects Very little is known about Chry-sophy!JumperPMlchrum and research is neededto evaluate its usefulness as a timber tree.

However, its wood properties are not out-standing in comparison with other Sopotoceuespecies.

Major references Bolza & Keating, 1972;Burki11, 2000; Hawthorne, 1995; Takahashi,1978; Voorhoeve, 1979

Other references Aubr6ville, 1959d; Au-br6ville, 1964; Bouquet & Debray, 1974; Bryce,1967; CTFT, 1961g; Neuwinger, 2000; 0teng-Amoako (Editor), 2006; Plumptre, 1995; Saville& Fox, 1967; Taylor, 1960.

Authors R. H. M. J. Lemmens

CHRYSOPHYLLUMPRUNIFORME PierreexEngl

Protologue Monogr. afrik. Pflanzen-Fam. a42 (1904).

Family SapotaceaeChromosome number2n = 26.28Synonyms Donello pruniformis (Pierre ex

, I

Engl. ) Aub"6v. & Penegr. (1961).Origin and geographic distribution Chryso-

phyllum pruniforme is widely distributed fromSierra Leone east to western Uganda andwestern Tanzania

Uses In Tanzania the wood is used for con-

struction, grain mortars and beehives. In WestAfrica it is used for house construction. In

Congo a tea made from the bark is drunk totreat cough. In some regions the frult pulp isreported as edible, but in others as medible

Properties The wood is yellowish white andmoderately hard

Botany Medium-sized tree up to 30(-40) intall; hole up to 80(-100) cm in diameter,straight and cylindrical, fluted or srightly but-tressed at base; bark surface dark brown toblackish, fissured, inner bark pale brown, fi.brous, exuding a little latex; young branchesreddish brown hairy. Leaves distichously al-ternate, simple and entire; stipules absent;petiole 0.5-I cm long, slender; blade ellipticalto ovate-oblong, 4-Ile13) cm x 2-5(-6.5) cm,asymmetricalIy curieate at base, long-acuminate at apex, leathery, glabrous, pin-nately veined with numerous, closely spacedlateral veins. Flowers in axillary fascicles,regular, 5-merous; pedice1 2-3.5 mm long; se-pals free, orbicular, up to 2 mm long, slightlypubescent to glabrous outside; corolla with upto I min long tube and rounded lobes up to 1.5min long, ciliate at margins, greenish; stamensinserted near base of corolla tube, oppositecorolla lobes; ovary superior, conical to globose,long-hairy, 5-celled, style short. Fruit a globoseto ovoid berry up to 5 cm x 4 cm, yellow whenripe, glabrous, up to 5-seeded. Seeds ellipsoid,flattened, up to 2.5 cm x 1.5 cm, shiny brownSeedling with epigealgerminationChrysophyllum comprises about 70 species andoccurs throughout the tropics. Tropical Amer-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Aneuchrysophyllumand sect. Done!IQ) contain African speciesChrysophyllum pruniforme belongs to sectDonello, characterized by the presence of nu-merous, closely spaced lateral veins in theleaves. It is closely related to ChrysophyllumDirtdijolium J. M. Wood & Franks from Eastand southern Africa, and the two species haveoften been confused

Ecology Chrysophyl!urn pruniforme occursin lowland rainforest, both primary as well as

secondary forest, in East Africa up to 1500 inaltitude. It is usually found on well-drainedsoils. It is fairly common in West and CentralAfrica. Seedlings of Chrysophyllum pruniformeare shade tolerant.

Management The 1000-seed weightis about715 g. In Sierra Leone Chrysophyllum pruni-forme sometimes constitutes more than 9% ofall trees with a girth of over 30 cm, and up to90 stemsftia have been counted

Genetic resources and breeding Chryso-phyllum pruniforme does not seem to be liableto genetic erosion because it is widespread andin several regionsfairly common

Prospects Very little is known about Chry-sophy!Ium pruniforme. In the past it has beenrecorded as a timber tree of little interest andeven as a weed species in commercial forest,but research is warranted to evaluate its role

in sustainably managed natural productionforest in relation to its wood properties.

Major references Burki11, 2000; Haw-thorne, 1995; Hemsley, 1968.

Other references Aubr6ville, 1959d; Au-br6ville, 1961; Aubr6ville, 1964; de in Mens-bruge, 1966; Lovett at a1. , 2006; Neuwinger,2000; Saville & Fox, 1967; Voorhoeve, 1979.

Authors R. H. M. J. Lemmens

CHRYSOPHYLLUM 165

CHRYSOPHYLLUMSUBNUDUMBaker

Protologue Onv. , F1. trop. Mr. 3: 499 (1877)Family SapotaceaeChromosome number2n=26

Synonyms Gombeyo subnudo (Baker) Pierre(1891).

Vernacular names Adasema (En). minigu6riargent6, longhi, longhirouge (Fr)

Origin and geographic distribution Chryso-phyllum subnudum is widely distributed fromSierra Leone east to DR Congo.

Uses The wood (trade name: longhi) is suit-able for construction, flooring, interior trim,joinery, furniture, mine props, vehicle bodies,Implements, panelling, veneer and plywood. InGabon and Congo a bark decoction is used aspurgative and for treatment of intestinal coin-plaints. The sweetly acidulous fruit pulp is edible.

Production and international trade The

timber of Chrysophyl!urn subnudum is tradedinternationally in small amounts, but quanti-tative data are not available due to mixture

with other Sopotoceoe species.Properties The heartwood is yellow-grey to

pinkish brown and indistinctly demarcated

.

166 TIMBERSl

from the whitish sapwood, which is up to 10 cmwide. The grain is usually straight but some-times wavy, texture fine. The wood is lustrousThe wood is fairly heavy, with a density of720-800 kg/ina at 12% moisture content. It airdries well without difficulties. The rates of

shrinkage are moderate to high, from green tooven dry 4.1-6.9% radial and 8.0-10.5% tan-gentialAt 12% moisture content, the modulus of rup-ture is 172-202 N/mm2, modulus of elasticity11,670-16,180 N/mm2, compression parallel tograin 62-84 N/min2, shear 7.3-8.0 N/mm2cleavage 18-23 N/mm and Chalais-Meudonhardness 4.6-6.6. The wood is moderatelyshock-resistant.

The wood is fairly easy to saw, works wellwithhand and machine tools, and it can be planedto a smooth finish. It holds nails and screws

well; pre-boring before nailing is recommendedbut not necessary. The wood takes polish andvarnish well. It is only moderately durable andliable to attacks by blue stain, termites andmarine borers. The sapwood is moderately resis-tant to preservatives, the heartwood resistant.

Botany Medium-sized tree up to 25(-35) intall; hole up to 60(-70) cm in diameter, straightand cylindrical, often shortly buttressed atbase; bark surface greyish brown, smooth tofissured, inner bark fibrous, brown, exuding asticky latex; young branches reddish brownhairy. Leaves arranged spiralIy, simple andentire; stipules absent; petiole I-2 cm long,grooved above, greyish hairy; blade narrowlyelliptical to oblong-lanceolate, 8-20 cm x 2.5-5cm, curieate at base, acuminate at apex, Ieath-ery, densely silvery grey velvety hairy beneathbut glabrescent, pinnately veined with 8-12pairs of lateral veins. Flowers in axillary fasci-cles, regular, 5-merous; pedice1 2-5 min long;sepals free, broadly ovate, c. 2 min long, red-dish brown pubescent outside; corolla with c2.5 mm long tube and rounded lobes c. 1.5 minlong, ciliate at margins, greenish white; sta-mens inserted near base of corolla tube, OPPo-site corolla lobes; ovary superlor, conical toglobose, long-hairy, 5-celled, style short. Fruita globose berry c. 3 cm long, slightly 5-10bed,greenish to yellowish, glabrous, up to 5-seededSeeds ellipsoid, flattened, up to 2.5 cm x 1.5cm, brown. Seedling with epigeal germination;hypocoty1 6-12 cm long, glabrous, epicoty1 0.5-3 cm long, pubescent; cotyledons ovate to ellip-tica1, 3-5 cm x 2-3.5 cm, shortly stalked, leafy,glabrous.Chrysophy!Ium comprises about 70 species and

occurs throughout the tropics. Tropical Amer-ICa is richest in species (about 45), followed bycontinental Africa (about 15), Madagascar(about 10) and tropical Asia and Australia (to-gether 2). The genus has been subdivided into6 sections, 2 of which (sect. Aneuchrysophyllumand sect. Donello) contain African speciesChrysophyllum subnudum belongs to sectAneuchrysophy!IumIn C6te d'Ivoire and Ghana Chrysophyl!urnsubnudum flowers in December-June and

fruits in November-February; in Gabon itflowers around August. The seeds are dis-persed by animals that feed on the fruits suchas elephants and possibly also chimpanzees

Ecology Chrysophy!!urn subnudum occursin lowland semi-deciduous forest to evergreenforest, usually as an understorey tree, and iscommonin many regions.

Management Chrysophyl!urn sub, Iudumoften regenerates abundantly, and seedlingsare shade tolerant. The 1000-seed weight isabout 770 g. Seeds start to germinate 2-8weeks after sowing. The germination rate var-ies from 25% to 90%. The logs have to betreated on felling and the wood immediatelyafter sawing. Fresh logs sink in water andcannot be transported by river.

Genetic resources and breeding Chryso-phyl!urn subnudum is not liable to genetic ero-slon because it is widespread and in many re-gions common.

Prospects Very little is known about Chry-sophyllum subnudum and research is neededto evaluate its usefulness as a timber tree on a

wider scale. However, because of its small sizein comparison with other Sopotoceoe species, itseems of secondary interest only.

Major references Bolza & Keating, 1972;Burki11, 2000; de Koning, 1983; Hawthorne,1995; Takahashi, 1978

Other references Aubr6ville, 1959d; Au-br6ville, 1961; Aubr6ville, 1964; de in Mens-bruge, 1966; Durand, 1983a; Neuwinger, 2000;Oteng-kirioako (Editor), 2006; Raponda-Walker& Sillans, 1961

Authors R. H. M. J. Lemmens

CHUKRASIATABULARISA. Juss

Protologue Bull. Sci. Nat. Geol. 23: 241(1830)

Family MenaceaeChromosome number2n=26

Synonyms Chuhrosio uelutino on. Roem. )

CDC. (1878).Vernacular names Chickrassy, Chittagong

wood, Burma almondwood, East Indian ina-hogany (En)

Origin and geographic distribution Chu-fryosio tobuloris originates from tropical Asia(from India and SriLanka eastwardsto Borneoand China). It has been planted in many coun-tries outside tropical Asia, in Africa in Nigeria,Garnero0n and northern South Africa, andelsewhere e. g. in Hawaii, Puerto Rico, CostaRica and Australia

Uses In tropical Asia, especially India,Myanmar, Bangladesh, Vietnam, Thailand andsouthern China, the wood is highly prized forhigh-grade cabinet work, decorative panelling,interior joinery such as doors, windows andlight flooring, and for carving, toys andturnery. It is also used for light to medium-heavy construction work, e. g. for posts, beams,scantlings and planks, and for railway sleep-ers, ship and boat building, furniture, musicalinstruments, packing cases, sporting goods,lorry bodies, mallet heads, anvil blocks, brushwares, drawing equipment, rifle butts, veneerand pulpA bark extract has powerful astringent proper-ties and is used as a febrifuge and to treat di-arrhoea. In India Chukrosio tubuloris is

planted as a shade tree in coffee plantations,and in Vietnam and Malaysia as an ornamen-taltree

Production and international trade Chu-

forOSio tabuloris timber is traded in small

amounts and often together with timbers fromother Meltoceoe genera such as Cedrelo andToo, IQ. It may also be marketed under mixedhardwood with a variety of other timbers. In

Thailand a production of 3200 ina was recordedin 1966, increasing to 9800 ms in 1989,whereas a production of less than 350 in31yearis reported from India. Production in Africa isinsignificant

Properties The heartwood is pale reddishbrown, yellowish red to red, darkening to darkyellowish brown, reddish brown to mediumdark brown on exposure, sharply differentiatedfrom the yellowish white, pale yellowish brown,pinkish brown or greyish brown sapwood; darkstreaks may be rather prominent. The grain isinterlocked and sometimes wavy, texture inod-erately fine but uneven. Freshly cut wood isfragrant, but dried wood has no characteristicodour or taste. Planed surfaces have a lustrous

satiny sheenThe wood is moderately heavy. The density is625-880 kg/ina at 15% moisture content. Therates of shrinkage are rather low: from green to15% moisture content c. 1.3% radial and 1.7%

tangential, from green to oven dry 3.9% radialand 6.0% tangential. Usually the wood driesfairly rapidly without degrade, but a slighttendency to check and warp and some liabilityto collapse have been reported. Fine hair sur-face checks may develop when drying thickboards

The wood is moderately hard. At 15% moisturecontent, the modulus of rupture is 82-101N/mm2 modulus of elasticity 10,800-14,300N/mm2, compression parallel to grain 47-64N/mm2, compression perpendicular to grain11-12 N/min2, shear 15-18 N/mm2, cleavage c.60 N/mm radial and 71 N/mm tangential andJanka side hardness 8990-9230 N

Tests in Malaysia showed that the wood is dif-ticult to saw, cross cut, turn and bore, but easyto plane. It produces a moderately smooth fin-ish, but some picking up of grain may occur onquarter-sawn material during planing andmoulding. However, tests in other areas

showed that the wood can be easily sawn andmachined. It has good nailing and screw-holding properties, it can be stained effectiveIyand polished excellently. The steam bendingproperties are rated as good. It can be readilypeeled and sliced into veneers and these can beglued satisfactorily to produce decorative ply-wood, fire-retardant treated plywood, and ply-wood suitable for concrete shuttering and ina-rine construction

The wood is considered non-durable to moder-

ately durable under exposed conditions. Theresistance to termite attack varies from good topoor. The wood is resistant to preservative

CHUKRAsiA 167

.

" . -

* * #,.

Chuhrosio tabuluris -pionted

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168 TIMBERSl

treatment.

Young leaves and bark have a high tannin con-tent, and the bark yields a reddish gum. A leafextract has been reported to exhibit consider-able antimalarial activity, as well as antibacte-rial and antifungal activities; the essential oilpresent in the leaf is responsible for these ac-tivities. The oil consists of oxygenatedmonoterpenes (42.8%, mainly carvacrol, thymoland borneol), phenyl propanoids (25.2%,mainly (E)-methylisoeugenol and myristicin)and smaller amounts of sesquiterpene hydro-carbons, monoterpene hydrocarbons and oxy-genated sesquiterpenes. Root extracts showedantifeedant activity against Spodoptero in-sects, with phragmalin 11monoids (tabulalinand tabulalides A-E) asthe active compounds.

Adulterations and substitutes The woodof Cedrelo odoroto L. and Toono ciliato

MRoem. , both planted in Africa, is similar tothat of Chuhrosio tobuloris, although the me-chanical properties of the wood of the formertwo species are inferior to those of Chuhrosiotubuloris wood

Description Deciduous, medium-sized tolarge tree up to 30(-40) in tall; hole branchlessfor up to 20(-25) in, with a diameter of up to

120 cm, with convex buttresses up to 1.5 inhigh or without buttresses; bark surface rustybrown or deep brown, deeply fissured orcracked, with lenticels, inner bark reddish;crown spreading. Leaves alternate, up to 50 cmlong, panpinnate with up to 24 leaflets in Iar-ger leaves; stipules absent; petiole 4-9 cm long;leaflets alternate, shortly stalked, ovate to ob-long, asymmetric, 4-17.5 cm x 2-6.5 cm, apicalones largest, acute to acuminate at apex, en-tire, glabrous to pubescent, pinnately veinedInflorescence an axillary panicle, often appear-ing terminal, up to 30 cm long. Flowers func-tionally unisexual, regular, 4-5-merous,sweetly scented; pedice1 2-4 min long; calyxshallowly cup-shaped, c. 3 mm in diameter,with short lobes; petals free, narrowly oblong tospatulate, I-1.5 cm long, contorted, cream-coloured to yellowish, often tinged pink; sta-mens 8-10, filaments united into a cylindricaltube, with the arithers attached to the margin;ovary superior, flask-shaped, pubescent, 3-5-celled, style slender, stigma head-shaped. Fruitan ovoid or ellipsoid capsule (2.5-)3.5-5 cmlong, woody, opening by 3-5 valves from theapex, valves splitting into 2 layers, many-seeded. Seeds c. 12 min long, flat, with largeterminal wing. Seedling with epigeal germina-tion; cotyledons leafy; first 2 leaves opposite,subsequent ones arranged spiralIy, seedlingleaves often imparipinnate and hipinnate withIncised or lobed leaflets

Other botanical information Chuhrosio

comprises a single, rather variable species. It isdistinctive of the tribetheamong genera

Swietenieoe of the subfamily Swietenioideoe (towhich mahogany (Suitetento) and the impor-tant African timber genera Britondrophrogmoand Khoyo also belong), characterized by thecomparatively large flowers, the more or lessentire stamen tube, and the large number ofseeds arranged in layers, alternately 'head-to-toe'. Molecular phylogenetic studies indicatedthat Chuhrosio is quite distinct from othergenera in the tribe and separation from it maybe warranted

Some authors regard Chuhrosio uelutino(MRoem. ) CDC. as a species (or variety) dis-tinct from Chuhrosio tobu!ons, being a smallertree with more fissured bark and harder wood,and more hairy. However, intermediates are

^I.

4\

\

11

,:;! \

Chuhrusio tobuloris - I, tree hobit, . 2, floweringtwig, . 3, flower in longitudino! section, 4, de-hisced fruit.Source. .PROSEA

^ 2

\

--. J

3

common.

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; (24: intervessel pits minute (S 4 pin));25: intervessel pits small(4-7 pin); 30: vessel-ray pits with distinct borders; similar to in-tervesselpits in size and shape throughout theray cell; 42: mean tangentialdiameter of vesselIumina 100-200 pin; (46: S 5 vessels per squareminimetre); 47: 5-20 vessels per square milli-metre; 58: gums and other deposits in heart-wood vessels. Tracheids and fibres: 61: fibreswith simple to minutely bordered pits; 66: non-septate fibres present; 69: fibres thin- to thick-walled. Axial parenchyma: (78: axial paren-chyma scanty paratrachea1); 79: axial paren-chyma vasicentric; (83: axial parenchyma con-fluent); 85: axial parenchyma bands more thanthree cells wide; 89: axial parenchyma in mar-ginal or in seemingly marginal bands; 92: four(3-4) cells per parenchyma strand. Rays: 98:larger rays commonly 4- to 10'seriate; (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. Mineralinclusions: (136: prismatic crystals present);(142: prismatic crystals in chambered axialparenchyma cells)re. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development In India growthof seedlings proved moderately fast in the first2 years' After 2 years the plants had reached aheight of 1.2-2.1 in, after 3 years 2.8-3.4 inwith a hole diameter of 4-5 cm and after 6

years 5.5 in with a hole diameter of 15 cm, In-dicating a mean annual diameter increment of2.5 cm. A few trees planted on a deep alluvialsoil in northern South Africa were on average37 in tall and 63 cm in hole diameter 49 yearsafter planting, but on a more shallow soil only25 in tall and 47 cm in hole diameter 51 yearsafter plantingChuhrosio tubu!onsnowers and fruits annuaUyThe trees may start flowerlrig when 5 yearsold. The winged seeds are dispersed by wind.

Ecology Under natural conditions in tropi-cal Asia, Chuhrosio tobuloris is found scat-tered in lowland evergreen rain forest or de-ciduous forest up to 900(-1400) in altitude, inareas with an annual rainfall of 1800-3800min and mean annual temperature of 14-27'CIt Is a light-demanding pioneer species, coin-mon in former shifting cultivation areas.Chuhrosio tubuluris should not be planted onsites with heavy soil or excessive moisture. inimpenetrable soillayer less than 60 cm belowsoillevelmay cause failure of plantations

Propagation and planting There

71,000-100,000(-160,000) seeds per kg, andeacll fruit contains about 700 seeds. Germina-tion is fair to easy, up to 90% in I-4(-6) weeksSeeds retain good viahinty for up to 40 monthswhen stored in a cold room (4'C) or freezer,with a viabillty rate of 60-80%, but they canalso be stored at room temperature. The ger-mination rate in the nursery is generally lessand I kg of seed gives about 10,000 viable seed-lings. The seeds can be separated by threshingsun-dried capsules, and do not require pre-treatment. They are sown In light porous soilwith overhead shade or under a mulch cover.Best results have been obtained by raisingseedlings in well-drained boxes and pots beforetransplanting. Watering of seedlings should besparse because they are sensitive to fungaldamping-off. Seedlings are pricked out andtransplanted when about I month old and 6-8cm tall. Planting takes place into open condi-tions. Stump planting is practised in India andChina. Stem and root cuttings are also used forplanting, and grafting, mainly patch budding,for seed orchards. A method for efficient invitro regeneration through organogenesis hasbeen developed.

Management In 1980 a commercial planta-tion of 25 ha was established in northern SouthAfrica, with good results. The spacing was 5 inx 5 in. Weeding around young trees is neces-sary at least once a year. Final stocking isabout 100 treesA1a. Planting trials in Camer-o0n were not successful, with trees showingpoor adaptation. The trees coppice profuselyafter cutting.

Diseases and pests Like many related Me-jigceoe species, Chuhrosio tubuloris is attackedby the shoot borer Hypsipylo robusto, but only40% of the attacked trees develops a inalforma-tion or a fork. This is also the case In planta-tions in Africa. In some countries, e. g. in Laos,the attack of shoot borers discourages planta-tion development. However, It has been re-corded that there is no damage by shoot borersin trees over 8 years old, and some trees showresistance.

Harvesting In Vietnam the rotation periodin plantations for the production of sawlogs is20-25 years, and the cutting cycle in naturalforest in India is 60 years

Genetic resources In many regions in tropi-cal Asia where Chukrosio tobuloris occurs

naturally, it has been exploited heavily for Itstimber, and has become liable to genetic ero-SIon. In some countries it has been bannedfrom logging, e. g. in Vietnam. In India a germ-

CHUKRASiA 169

are

,

170 TIMBERSl

PIasm bank and a seed orchard have been es-tablished after selection of superior trees. Since1999, systematic studies of the genetic vana-tion of Chuhrosio tubu!ons have been under-taken, including the establishment of prove-nance trials, coordinated by CSJRO, Australia.

Prospects Chuhrosio tabuloris is a fairlyfast-growing tree with high-quality wood. It isthus a plantation species with good potentialIn tropical Asia it is rated as having high plan-tation potential on exposed sites, and it mighthave the same potential in Africa, although theresults from the few tests done were variable.

Tree improvement efforts should focus on pro-ductivity characteristics and on ways to mini-inize damage by Hypsipylo. Mixed plantingmay reduce damage by shootborers

Major references Chudnoff, 1980; Ho &Noshir0, 1995; Kannganire & Pinyopusarerk,2000; Mahberley, Pannell & Sing, 1995; Naka-tani at a1. , 2004; von dem Bussche, 1982a; vondem Bussche, 1982b; World Agroforestry Gen-tre, undated.

Other references Delwaulle, 1979; Inside-Wood, undated; 00n at a1. , 2000; Richter &Danwitz, 2000; Shukla, Sharma & mill-Negi,1993; Thangaduraiet a1. , 2003

Sources of illustration Ho & Noshir0, 1995.Authors R. H. M. J. Lemmens

Based on PROSEA 5(2): Timber trees: Minorcommercialtlinhers

elliptical to almost lanceolate, 8-22 cm x 2.5-7cm, base curieate, apex acuminate, thinlyleathery, glabrous, brown, shiny, pinnatelyveined with 8-13 pairs of lateral veins. Inflo-rescence a fascicle arising from the trunk andbranches. Flowers unisexual or bisexual, regu-Iar, whitish; pedice1 18-20 min long, jointed;calyx shortly campanulate, 15-20 mm long, 4-6-10bed, lobes 10-12 min long; corolla absent;male flowers with an androphore 2-6 min longcarrying I whorl of 16-20 stamens; female andbisexual flowers with superior ovary consistingof (3-)5-8 carpels. Fruit consisting of I-6 0voidto globose follicles 3.5-4.5 cm x 2-3 cm, with c3 mm long stipe, rounded at apex, glabrous,red or brown, exuding a inucilaginous gumwhen fresh, indehiscent, I-2-seeded. Seedsovoid, 1.5-2 cm x c. I cm; seed coat fibrousGolo comprises about 100 species and is re-stricted to continental Africa

Ecology Golo couli/10ro occurs in lowlandsemi-deciduous forest up to 600 in altitude.

Genetic resources and breeding Golocoulifioro has a limited area of distributionand could be threatened by habitat degradationand felling, although it has not been classifiedby IUCN as endangered or vulnerable

Prospects The wood of Cold eduliftoru isconsidered to be of fairly good quality, but thetree is too smallto become an important sourceof timber.

Major references Burki11, 2000; Germain &Barnps, 1963; Hall6, 1961.

Other references CJB & SANB1, 2006;Keay, 1958e

Authors M. Brink

COLACAULIFLORAMast

Protologue 011v. , F1. trop. Mr. I: 221 (1868)Family Sterculiaceae (APG: Malvaceae)Vernacular names KOIa mahogany (En)Origin and geographic distribution 0010

eduli/10ro is distributed in southern Nigeria,Gamero0n, Gabon, Congo and DR Congo

Uses The wood of Golo cou!iftoro is suitablefor carpentry and construction.

Properties The heartwood of Cola eauli/10roIs red-brown, with fine texture; the sapwood iswhitish. The wood is durable and reportedIyresistant to termites. When large enough it isconsidered a fairly good timber of the Africanmahoganytype.

Botany Straggling shrub or smalltree up to9 in tall; branches sparsely hairy to glabrous;bark whitish grey, sometimes with lenticelsLeaves alternate, simple and entire; stipulesnarrowly lanceolate, up to 5 mm long, shortlyhairy, caducous; petiole 3-12(-14) min long,sparsely pubescent, brownish; blade narrowly

COLACLAVATAMast.

Protologue 011v. , F1. trop. Mr. I: 222(1868).

Family Sterculiaceae (APG: Malvaceae)Origin and geographic distribution Cold

CIOuoto is distributed in Somalia, Kenya, Tan-zania, Malawiand Mozambique.

Uses The wood of Golo cloudto is used for

construction, tool handles, bows, animal traps,fencing and as firewood. The tree is suitable forproviding shade and as a wayside tree

Properties The wood of Cold cloudto isheavy, hard, tough and termite-resistant.

Botany Evergreen, dioecious medium-sizedtree up to 30 in tall; hole often branched low;bark brownish grey to pale green, smooth;branchlets first stellate-pubescent but soon

glabrous. Leaves alternate, simple and entire;stipules c. 3 min long, caducous; petiole up to6.5 cm long, slightly swollen at base and apex;blade narrowly obovate to elliptical, up to 13(-23) cm x 6(-10) cm, base curieate, apex obtuseto obtusely acuminate, leathery, glabrous, pin-nately veined with c. 10 pairs of lateral veinsInflorescence an axiUary fascicle on I-3-year-old branches. Flowers unisexual, regular; pedi-ce1 5-10 mm long, pubescent; calyx stellate-pubescent; petals absent. Fruit consisting of I-3 club-shaped follicles c. 2 cm x I cm, narrow-ing into a stalk-like base, stellate-pubescent, I-seeded. Seed oblong-ellipsoid, c. 2 cm x 1.5 cm,shiny brown.Golo comprises about 100 species and is re-stricted to continental Africa. Some other Cola

spp. are used as a source of wood in East At-0010 discog!ypremnophyllo Brenan &

A. FD. Jones is a smalltree up to 10 in tall oc-curring in riverme and lowland forest in Tan-zania and perhaps Mozambique; its wood isused for poles, tool handles, animal traps andfirewood, and it is also a shade tree. The woodof Cola ulolomo Brenan, occurring in lowlandforest in Kenya, Tanzania and Malawi, is usedfor poles, tool handles, firewood and charcoalproduction; this species Is also used as a shadetree. Colo usumborensis Engl. is a smalltreeup to 15 in tall occurring in submontane forestin Tanzania. Its wood is hard, heavy and tough,and is used for poles, animal traps, tool han-dles, firewood and charcoal production; ropeand medicines are also obtained from the treeLow-altitude populations of Cola usumborensishave sometimes been confounded with Cold IuheiCheek, a tree from eastern Tanzania, up to 10in tall and sometimes coppiced for poles. GoloIukeiis classified as endangered in the IUCNred list of threatened species, due to habitatloss and growing local population pressure

Ecology Golo CIOuoto occurs in riverIne for-est and forest edges up to 600 in altitude.

Management Golu CIOuoto is recorded to besometimes cultivated on farms in the Usam-bara Mountains of Tanzania, but details arelacking.

Genetic resources and breeding Cold CIO-uoto is classified as data deficient in the TUGNred list of threatened species, indicating thatthere is inadequate information to assess itsrisk of extinction

Prospects Although the wood of Colacloudto is considered to have favourable prop-erties, too little information on the wood prop-erties is available to properly evaluate its pros-

TICa

pects for increased use as a source of timberMajor references Lovett at a1. , 2006; Thu-

Iin, 1999b; Wild & Gonealves, 1979Other references Bandeira, 1998; Cheek,

2002a; Cheek, 2003; CJB & SANB1, 2006;Muir, 1998; Wild, 1961

Authors M. Brink

COLAGREENWAYIBrenan

Protologue Kew Bull. 1956: 144 (1956).Family Sterculiaceae (APG: Malvaceae)Synonyms Cold microcorpo Brenan (1956)Vernacular names Hairy cola (En).Origin and geographic distribution Golo

greenwoyiis distributed in DR Congo, Kenya,Tanzania, Malawi, Zambia, Namibia, Zim-babwe, Mozambique and South Africa.

Uses The wood of Golo greenwoyiis used forconstruction, tool handles, bows, pegs, animaltraps, firewood and charcoal. It is used in tra-ditionalmedicine, but uses are unspecified

Properties The wood is heavy, hard andtough. Ethanol and dichloromethane extractsof the twigs have shown anti-inflammatoryactivity, and an ethyl acetate extract moderateantibacterial activity

Botany Evergreen, usually dioecious, me-dium-sized tree up to 24 in tall; bole up to 50cm in diameter; bark pale brown to grey,rough; young branches densely dark brownhairy. Leaves alternate, simple and entire;stipules subulate-lanceolate, 3-6 mm long,caducous, brown hairy; petiole up to 6 cm long,with pulvinus just below the blade; blade ellip-tical to oblanceolate or obovate-elliptical, up to15 cm x 7 cm, thinly leathery, base broadlycurieate or narrowly rounded, apex obtuselyacuminate, glabrous on both sides at maturity,purplish red when young, pinnately veinedwith 8-12 pairs of lateral veins. Inflorescencean axiUary fascicle on I-3-year-old branchesFlowers unisexual, rarely some bisexual, regu-Iar; calyx 6-11.5 mm long, deeply 4-6-lobed,tube up to 1.5 mm long, lobes elliptical toobovate-elliptical; corona absent; male flowerson jointed pedice1 4-7 min long, androphore2.5-4.5 mm long with 5-12 sessile arithers in asingle row around the apex and 4 rudimentarycarpels immersed in the top; female flowerswith pedicel up to I(-2) cm long with partabove joint markedIy grooved, ovary superior,consisting of 3-5 carpels, styles up to 3 minlong, rudimentary stamens 5 at base of carpels.Fruit consisting of 4-5 obliquely ellipsoid to

COM 171

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172 TIMBERSl

obovoid follicles I-2 cm x 1.5-2 cm, apex

rounded or inucronate, orange-yellow to deeporange when ripe, rusty hairy, I-2-seededSeeds ellipsoid, 13-16 mm x 8-11 mm, testabrown, somewhat wrinkled.In Zimbabwe Co!o greener^oyiflowers in Octo-her-November. 0010 comprises about 100 spe-cies and is restricted to continental Africa.

Ecology Catogreenuioyioccurs in dense wood-land or evergreen forest up to 2200 in altitude.

Genetic resources and breeding 0010greenwoyi has a wide distribution and thereare no Indications of threat by genetic erosion

Prospects The wood of Golo greenwoyiislocally used for various purposes. Too littleinformation is available on its wood propertiesto assess its potential for wider use.

Major references Lovett at a1. , 2006; Ver-doom, 1981; Wild & Gongalves, 1979.

Other references CJB & SANB1, 2006;Germain & Barnps, 1963; Hyde, 2004; Reid ata1. , 2005; Wild, 1961

Authors M. Brink

pounded with clay, and by bathing the afflictedparts in a leaf decoction. In C6te d'Ivoire abark-decoction is applied as vaginal doucheagainst sterility, whereas a preparation of theinner bark is drunk against cough. In CentralAfrica a bark decoction is drunk against inter-costalpain

Properties The heartwood of 0010 latentiois pinkish brown, sometimes figured; the sap-wood is grey-white. The grain is mostlystraight, texture coarse. The wood has a den-sity of about 590 kg/ina at 12% moisture con-tent. It is tough, strong and flexible, bendingwithout breaking. At 12% moisture content,the modulus of rupture is 74 N/mm2, modulusof elasticity 10,100 N/min2, compression paral-Iel to grain 51 N/min2, Janka side hardness3980 N and Janka end hardness 4230 N. The

wood is difficult to work and does not give asmooth finish. It splits easily, and is not dura-ble

Botany Large deciduous tree up to 50 in tall;hole straight, up to 80 cm in diameter, usuallywith concave buttresses I-2 in high and about1.2 in wide at the base; outer bark grey tobrown, smooth, sometimes slightly fissured orcracked, inner bark pink, fibrous; crown largeand spreading; twigs and buds hairy. Leavesalternate, simple, entire or lobed; stipules tri-angular, up to 8 min long, caducous; petiole upto 30 cm long, sparsely hairy, glabrescent;blade broadly ovate, up to 35(-45) cm x 30(-40)cm, base cordate, apex acute, hairy on bothsides when young, glabrescent, becoming red-dish at drying, basal veins 7-9, lateral veins in4-7 pairs. Inflorescence a panicle up to 15 cmlong in axils of upper leaves or on defoliatedbranches, reddish pubescent. Flowers unisex-ual or bisexual, regular, yellow, pink or red,scented; pedicel up to 15 min long, jointed; ca-Iyx campanulate, 5(-7)-lobed, 7-15 min long;corolla absent; male flowers with a shortlyhairy androphore with 2 whorls of 7-10 sta-mens each; female and bisexual flowers withovary generally consisting of 4 carpels. Fruitconsisting of usually 4 indehiscent follicles, onrobust, 7-10 cm long pedicel; follicles ovoid toglobose, up to 6.5 cm long, with a short andslender beak, red or pink, wrinkled, glabrous,4-8-seeded. Seeds ovoid, 2.5-3 cm x 1.5 cm,glabrous, surrounded by a inucilaginous pulp;anI c. 2 min thick. Seedling with epigeal ger-mination

0010 comprises about 100 species and is re-stricted to continental Africa. Cola bruneelii DeWild. , a shrub or smalltree up to 10 in tall also

COLALATERITIA K. Schum.

Protologue Notizbl. Bot. Gart. Berlin-Dahlema 307 (1899).

Family Sterculiaceae (APG: Malvaceae)Chromosome number2n = 40, 42Vernacular names Petit ouara (Fr). Amo-

relra, morelra (Po)Origin and geographic distribution 0010

lotoritio is distributed from Guinea to south-

eastern DR Congo. It has been introducedelsewhere, e. g. into Cape Verde, where it hasnaturalized

Uses The wood of Cola Idleritto is used in

local carpentry and for making casks and bowsIn DR Congo the wood of the buttresses is con-sidered good for making pails and tubs forhousehold use, and clappers made from thewood are attached to hunting-dogs.The fruit pulp is eaten and the seed is chewedin the same way as that of the true kola(mainly Golo acuminoto (P. Beauv. ) Schott &Endl. and 0010 nthdo ryent. ) Schott & Endl. )In C6te d'TVoire leafbuds and young leaves areeaten as a vegetable; they are much appreci-ated during feasts and initiation ceremoniesFibre from the bark is made into cordageIn Sierra Leone sun-dried plant parts aresmoked in a pine for treatment of tuberculosisIn Liberia eruptions, especially on the nostrils,are treated with an ointment of the inner bark

.

occurs in dense humid forest in DR Congo. Itswood is locally used for making tool handles;its leaves and seed pulp are edible.Within 0010 loteriti0 2 varieties are distin-

guished: var. latentto, with peduncle 5-15 minlong, distributed from southern Nigeria to Ga-bon, and var. ingc!oudii(Achev. ) Brenan &Keay, with peduncle less than 3 min long, dis-tributed from Guinea to south-western Nigeria.In Sierra Leone the tree is leafless in April andMay;flowering has been observed from Octoberto May and ripe fruits from February to Sep-ternber. In C6te d'Ivoire Cold loteritio flowers

from August to April; fruiting is from Novem-her to July. Rodents eat the fruits and maydisperse the seeds

Ecology Cold loteritio occurs in rainforest,gallery forest and secondary forest, up to 1000in altitude

Management Cold latentio is sometimesplanted, e. g. in Gamero0n and Cape Verde, butmanagement details are lacking. The 1000-seed weight is 1.7-3.3 kg. Germination takesplace 6-21 days after sowing; the germinationrate is usually high

Genetic resources and breeding In viewof its wide distribution, 0010 loteritio is notthreatened by genetic erosion

Prospects Cola luteritio is a useful source ofwood for local use and of other products, suchas food, fibre and medicines. However, thequality of its wood is probably too poor to ex-Dect increased importance as a source of tim-her

Major references Aubr6ville, 1959b; Bur-kill, 2000; Germain & Barnps, 1963; Hall6,1961; Lovett at a1. , 2006

Other references Adebola & Morakinyo,2005; Carriere, 1999; de to Mensbruge, 1966;Gongalves, 1996; ITvine, 1961; Keay, 1958e;Neuwinger, 2000; Saville & Fox, 1967; Taka-hashi, 1978; T6r6, 2000

Authors M. Brink

making bows. Saplings are used for spearshafts in Nigeria. Twigs serve as chew-sticksThe wood is also used as firewood. The barkfibre is made into cordage. Bark of young sap-lings can be made into very fine cloth;it is alsochewed by women to colour the mouth. Thefruits are eaten raw and the foliage has beenmentioned as a possible fodder. In Nigeria seedpreparations are used against diarrhoea anddysentery. In Niger Cold IOUrifo!to is madeinto love potions.

Properties The wood of Cold IOUrtfo!to isyellow-grey. The texture is fine. The wood isstrong and elastic, bending without breaking,but sometimes splitting

Botany Medium-sized tree up to 30 in tall,often low-branching with arching, sometimesrooting branches; bole up to 80 cm in diameter;young parts rusty stellate-hairy. Leaves alter-nate, grouped at the end of twigs, simple andentire; petiole 2-8 cm long; blade oblong, ellip-tical or oblanceolate, 5-25 cm x 2.5-13 cm,curieate at base, apex obtuse or broadly acumi-nate, leathery, covered with reddish stellatehairs when young, but becoming glabrous laterInflorescence a short axillary cyme, denselyreddish hairy. Flowers unisexual, regular, yel-low or brownish; calyx 4-6 min long, with veryshort tube, 5-10bed with lanceolate lobes, rustybrown hairy outside, margins fringed; corollaabsent; male flowers with arithers on andro-phore in I row; female flowers with pedicel upto 1.5 cm long, ovary superior, consisting of 4-6carpels. Fruit consisting of 4-5 fontcles onshort pedicel; follicles obovoid, up to 5 cm long,brown or red, wrinkled, reddish pubescent, 4-5-seeded. Seeds c. 2 cm x 1.5 cm x I cm, coin-

PIetely covered by a yellow anIIn C6te d'TVoire 0010 IOUrifolio flowers fromAugust to March; fruiting is from June to Feb-ruary. Golo comprises about 100 species and isrestricted to continental Africa. Also present inWest Africa is Cold buntingii Baker f. (bushkola), a tree up to 12 in tall, occurring in theunderstorey of evergreen forest from Liberia toGhana. Its heartwood is orange-brown; thewood is rather hard (Janka side hardness 9490N at 12% moisture content) and heavy (density830 kg/ms at 12% moisture content). In Liberiathe stems are used for native house construc-

tion and as pestles; the fruit pulp is edibleEcology Cold IOUrifolio occurs on riverbanks

in forest and moist savanna; it is often one ofthe dominant species in gallery forest

Genetic resources and breeding In viewof its wide distribution the risk of genetic ero-

COLA 173

COLALAURIFOLIA Mast.

Protologue 011v. , F1. trop. Mr. I: 222 (1868)Family Sterculiaceae (APG: Malvaceae)Chromosome number2n=42

Vernacular names Laurel-leaved kola (En)Komonb610 (Fr).

Origin and geographic distribution ColdIOUrifolio is distributed in West Africa fromSenegal to Nigeria

Uses The wood of Cold IOUrifolio is used for

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174 TIMBERSl

sion of 0010 joltrifolio seems low, although it isconsidered threatened in Senegal.

Prospects 0010 IOUrifolio is a useful localsource of wood, fibre, edible fruits and tradi-tionalmedicine. Too little is known on its wood

properties to evaluate its prospects as a sourceof timber

Major references Aubr6ville, 1959b;Burki11, 2000; Keay, 1958e

Other references Amb6, 2001; Irvine, 1961;ICryn & Fobes, 1959; MEPN, 1997; Natta, Sin-sin & van der Maesen, 2002; Saadou, 1993;Takahashi, 1978.

Authors M. Brink

COMMiPHORAPTERocARPAH. Ferner

Protologue Mein. Mus. natl. Hist. nat. ,Paris n. s. 18: 282-283 (1944)

Family BurseraceaeOrigin and geographic distribution Coin-

in iphoro pterocorpo is endemic to Madagascar,where it occurs in the south-western part of thecountry

Uses The wood of Comintphoro prerocorpoand that of various other Comintphoro speciesis known as 'arofy' in Madagascar. It is locallymuch used for construction, Joinery, shutteringand vehicle bodies. A decoction of the bark of

Comintphoro prerocorpo is used for treating ofulcerated wounds.

Properties The pale brown heartwood is notdistinctly demarcated from the sapwood, but onexposure the sapwood becomes more greyish.The grain is usually straight. The wood islightweight, with a density of about 410 kg/inaat 12% moisture content. It dries easily andrapidly, without risk of distortion. Shrinkagerates from green to oven dry are low: 1.6% ra-dial and 5.1% tangential. At 12% moisture con-tent, the modulus of rupture is about 64N/mm2, modulus of elasticity 6300 N/min2compression parallel to grain 26 N/min2, shear3.7 N/min2, cleavage 18 N/min and Chalais-Meudon side hardness 0.9. The wood saws and

works easily. It holds nails well, and gluingand painting properties are good. The wood isnot durable; it is liable to attack by termites,Lyetus borers and fungi. It is permeable to pre-servatives.

The approximate chemical composition of thewood is: cellulose 33.2%, pentosans 13.8%, 11g-rim 24.6%, ash 2.1% and silica 0.02%. Thesolubility in water is 2.3% and in alcohol-benzene 19.2%. Wood fibres are c. I min long,

with a diameter of 36 pin and a lumen diame-ter of 30 prn. The inflammability of the wood ishighThe plant is recorded to contain navonoids,Ieucoanthocyanins and tannins.

Botany DIDecious shrub or smalltree up to15 in tall; bark with annular appearanceLeaves alternate, often clustered at tips ofbranches, up to 30 cm long, imparipinnatelycompound, with (7-)9 leaflets, glabrous; rachisfinely grooved, whitish; petiolules thin, lateralones 15-25 min long; leaflets orbicular orbroadly ovate, 3-4.5 cm x 2.5-3.5 cm, baserounded, apex acuminate with acumen c. 1.5cm long, pinnately veined with 5-7 pairs oflateral veins. Inflorescence an axillary spikewith flowers in fascicles. Flowers unisexual,regular, small. Intructescences a few togetheron twigs below the leaves. Fruit a compressedovoid drupe 1.5-2 cm long, winged, dehiscentwith 2 valves, with stone c. 12 mm long, cov-ered by false anI at base. Fruiting of Comint-phoropterocorpo is in December.Comintphoro comprises about 200 species andoccurs in tropical and subtropical Africa, Asiaand South America. It is most abundant in the

drier parts of eastern and southern Africa andin Madagascar. In Madagascar the number ofspecies is estimated at 50. Several species ofComintphoro are important for their gum res-ms. the best known being myrrh from Comint-phoro myrrho (Nees) Engl. from Somalia andYemen.

Comintphoro origolensis Engl. CSand corkwood')is a shrub or smalltree up to 5 in tall, distrib-uted in Angola, Zambia, Namibia, Botswana,Zimbabwe and South Africa. Its wood is used

for carving household utensils. The root is animportant source of water in the Kalahari de-sert. In Zimbabwe the root is used in thetreatment of diarrhoea.

Comintphoro Iuluotomentoso Engl. is a smalltree up to 12 in tall, distributed in southernTanzania and northern Mozambique. Its softwood Is used for carvings, spoons, water potsand beehives. The tree is also used for live

fences and for marking boundariesComintphoro glanduloso Schinz ('tall commoncorkwood'; synonym: Comintphoro pyroco, I-thoides Engl. subsp. glo, Iduloso (Schinz) Wild)is a shrub or smalltree up to 10 in tall, distrib-uted from Angola, DR Congo and Tanzaniasouth to South Africa. Its wood is used for

household utensils, and stems are oftenplanted to develop Into live fences.Comintphoro gloucescens Engl. ('blue-leaved

corkwood') is a shrub or smalltree up to 8 intall, occurring in southern Angola and Na-inibia. Its wood is made into household utensilsand the tree is browsed by livestock.Comintphoro hildebrondtii(Engl. ) Engl. is asmall tree up to 10 in tall, occurring in Ethio-pia, Djibouti, Somalia, Kenya and Uganda. Itswood is used for making smallmilk vessels; thestem yields a gumComintphoro woodii Engl. ('forest corkwood';synonym: Coinmjphoro coryoefoli0 011v. ) is asmalltree up to 15 in tall distributed in Mo-zambique and South Africa. Its wood is light-weight and is used for fishing floats. Gum isobtained from the bark. Stem cuttings are of-ten planted to develop into live fences.

Ecology Comintphoro prerocorpo occurs indrier forest and thicket, at 100-900 in altitude.

Genetic resources and breeding Comint-phoro pterocorpo is not considered threatenedby genetic erosion.

Prospects The wood of Comintphoro ptero-euroo is soft and not durable. Nevertheless, itis locally much used in Madagascar, due to itsabundance and its well-shaped hole, and it islikely to remain a useful localsource of wood.

Major references Cainiez & Gu6neau,1972; Gu6neau, Bedel & Thie1, 1970-1975;Gunb-Fakim & Brendler, 2004; Perrier de inBathie, 1946; Sallenave, 1971

Other references Chinemana at a1. , 1985;Coates Palgrave, 1983; Dale & Greenway,1961; Doat & Valette, 1980; Ginett, 1991;Lovett at a1. , 2006; Raharimampionona & Phil.tipson, 2006; Raharimampionona at a1. , 2007;van der Walt, 1986; van Wyk & van Wyk, 1997

Authors M. Brink

.

CupRESSus 175

,

Cupressus lusttonico -planted

vador and Nicaragua. In the 20th century it wasintroduced as a plantation forest tree intotropical Africa, where it is widely planted athigher elevations. It is also planted in SouthAfrlca

Uses The wood is used for construction, fur-niture, poles and posts. It is also suitable forlight flooring, ship and boat building, vehiclebodies, agricultural implements, boxes andcrates, interior trim, Joinery, toys and novel-ties, turnery, draining boards, veneer and ply-wood, hardboard and particle board. The woodis used for paper making, e. g. in Ethiopia andKenya. It is a good fuelwoodEssential oil from the leaves, twigs andbranches of the tree is used as an adjuvant andperfume in soaps, room sprays, deodorants andother products. Cupressus lusttunico is plantedas an ornamental(e. g. as a Christmas tree), asa shade tree, and in windbreaks and livefences. It is used for making toothbrushes andbrooms.

The bark is used as an astringent. The leavesare used to treat catarrh and headache, leafsap to treat skin diseases. Essential oil fromthe leaves is used in the treatment of Theurna-tism, whooping cough and as a styptic. Thevapour from a leaf decoction is inhaled severaltimes a day for treatment of flu. Some ethnicgroups in Mexico use the leaves against cancer.In Gamero0n the leafjuice is used to cure skindiseases and the leaves are used to protectstored grain from Insects.

Production and international trade Forest

plantations of Cupressus lusttonico cover anestimated 70,000-80,000 ha in Kenya. InEthiopia the area was estimated at 10,000-

CupREssusLUSiTANicAMill.

Protologue Gard. dict. ed. a Cupressus n. 3(1768)

Family CupressaceaeChromosome number2n=22

Synonyms Cupressus benthomiiEnd1. (1847),Cupressus JindleyiKlotzsch ex End1. (1847).

Vernacular names Mexican cypress, EastAfrican cypress, Portuguese cedar, cedar of Goa(En). Cypr6s du Portugal, c6dre de Goa, cypr6sde Goa, cypr6s du Mexique (Fr). Cipreste doBUGaco, falso cedro do Bugaco, cedro do Bugaco(Po). Msanduku (Sw).

Origin and geographic distribution Cu-pressus lusttonico is naturally distributed inMexico, Guatemala, Belize, Honduras, EI Sal-

o0

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,^

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176 TIMBERSl

15,000 ha at the end of the 1980s. Essential oilis exported from Madagascar

Properties The heartwood is yellowish, palebrown or pinkish, sometimes streaked orvariegated; the sapwood is paler and 3-7.5 cmwide. The wood becomes paler on exposure.The grain is straight to irregular, texture fineand even. Resin cells may be present, appear-ing as occasional brown streaks. Freshly sawnwood has a faint cedar-like scent

The physical and mechanical properties of thewood are very variable. The density is 380-545(-650) kg/in' at 12% moisture content.Shrinkage rates are 1.7% radial and 3.1% tan-gential from green to 12% moisture content;from Madagascar 4.4% radial and 6.8% tangen-tial from green to oven dry has been reportedThe wood seasons rapidly with minimal split-ting or surface checking; in kiln drying, hightemperatures may result in distortion. At 12%moisture content, the modulus of rupture is69-85(-165) N/mm, , modulus of elasticity7030-9590(-10,300) N/min2, compression par-allel to grain 23-41(-56) N/min2, shear 11.4N/min2, cleavage 25 N/mm radial and 46 N/mintangential, and Janka side hardness 2050-2720 N

The wood saws well and is easy to work withhand and machine tools. It finishes well and

has good nail-holding properties, but boring aclean hole is difficult. It stains and polisheswell. The wood is suitable for moulding andpeeling.Reports on the durability of the heartwood arecontradictory. The sapwood is not susceptibleto Lyetus borers, but susceptible to attacks byother insects and fungi. Both the heartwoodand the sapwood are resistant to impregnationwith preservatives, but incising Improves re-tention

The wood fibres are (IIJ1.9-2.5(-29) mmlong and (13-)27-32(-39) prn wide, with a cellwall thickness of 3-3.5 pin. The fibres contain62-63% bolocellulose (with 39-40% or-cellulose)and 31-33% 11gnin. The wood can be pulpedsatisfactorily using the sulphate process, withpulping yields of 38-51%. Unbleached pulp hasacceptable tensile and bursting strengths, butthe tearing strength is too low for use in pack-aging paper; bleached pulp is suitable for print-ing and writing papersDistillation of leaves, twigs, fruits or flowersyields 0.05-3% essential OIL reported coinposi-tions vary widely. Essential oilfrom the leavesand hexane leaf extracts has shown antifungalactivity against skin pathogens, which sup-

ports the use against skin diseases in Camer-o0n. A crude ethanol extract of the leaves has

shown cytoxicity in a range of cancer celllines,with celldeath being due to apoptosis.

Description Evergreen, monoecious, medium-sized to fairly large tree up to 35 in tall; holestraight, cylindrical, up to 200 cm in diameter,often buttressed; outer bark of young treessmooth, orange-brown to red-brown, becomingvertically grooved, grey and exfoliating in largestrips with age; crown pyramidalin youngtrees, flat-topped in older ones; branchesspreading or ascending, ends often droopingLeaves decussately opposite, simple, scale-like,on ultimate branchlets rhombic and I-2.5 min

long, on leading branchlets up to 10 min long,apex incurved and acute, margin minutelytoothed, green or glaucous-green. Male coneterminal, solitary, oblong, more or less quad-rangular, 3-5 min x 2-2.5 mm, yellowish greenwhen young, becoming pale brown when ina-ture; scales 10-16(-18), decussately opposite,peltate, slightly keeled, each bearing 3-4 pol-Ien sacs. Female cone terminal, solitary or

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Cupressus lusttonico - I, tree hubit, . 2, bronchletwith mole cones, . 3, brunchlet with fernole cone, .4, seeds.Redrown grid odopted by Achingd Satin Nur-hornon

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grouped, mature one globose-angular, 10-18(-diameter, green or purplish-20)

glaucous maturing to brown; scales 6-8(-10),decussately opposite, peltate, bossed, rough,pale brown to reddish brown, each 8-12-seeded. Seeds somewhat angular and slightlyflattened, 3-4.5 min x 3-4 mm, brown or yel-lowish brown, usually with 2 wings I-1.5 mmwide

Other botanical information Cupressuscomprises about 15 species, distributed inNorth and Central America, the Mediterra.nean region, the Middle East, the Himalayasand China. Cupressus ingcrocorpo Hartw. exGOTdon (Monterey cypress) from California(United States), a tree up to 25 in tall with ahole diameter up to 170 cm, has also been in-troduced into tropical Africa, e. g. in Kenya,Tanzania, Zimbabwe and Madagascar. InKenya it was formerly common, but it has be-come rare. In Madagascar it was successfullyplanted as an ornamental and for windbreaks;here the wood is rarely used except as fire-wood. Cupressus semperuirens L. onediterra-nean cypress) from the Mediterranean region,a tree up to 40 in tall with a hole diameter upto 200 cm, has been planted as a timber tree inTanzania. The wood is valued for its durabUityIt is unclear, however, to what extent it is dis-tributed and used in tropical Africa nowadaysIn Cameroon decoctions of the branches andleaves are reported to be used to treat peptlculcers, haemorrhoids and menstruation prob-Iems. Cupressus torulosu D. Don (Himalayancypress or Bhutan cypress) from the Himalayasand China, a tree up to 40 in tall with a holediameter up to 150 cm, has been introducedinto various tropical African countries, lnclud-ing Kenya, Tanzania, Malawi and Zimbabwe.The wood is considered suitable for cabinetwork, furniture, art articles, construction, fenceposts, poles and railway carriage making, butits actual distribution and use in tropical at-rica is unclear. In the timber trade no distinc-tion is made between the different Cupressusspp.

Anatomy Wood-anatontical description OAWAsoftwood codes):Growth rings: 40: growth ring boundaries dis-tinct; (41: growth ring boundaries indistinct orabsent); 43: transition from earlywood to late-wood gradual. Tracheids: 44: tracheid pittingin radial walls (predominantly) uniseriate (ear-Iywood only); (55: latewood tracheids thick-walled (double wall thickness larger than ra-diallumen diameter)); 56: torus present (pits

mm In

in earlywood tracheids only). Axial paren-chyma: 72: axial parenchyma present; 73: axialparenchyma diffuse (evenly scattered through-out the entire growth increment); 76: trans-verse end wafts smooth. Ray composition: 80:ray tracheids absent or very rare; 81: cell wallsof ray tracheids smooth; 85: end walls of rayparenchyma cells smooth (unpitted); 87: bon-zorital walls of ray parenchyma cells smooth(unpitted); (89: indentures present). Cross-fieldpitting: 93: cross-field pits cupressoid; 98: I-3pits per cross-field (earlywood only). Ray size:103: average ray height medium (5-15 cells);107: ray width exclusively uniseriateor. Baas & I. Hemz)

Growth and development Initial growthof Cupressus lusttonico is fast, with an annualincrease in height of up to 1.5(-2) in during theearly years, Flowering occurs in the driest timeof the year. Female cones take 2 years to ina-ture. Trees normally start bearing frult whenthey are 6-9 years old.In Burundithe average height of trees was 60cm after I year, 125 cm after 2 years, 15 inafter 15 years, and 25 in after 37 years' Thehole diameter was 39 cm and 75 cm after 15and 25 years, respectively. In the Usambaramountains in Tanzania (altitude 2200 in, aver-age annual rainfall 900-1200 min) 57-year-oldCupressus lusttonico, planted at a density of155 treesA1a, had an average height of 28 in, anaverage diameter at breast height of 54 cm,and a standing volume of 400 ina per ha. Invarious locations in Kenya 24-26-year-oldplantations, with a density of 213-401trees/ha,had a standing volume of 171-443 ina per haIn Madagascar good growth of Cupressus lust-tonico, planted between 1200 in and 2000 inaltitude with an annual rainfall of about 2000min, leads to a tree height of 20 in after 15

and an annual increment in standingyears,

volume of 16 in3ftia

Ecology Cupressus lusttonico occurs at (500-)1000-4000 in altitude in areas with an averageannual temperature of 12-30'C, an averageannual rainfallof 800-1500 mm and a dry sea-son not longer than 2-3 months, but also invery humid climates with an average annualrainfall up to 4000 min. In general, it is notdamaged by occasional frost or snow. It prefersdeep, moist, well-drained, fertile, neutral toslightly acidic loamy soils. It does not toleratewaterlogged soil. Natural regeneration is goodin clearings and in burnt areas. Cupressus lust-tonico has become an invasive species in Ma-lawi.

CupREssus 177

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178 TIMBERSl

Propagation and planting Cupressus lust-tonico is normally propagated by seed, but rootcuttings can also be used, and micropropaga-tion with hypocotyl explants has been success-ful. Wildlings may also be planted. The 1000-seed weight is 3-6.5 g. Seed for propagationshould be collected when the cones start toturn brown. After the cones have been dried in

the sun untilthey open, the seeds can be sepa-rated by sieving and be sown in seedbeds.Normally germination takes 20-35 days. Seed-lings are planted out at a spacing of2-3 in x 2-3 in. In Kenya plantations for sawlogs areplanted at a spacing of 2.5 in x 2.5 in (1600treesA1a; later thinned to about 250 trees/}Ia),those for pulpwood at 2.75 in x 2.75 in (1320treesA1a). To prevent soil erosion, Cupressuslusttonico is sometimes underplanted withother plants, but it is not suitable for inter-cropping with crops

Management Weeding is necessary duringthe first years after planting. The trees arenormally pruned and thinned several times. InKenya plantations for sawlogs are recoin-mended to be thinned 3-4 times, resulting in afinal density of about 250 treesA1a. Whengrown as a live fence the trees are trimmed.Cupressus lusttonico needs to be protectedfrom fire

Diseases and pests Cupressus lusttonico isseverely attacked by the cypress aphid Ginorocupressi, causing branches to turn yellow andto dry out. Trees may die, but partial or totalrecovery has also been observed. In Africa Ci-noro cupressi was first discovered in Malawiin1986, and since then it has also been detectedin various Cupressoceoe in DR Congo, Rwanda,Burundi, Kenya, Uganda, Tanzania, Zambiaand Zimbabwe. In the early 1990s, it was esti-mated that over 75,000 ha of Cupressus jusi-tonico in Kenya were attacked, 15,000 ha inTanzania, 4500 ha in Uganda, and over 60,000ha in Rwanda and Burunditogether. Biologicalcontrol of Ginuro cupressi using natural ene-inIes is being investigated. Differences in sus-ceptibility between genotypes of Cupressuslusttonico and hybridization with less suscep-tible Cupressus spp. , such as Cupressus toru-1080, may offer scope for selection and breedingfor resistance.

Harvesting Cupressus lusttonico producespoles at 10 years after planting, and generalpurpose timber after 20 years. In Kenya plan-tations for sawlogs are harvested when theyare 30 years old, whereas plantations for pulp-wood are harvested when they are 15-20 years

old.

Yield In plantations in Burundi, the indus-trial wood volume of individual trees rangesfrom about I ina in trees with a hole diameter

of 38 cm to about 2.6 ms in trees with a hole

diameter of 57 cm. Yield tables for Kenya indi-cate possible timber yields of about 400 ina perha after 25 years and about 500 ing per ha after30 years, but actual yields are lower, due tofactors such as suboptimalthinning and prun-ing practices. Logs are generally well shaped,straight and cylindrical, but wood fromyounger plantations contains a fair amount oflow-grade knotty material

Genetic resources In its natural distribu-

tion area in Central America, Cupressus lust-tonico is not considered to be at risk. Moreover,it has been planted over large areas outside itsnatural distribution area. Long-term seed stor-age behaviour is orthodox.

Breeding The main selection and breedingobjective in Cupressus lusttonicu should beresistance to the cypress aphid. Accessionsfrom Kenya, Uganda and Tanzania have beenincluded in progeny trials in Tanzania, andhave been planted in seed and breeding or-chards.

Prospects Cupressus lusttonico has becomean important forest plantation tree in tropicalAfrica, but nowadays the cypress aphid is aserious threat. This pest is difficult to controland planting of Cupressus lusttonico is notrecommended until a solution becomes avail-able

Major references Bein at a1. , 1996; Bolza &Keating, 1972; Bryce, 1967; Chudnoff, 1980;Fanon, 2005; Maundu & Tengnas, 2005;Muchiri, 1991; Obiri, 1994; Palmer at a1. , 1986;Takahashi, 1978

Other references BOTota, 1979; Brenan &Greenway, 1949; Carmo & Fraz50, 1989;Chauvet, 1968; Chifundera, 2001; Ciesla, 1991;Franco & Schwarz, 1985; Oilbert & Bellefon-tame, 1973; Hemz, 2004; 1<1wuso & Malteki,2003; Kuiate at a1. , 2006a; Kuiate at a1. , 2006b;Lop6z at a1. , 2002; Madoffe & Chainshama,1989; Moorthy at a1. , 1977; Ngugi, Mason &Whyte, 2000; Noumi & Dibakt0, 2000; Obiri,Giathi & Massawe, 1994; Orondo & Day, 1994;Pukkala & Fobjonen, 1993; Wimbush, 1957

Sources of illustration Fanon, 2005.Authors M. Brink

,

CYLICODISCUSGABUNENSIS Harms

Protologue Engl. & Prantl, Nat. Pflanzen-fom. , IIJV Nachtr. I: 192 (1897)

Family Mimosaceae (Leguminosae - Mimo-soideae)

Synonyms Pintodenio gobunensis (Harms)Roberty (1954)

Vernacular names Okan, denya, Africangreenheart (En). Okan, bouemon (Fr).

Origin and geographic distribution Cyli-codiscus 806une, ,sis occurs in the forest zonefrom C6te d'Ivoire to Gabon and Congo.

Uses The wood (trade names: okan, denya)isused for heavy construction including marineconstruction, sluice gates and bridges, heavyflooring, joinery, vehicle bodies, mine props,shipbuilding especiaUy for decking, furnitureincluding garden furniture, sporting goods,agricultural Implements, railway sleepers,carving and turning. In Nigeria it is used formaking pontoons. It is also used as fuelwoodand for charcoal productionThe foliage is browsed by sheep and goats. Thebark serves as a soap substitute and as a fishpoison

A bark decoction administered by enema isused to treat stomach-ache, and it is drunk orused in a bath as an anodyne and against vom-iting, venereal diseases, malaria, psoriasis andrheumatism. A macerate of the leaves is used

against inIgralneProduction and international trade The

annual export of okan logs from Gabon aver-aged 13,600 ina in 2000-2004. In 2003 Camer-o0n exported about 70 ing of okan logs andabout 7900 ms of sawnwood, of which 6000 in3to Europe; in 2004 it exported about 8500 ing of

okan sawnwood, of which 6700 mato Europe.Properties The heartwood is yellowish

brown, often with a slight green tinge, on expo-sure darkening to reddish brown with a yellow-ish or greenish tinge; it is distinctly demar-cated from the 5-8 cm thick, pale pink sap-wood. The grain is interlocked, texture moder-ately coarse. The wood is often slightly stripedand lustrous, and has an unpleasant smellwhen freshly cut.The wood is very heavy and hard. At 12% ino1s-ture content, the density is 770-1100 kg/inaThe rates of shrinkage during drying are inod-erate to high, from green to 12% moisture con-tent 3.0% radial and 3.5% tangential, and fromgreen to oven dry 4.0-7.3% radial and 7.2-10.4% tangential. The wood should be air-driedrather slowly or be carefully kiln-dried, toavoid deformation. After drying, the wood ismoderately stable in service.At 12% moisture content, the modulus of rup-ture is (82-)129-230 N/mm2, modulus of elas-ticity 14,700-22,600 N/min2, compression par-allel to grain 64-108 N/min2, shear 8-22N/min2 cleavage 12-29 N/mm, Janka sidehardness 10,600-12,800 N and Janka endhardness 11,340-13,740 N.The wood is difficult to saw and work, withsome dulling effect on cutting edges, and forceis required. Stellite-tipped saws are recoin-mended. It is difficult to obtain a smooth finlshbecause of the interlocked grain. Planing showsbest results at a cutting angle of 15'. The woodholds screws and nails well, but pre-boring isneeded. The glulrig, staining, polishing andpainting properties are satisfactory, but foroptimal polishing effect the use of a filler isrequired. Turning gives good results. The woodis not suitable for veneer or plywood produc-tlOn

The heartwood is very durable. It shows excel-lent natural resistance to wood decay fungi andis also resistant to dry-wood borer, marineborer and termite attacks. It does not absorbpreservatives. The sapwood is liable to fungiand powder-post beetles and should be re-moved before the wood is processed; it is resis-

The resistance to wood-tant to impregnationrotting fungiis due to the presence of coin-pounds such asisookanin and okaninThe intake and digestIbility of the foliage Ishigh for goats. F1avonoids, saponins, tannins,polyphenols, coumarins, triterpenes and/orsterols, and reducing sugars were detected inthe ethyl acetate extract of the stem bark. Theextract also showed antimicrobial activity

CYLICODISCUS 179

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180 TIMBERSl

against, among others, Stophylococcus aureus,Proteus uulgoris and Bacillus cereus. In testswith rats, an ethyl acetate extract of the barkproduced significant anti-diarrhoeal effect. Thebark showed significant antiplasmodialactivityin tests with mice, which supports the useagainst malaria in traditional medicine.

Adulterations and substitutes The woodof Cylicodiscus gobunensis is considered a sub-stitute for that of azob6 (LOphtr0 o10to Banksex P. Gaertn. ) from Africa and greenheart(Ohiorocordium rodiei (Schomb. ) Rohwer,H. G. Richt. & van der Werff) from tropicalAmerica. The timber of P4ptodeniostrum ofri-corium (Hook. f. ) Brenan is similar and alsotraded as African greenheart

Description Large to very large tree up to60 in tall; hole straight and cylindrical, branch-less for up to 30(-35) in, up to 170(-300) cm indiameter, broadened at base, sometimes withlow, thick buttresses or fluted; bark surfacewith rectangular scales, blackish brown, innerbark creamy to yellow, fibrous, with some vis-cid, yellowish exudate; crown spreading, hemi-spherical; twigs glabrous, blackish. Leavesalternate, bipinnately compound with I-2

pairs of pinnae; stipules absent or early cadu-cous; petiole c. 2 cm long, with gland at apex;leaflets alternate, 5-10 per pinna, with petio.Iule 2-3 min long, elliptical to lanceolate, 4-10cm x 2-5 cm, usually slightly asymmetrical atbase, acuminate at apex, glabrous. Innores-cence an axillary or terminal spike-like falseraceme up to 15 cm long, hairy, densely flow-ered. Flowers bisexual, regular, 5-merous;pedicel c. 0.5 mm long; calyx with c. 1.5 mmlong tube, toothed, reddish; petals free, oblongto lanceolate, c. 3 mm x 0.5 mm, whitish oryellowish; stamens 10, free, c. 5 min long;ovary superior, ellipsoid, c. 1.5 min long, withc. I min long stipe, hairy, style slender. Fruit aflattened linear pod up to 90 cm x 5 cm, red-dish brown, scaly, reticulately veined, dehis-cent at one side, many-seeded. Seeds oblong,flat, up to 7.5 cm long including the paperywing surrounding the seed, brown. Seedlingwith epigeal germination but cotyledons re-maining within seed coat; hypocotyl c. I cmlong, epicoty15-10 cm long

Other botanical information Cylicodiscuscomprises a single species and seems to have arather isolated position

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; 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 F1m;43: mean tangential diameter of vesselluminaZ 200 pin; 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 fibrespresent; 69: fibres thin. to thick-walled; (70:fibres very thick-walled). Axial parenchyma:79: axial parenchyma vasicentric; 80: axialparenchyma anform; 81: axial parenchymalozenge-allform; (83: axial parenchyma conflu-ent); 91: two cells per parenchyma strand; 92four (3-4) cells per parenchyma strand. Rays(97: ray width I-3 cells); (98: larger rays coin-monly 4- to 10'seriate); 104: all ray cells pro-cumbent; 1/5: 4-12 rays per mm. Mineral in-clusions: 136: prismatic crystals present; 142prismatic crystals in chambered axial paren-

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Growth and development Seedlings toIer-ate shade, and young trees can be found insmall gaps in the forest. The holes of youngtrees are spiny. The trees are deciduous or ev-ergreen. Adult trees have huge crowns thatspread in the upper canopy of the forest. Youngleaves are red. In Gabon trees flower in No-

veinber-April and fruits mature in May-August. In Nigeria trees flower in February-April and June-September, whereas fruitingtrees have been observed in November-MarchIn Ghana fruits are ripe in January-Februaryand MayJune. The winged seeds are dis-persed by wind. They are eaten by several pn-mate species. Natural regeneration Is oftensparse

Ecology Cylicodiscus gobunensis can befound in deciduous as well as evergreen forest,but is most abundant in well-drained sites inmoist semi-deciduous forest. In Gabon it occursscattered in primary forest, but may be locallymore abundant in old secondary forest.

Propagation and planting The 1000 seedweight is about 100 g. The seeds do not showdormancy, and germinate in 5-12 days. Wild-lings are collected for planting

Management In general Cylicodiscus go-bunensis occurs scattered and in low densitiesin the forest. In Gabon the average bole volumeis 0.3 mama, but in south-western Gamero0n2.1 mama for hole diameters over 60 cm. InGhana the average stock is estimated at 267ms/kin2 and the total exploitable volume(above 70 cm bole diameter) 866,400 ms withan annual allowable cut volume of 21,600 in3.

Harvesting Felling of the trees is difficultbecause the hard wood has a serious dullingeffect on saws.

Yield One big tree may provide 15-20 ina ofusable timber

Handling after harvest Logs are durableand can be left in the forest for some time be-fore transport. However, if they are left for along time, deep splits develop on exposedtransverse sections, which extend into the logThe logs sink in water and therefore cannot betransported by river.

Genetic resources Cylicodiscus gobunensisis fairly widespread in West and Central at-rica, the export volumes are low, and it is notcommonly cut for local applications because ofthe hardness of the wood. Therefore, it does notseem to be threatened at present

Prospects There is an export market for the

naturally durable timber of Cy!icodiscus go-bunensis. It may replace other durable timberswhich have become rare due to overexploita-tion, such as Miltcio species. However, al-though there is a lack of data, Cylicodiscusgobunensis probably grows slowly, and thismay hamper its sustainable exploitation fromnatural forest.

Major references ATIBT, 1986; Bolza &Keating, 1972; CTRAD Forestry Department,2003; CTFT, 1955; 060n, Chadenson & Haute-vine, 1980; Farmer, 1972; Okokon, Ita &Udokpoh, 2006; Takahashi, 1978; Viniers,1989; Wilks & Issemb6, 2000

Other references Adjanohoun et al. (Edi-tors), 1988; Agyeman at a1. , 2003; ATIBT, 2004;ATIBT, 2005; Ayarkwa, 2000; Bouquet, 1969;Burki11, 1995; de in Mensbruge, 1966; Dia-fouka, 1993; FORTG (Forestry Research Insti-tute of Ghana), 1998; Hawthorne & Jongkind,2006; InsideWood, undated; Keay, 1989;Kouitcheu-Maheku at a1. , 2005; Kouitcheu-Maheku at a1. , 2006; Neuwinger, 2000; Nor-mand & Paquis, 1976; 0wusu, 1998; Phong-phaew, 2003; Raponda-Walker & Sinans, 1961;Vivien & Faure, 1985

Sources of illustration Viniers, 1989;Vivien & Faure, 1985

AuthorsJ. Ayarkwa & F. W. Owusu

DACRYODES 181

DACRYODESBUETTNERl(Engl. ) H. J. Lain

Protologue Bull. Jard. Bot. Buitenzorg, ser.3, 12: 336 (1932)

Family BurseraceaeSynonyms Pochy!o6us buettneri(Engl. ) Engl

(1897)Vernacular names Oatgo, assia (En). Ozigo

(Fr)Origin and geographic distribution Doery-

odes buettneri is a characteristic species ofwestern Central Africa, where its natural dis-tribution is very similar to that of AucoumeoMumeono Pierre (okoum6). It occurs mainly inGabon, where it can be found throughout thecountry, and Equatorial Guinea. Small popula-tions have been recorded from southern Cam-eroon, around the loop of the Dja river, as wellas along the southern border, from Campo toAmbam and south of Djoum; from here its dis-tribution extends into the Sonank6 and Mbomo

regions in northern Congo. Doeryodes buettneriis also present in southern Congo (Mayombe)

Uses The wood (trade name: ozigo)is impor-tant in Gabon and Equatorial Guinea. It is

.

182 TIMBERSl

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Doeryodes buettneri- wild

especially used for plywood (for boxes, packag-ing or furniture) and no longer used much inmassive form, although it can be sawn intoplanks for interior joinery, furniture, carpen-try, moulding, parquetry, interior stairs, panel-ling, vehicle bodies, boat hulls, boxes and poles.Locally, the wood is used for making canoes. Itis suitable for papermaking.In Gabon Doeryodes buettneriis an importantfruit tree. Its fruits resemble those of the but-

ter fruit tree (Doeryodes edulis (G. Don)H. J. Lain), but they are smaller. The oil-richpulp is locally consumed after soaking the fruitin boiling water for about one minuteThe powdered bark is used medicinalIy for thetreatment of burns. The resin is applied to ab-scesses, and it is used as a disinfectant andastringent.

Production and international trade The

volume of ozigo exported from Gabon has de-creased since the late 1990s, from 160,000 in3in 1994-1995, through 61,000 ing in 1999, toabout 20,000 ing and 30,000 ms in 2004 and2005, respectively. In 2005 it occupied 9th placein the list of the most important export timbersof Gabon. As an export timber, oatgo is muchless valued and less traded than okoum6. Theaverage port price and free-on-board price areabout Frs CFA 50,000/in3 (Us$ 931m, ) and FrsCFA 61,000/ms(Us$ 1/31m3), respectively. Thelatter is 35% lower than the price of okoum6According to ITTO statistics, Gabon exportedabout 1000 ina of sawn ozigo in 2004, at anaverage price of Us$ 2431m3, and also 1000 in3in 2005, at an average price of Us$ 2771m3.Despite the decline in international trade inozigo, it must be stressed that the wood is still

much exploited for the interior market in Ga-bon; numerous sawmills produce planks andrafters for construction purposes. In 1999 theremaining standing volume of ozigo was esti-mated at 40 million ing. In Cameroon onlyabout 200 ms of Doeryodes huettneri wood wasexploited in 1998-1999.The fruits are collected in the forest but not

much traded, although they can sometimes befound on local markets

Properties The heartwood is pinkish whiteto grey-cream; It Is not distinctly demarcatedfrom the 5-9 cm wide sapwood. The grain maybe interlocked, crossed or straight; the textureis rather coarse. The wood contains 0.1-0.5%silica

The wood is medium-weight, with a density of500-700 kg/in3 at 12% moisture content. Itdries rapidly, generally without problems, butdrying of thin pieces and plywood sheetsshould be done carefully, because of the risk ofdeformation due to the frequent presence ofinterlocked grain. The rates of shrinkage fromgreen to oven dry are 5.0-7.4% radial and6.3-9.8% tangential. Once dried, the wood ismoderately stable in service. The wood is inod-erately hard, elastic and resistant to shock. At12% moisture content, the modulus of ruptureis 101-161 N/min2, modulus of elasticity8300-13,900 N/min2, compression parallel tograin 38-70 N/mm2, compression perpendicu-Iar to grain 6.5 N/mm2, cleavage 13-25 N/minand Chalais-Meudon side hardness 2.2-5.0

The high silica content and the presence ofinterlocked grain may cause problems duringprocessing and make special equTpment neces-sary. The wood saws easily, but with seriousblunting effect, necessitating the use of stelliteteeth and tungsten carbide tools. The workingangle should be reduced as much as possibleQuartersawn pieces have a ribbon-like aspectand are sometimes marbled; Hat'sawn pieceshave better surfaces after processing. The peel-ing properties are good. Although the wood canbe peeled without prior steaming, steaming isrecommended to improve the quality of thesheets obtained; it is not recommended or use-ful for slicing. Gluing properties are good, butproblems may arise with glues based on phe-nol-formol. Nailing, screwing and finishingproperties are goodOzigo is not durable. It is liable to attack byfungi, termites and other insects, but resistantto marine borers. The wood is not recoin-

mended for temporarily or permanently humidconditions. It is difficult to impregnate with

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preservatives.The wood contains: cellulose 54.4%, pentosans15.7%, 11gnin 29.9% and ash 0.9%. The solubil-ity in I% NaOH is 15.4%.Distillation of the resin yielded 6.7% essentialoil, with as main components terpinen-4-o1(27%), p-cymene (19%) and u-pinene (13%)The essential oil showed in-vitro antioxidant

activity and antibacterial activity against arange of bacteria. The fruit pulp Is rich in oil,like that of Doeryodes edulis. The main coinpo-nents of the essential oil of the frult are or

pinene (29%) and 11monene (23%)Adulterations and substitutes The wood

is difficult to distinguish from that of otherDoeryodes species, such as Igaganga (Docryo-des igogongu Aubr6v. & Penegr. ), OSsabel(Do-cryodes normondii Aubr6v. & Fellegr. ) andsafukala tooth Doeryodes pubescerts ryermoe-sen) H. J. Lain and Doeryodes heterotricho(Fellegr. ) H. J. Lain)

Description Dioecious, medium-sized tolarge tree up to 40(-50) in tall; hole branchlessfor up to 20 in, up to 150 cm in diameter, oftensinuous or with knobs, base thickened; outerbark golden yellow, flaking, inner bark pinkish,

when cut slowly exuding a translucid resin witha terebinth-like odour; crown much-branched,with entangled branches and dark foliage hav-ing a brown tint when seen from below. Leavesalternate, imparipinnately compound wlth 5-8(-10) pairs of leaflets; stipules absent; rachis10-30 cm long, reddish brown hairy; petiolules8-LOG15) mm long, channelled; leaflets OPPo-site, oblong-lanceolate to lanceolate, 12-20 cmx 3-4 cm, base asymmetric, apex acuminate,upper surface dark green and shiny, lower sur-face reddish brown stellate hairy, pinnatelyveined with 10-15 pairs of lateral veins. Inflo-rescence an axillary or terminal panicle 30 cmx 4 cm, with cymose ramifications, stellatehairy. Flowers unisexual, regular; pedice1 I-2mm long, hairy; male flowers with 3 ovate se-pals 2-2.5 min long, hairy outside, glabrousinside, petals 3, 2.5-3 min long, hairy outside,stamens 6, with filaments I mm long, ovaryrudimentary; female flowers similar but some-what larger, with petals 3.5 min long, ovarysuperior, ovoid, glabrous. Fruit an oblong-ellipsoid drupe 2-4 cm x I-1.5 cm, purplishblue at maturity, pulp fleshy, up to 3 mm thick,green-yellow, stone with thin and smooth wall,I-seeded. Seed with cotyledons palmately di-vided into 5 segments.

Other botanical information Doeryodescomprises about 40 species, occurring in theAmerican, Asian and African tropics. In Africa,about 20 species have been found, allof sectionPochylobus. Gabon seems richest with ap-proximateIy 10 species. Docryodes buettneriiseasily recognized by its hairy leaflets, colouredreddish brown below.

Anatomy Wood-anatoimcal description (IAWAhardwood 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: intervessel pits medium (7-10 pin);32: vessel-ray pits with much reduced bordersto apparently simple: pits horizontal(scalari-form, gash-like) to vertical(palisade); 42: meantangential diameter of vessellumina 100-200Urn; 46: S 5 vessels per square minimetre; 47:5-20 vessels per square minimetre; 56: tyloses

Tracheids and fibres: 61: fibres with

simple to minutely bordered pits; 65: septatefibres present; 69: fibres thin- to thick-walledAxial parenchyma: 75: axial parenchyma ab-sent or extremely rare; 78: axial parenchymascanty paratrachea1; 92: four (3-4) cells perparenchyma strand; 93: eight (5-8) cells per

DACRYODES 183

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parenchyma strand. Rays: 97: ray width I-3cells; 106: body ray cells procumbent with onerow of upright and/or square marginal cells;1/5: 4-12 rays per min. Mineral inclusions:159: silica bodies present; 160: silica bodies inray cells; (161: silica bodies in axial paren.chyma cells).(M. Thiam, P. Derienne & E. A. Wheeler)

Growth and development Germinationtakes 30-37 days. The seeds can germinate inforest undergrowth and seem to tolerate lowerlight intensities than those of okoum6. Theseeds may form dense mats below mothertrees, but many of them disappear rapidlySome seedlings may persist in forest under-growth for several years, but more light seemsnecessary for subsequent growth. Young shootsof small plants have a characteristically pinkcolour. In mixed plantings with other species inEkouk (Gabon), the survival rate of Docryodesbuettneri was 30-70% after 3 years, withoutmuch mortality afterwards. When 10 years old,the trees had an average hole diameter of 8-9cm. Although they were in good health, alltrees had a sinuous hole. In Lop6 (Gabon) theannual increase in bole diameter was 4.5 minin dense forest and 6.8 mm in more open Mo-rontoceoe forest.

In north-eastern Gabon most trees seem de-

ciduous, but phenological studies over longperiods and with a sufficient number of indi-viduals are lacking. Leaf loss has been ob-served in July-August, December-Januaryand April. The trees remain leafless for lessthan a month only.In north-eastern Gabon flowering sometimesoccurs in March-April, but mostly in Septem-her-November. Fruiting is from July to March,with a high production of ripe fruits in Sep-ternber-November and February-March. Ittakes (3-)4-5(-6) months between the onset offlowering and the start of fruit production. Thefew data available from GarnerDon indicatethat flowering in this country is in March, andfruiting in September. Pollination is mainly byinsects. The trees fruit massively approxi-mately every 3 years; fruit abortion is commonSeed dispersal is mainly by animals, the fruitsbeing eaten by numerous birds, squirrels andmonkeys

Ecology Doeryodes buettnerioccurs in denseevergreen humid lowland forest (up to 700 inaltitude). It grows well on well-drained soils inequatorial or subequatorial climates with anaverage annual rainfall of 2000-3000 mm. It isalways less abundant near the coast and on

sandy soils than somewhat further inland andon clayey soils. In Gabon it is a tree of the up-per forest layer, most common in the easternpart of the coastal sedimental basin in forestswith Aucoumeo Mumeono and Desbordesio

glot4cescens (Engl. ) Tiegh. , and in the undulat-ing land of the central part of the country, inforest with Desbordesio gloucescens and van-ous Goeso!pintoceoe. In the north-eastern partof the country it is less common, although it islocally abundant. In the Lop6 national park incentral Gabon, Doeryodes buettneriis the sec-ond most characteristic species of the upperforest layer, after AMcoumeo k!umeono; thedensity is 2.7 trees with a bole diameter over70 cm per ha in dense forest, against 0.7treesA1a in more open Morontoceoe forest. Thetree is generally common in old secondary for-est, often in groups. Like AMcoumeo Atomeonu,it behaves as a hellophyte, reproducing in oldclearings, but less effectiveIy so than Au-coumeo hintneono. Docryodes buettneri hasalso been observed in young secondary forest,under Musongo cecropioides R. Br. or Xylopiogethtopico (Dunal) ARIch

Propagation and planting Doeryodes buett-iteri is only occasionally planted, mostly inhome gardens. The only known trials andplantings have been realized in Gabon(Ningh6-rimgh6, M'voum and Bokou6). Stumpswillsprout after 4-8 months, with sprouts fromthe base being more vigorous than those higheron the stump. Unsuccessful experiments withmacro cuttings have been carried out in Ma-kokou experimental station (Gabon)

Management No management practices ofDoeryodes buettneri are known, although treesare sometimes maintained in home gardens.During clearing, trees that are consideredcumbersome are felled. They are not conservedlike other species which are more highly val-ued, such as Iruingio gobonensis (Aubry-Lecomte ex O'Rorke) Baill. , or of which thewood is harder, such as Desbordesio g!duees-certs (Engl. ) Tiegh. Furthermore, if an ozigotree in the forest bears fruits abundantly, itmay be felled to facilitate harvesting.

Diseases and pests In Gabon desiccation ofthe crown has been observed in young planta-tions, but the cause is not clear. Elephantssometimes eat the young shoots and barkVarious predators may destroy the seeds byeating the fruits that have fallen on theground.

Harvesting In natural forest the averageexploitation intensity of ozigo is not more than

,

I-2 trees/ha, but where the species Is locallyabundant the intensity may be much higherThe holes are exploited when they have a di-ameter at breast height of over 70 cm.

Yield Oatgo logs have an average diameterof 70-100 cm. The mean wood volume of ex-

PIOited trees is 7-8 inaHandling after harvest The logs float in

water and can be transported by river. It isrecommended to treat the wood against fungiand insects and to avoid humid conditions bystoring the wood under shelter and not in con-tact with soil. Fruits collected in the forest arenormally consumed directly, but sometimes thepulp is conserved for later usage by drying overa fire after the seed has been removed.

Genetic resources Information on the ge-netic variation of Doeryodes buettneriis not yet

germplasm collection Isavailable, and noknown. In view of its abundance, the speciesdoes not seem threatened with genetic eroslon,except in some parts of its distribution area. InCameroon, for instance, it is considered threat-ened due to forest exploitation pressure andthe relatively low number of trees.

Prospects The wood of Doeryodes huettneriis considered a substitute of okoum6 for the

production of plywood. Although its exploita-tion for international trade is decreasing, it Isstill among the 10 main export timbers of Ga-bon. Furthermore, its utilization in 101Dery Lscurrently well developed in the interior marketof Gabon, indicating that the resource remainsunder considerable pressure. In spite of itsimportance as a source of wood, frults and tra-ditionalmedicines, the species has not been theobject of research. Its biology is poorly knownand it is not intentionally included in tradi-tional agroforestry systems. It seems useful toestablish collections of genetic resources, par-ticularly in Cameroon, where Doeryodes buett-iteriis considered a threatened species. Thismay also become the case in Gabon If its ex-PIOitation remains mainly in the hands ofsawmillers and smallexploiters

Major references Aubr6ville, 1948; Au-br6viUe, 1962a; Bolza & Keating, 1972; CTRADForestry Department, 2003; Normand & Man-aux, 1962; Onaria, 1998; Vivien & Faure, 1985;White & Abernethy, 1997; White, MCPherson &Ahemethy, 1999; Wilks & Issemb6, 2000

Other references ATIBT, 1986; Bourobou-Bourobou, 1994; Christy at a1. , 2003; Cordiez,2000; Cravo at a1. , 1992; CTFT, 1961f; Daly &Martinez-Habibe, 2002; de Saint-Aubin, 1963;Dowsett-Lemaire, 1996; G6rard at a1. , 1998;

Hecketsweiler, 1992; InsideWood, undated;Louis & Fontss, 1996; Louppe at a1. , 1999;Ngoye, 1994; 0bame at a1. , 2007; Senterre,2001; Takahashi, 1978; Vivien & Faure, 1988a;White, 2001.

Sources of illustration Aubr6ville, 1962a;White & Ahemethy, 1997

Authors G. Todou & C. Doumenge

DACRYODEs 185

DACRYODESiGAGANGAAubr6v. & Fellegr.

Protologue F1. Gabon a 86 (1962).Family BurseraceaeOrigin and geographic distribution Doery-

odes igogongo occurs in Garnero0n, EquatorialGuinea, Gabon and possibly Congo

Uses The wood (trade name:Igaganga)is usedfor veneer, plywood, joinery and parquetry. Itis also suitable for construction, furniture andcabinet work, vehicle bodies, ship and boatbuilding, handles, ladders, carvings, toys, nov-elties, turnery, poles and piles. It can be usedfor paper making. The fruit pulp is edible.

Production and international trade Thewood is very similar to 'ozigo' (wood of Doeryo-des buettneri (Engl. ) H. J. Lain) and 'OSsabel'(wood of Doeryodes normondii Aubr6v. &Fellegr. ), and it is often sold together with thelatter. Exports of Igaganga from Gabon arediminishing: 32,690 in3 in 2000, 13,350 ms in2001, 8360 ina in 2002, 9500 in3 in 2003, 7800main 2004 and 2630 main 2005

Properties The heartwood is pale brown orpink; it is not distinctly demarcated from the2-4 cm wide cream-coloured sapwood. Thegrain is straight or interlocked; the texture fineto medium. The wood contains silicaThe wood has a density of 580-670 kg/ina at12% moisture content. It dries rapidly andgenerally without problems. The rates ofshrinkage from green to oven dry are 3.8-6.2%radial and 6.8-9.6% tangential. Once dry, thewood is stable in service. At 12% moisture con-tent, the modulus of rupture is 95-155 N/mm2,modulus of elasticity 9600-13,100 N/mm2,compression parallel to grain 52-67 N/mm2,shear 8-12 N/min2, cleavage 14-25 N/mm andChalais-Meudon side hardness 2.5-4.7.Sawing is rather difficult due to the presence ofsilica, causing severe blunting of tools; stellitesaw teeth and tungsten carbide tools are rec-ommended. The wood finishes, nails, screwsand glues well. Peeling and slicing propertiesare good. The wood is not very durable, beingsusceptible to attack by fungi, termites, dry-

.

186 TIMBERSl

wood borers and marine borers. The heartwoodIs resistant to ImpregnationThe fruit contains an essential oil, the maincomponents of which are orcopaene (16%) andor-humulene (14%).

Botany Dioecious, medium-sized tree up to25 in tall; hole straight, cylindrical, branchlessfor up to 15 in, up to 90 cm in diameter; outerbark grey, flaking, inner bark brownish pink,fibrous, slightly fragrant, when cut exudingsmall amounts of a resin becoming amber-coloured on hardening. Leaves alternate, jin-panpinnately compound with 5-7 pairs of leaf-lets; stipules absent; petiole plano-convex, ra-chis scaly; petiolules c. 5 mm long; leaflets op-POSite, narrowly oblong to lanceolate, 7-11(-15) cm x 2.5-4 cm, base asymmetric, apexacuminate, lower surface slightly hairy or gla-brous, pinnately veined with 8-10 pairs of Iat-eral veins. Inflorescence a panicle 12-15 cmlong, reddish brown hairy. Flowers unisexual,regular; male flowers with pedice1 1.5 min long,sepals 3, c. 3 mm long, hairy outside, glabrousinside, petals 3, hairy outside, covered withglandular hairs inside, stamens 6, with glandu-Iar hairs. Fruit an ellipsoid drupe, blue-blackat maturity, stone ellipsoid, flattened, c. 3 cm x2 cm x 1.5 cm, hard, with rough surface, I-seeded

Doeryodes comprises about 40 species, occur-ring in the American, Asian and African trop-ICs. In Africa, about 20 species have beenfound, all of section Pochylobus. Gabon seemsrichest with approximately 10 species

Ecology Doeryodes igogongu occurs in low-land rainforest up to 550 in altitude.

Management Treatment of the logs withpreservatives after felling is recommended

Genetic resources and breeding Docryo-des igugongo is classified as vulnerable in the2007 IUCN Red list of threatened species, dueto the rate of logging that has taken place andgeneral habitat decline.

Prospects Because of overexploitation inthe past, Doeryodes igogongo has become vul-Derable. Therefore, its use should only be al-lowed where sustainable exploitation takesplace

Major references Aubr6ville, 1962; CIRADForestry Department, 2003; Normand & Man-aux, 1962; Takahashi, 1978; Vivien & Faure,1985.

Other references ATIBT, 2005; Bolza &Keating, 1972; Cravo at a1. , 1992; Daly &Martinez-Habibe, 2002; de Saint-Aubin, 1963;Oxford Forestry Institute, 1997-2004; Parant

at a1. , 2008; Sallenave, 1971; Vivien & Faure,1988a; World Conservation Monitoring Centre,1998a

Authors M. Brink

DACRYODES KLAiNEANA (Pierre) H. J. Lain

Protologue Bull. Jard. Bot. Buitenzorg, ser3, 12: 336 (1932)

Family BurseraceaeSynonyms Perchylobus deliciosus (Achev

ex Hutch. & Dalziel) Penegr. (1934).Vernacular names Monkey plum, African

cherry fruit (En)Origin and geographic distribution Doery-

odes hloineonu is distributed in forest regionsfrom Sierra Leone to Gabon

Uses The wood (trade name: adjouaba) isused in construction and for mortars, axe han-dles and wagons. It has also been recoin-mended for telegraph poles and railway sleep-ers. It is considered suitable for interior join-ery, furniture and parquetry. It is also used asfuelwood and has been used for paper makingThe fruit is eaten raw or cooked; the pulp isboiled or roasted to yield a kind of butter. Theground leaf is an Ingredient of an enema ap-plied against painful menstruation. In C6ted'TVoire Doeryodes Mumeono is used to treattachycardia and cough.

Properties The heartwoodis greyto yenowishgrey-brown with darker streaks, giving it adecorative appearance; the sapwood is whitishgrey and wide, up to more than 10 cm. Thewood contains silica. It is rather heavy, with adensity of 730-940 kg/in3 at 12% moisture con-tent. The wood dries quite fast and well. Therates of shrinkage from green to oven dry are6.4-8.0% radial and 8.3-10.3% tangential. Thewood Is hard. At 12% moisture content, themodulus of rupture is 1/1-229 N/mm2modulus of elasticity 12,900-18,700 N/mm2compression parallel to grain 55-78 N/mm2,shear c. 18 N/mm2, cleavage 18-28 N/mm andChalais-Meudon side hardness 5.8-7.9. Thewood is moderately difficult to work. Theheartwood is resistant to Impregnation withpreservatives.

The leaves contain tannins and traces of sapon-

Botany Dioecious, evergreen, medium-sizedtree up 25(-30) in tall; bole branchless for up to10 in, up to 60(-120) cm in diameter, lowerpart usually straight and slightly angular,higher part often twisted, base with low root

Iris

swellings or root spurs rarely higher than 60cm; outer bark grey-greenish or blackish, veryscaly, with irregular scales leaving yellowishmarks after shedding, inner bark pink-brownor red-brown, with turpentine smell, exuding awatery or gummy liquid when cut; crownrounded, dense. Leaves alternate, Imparlpin-nately compound with (2-)3(-4) pairs of leaf-lets, 15-30 cm long, rusty hairy when young,glabrescent but with persistent hairs on peti-o1e, rachis and inIdvein; stipules absent; petiole2.5-6.5 cm long, rachis 2-8 cm long; lateralpetiolules (2-)8-14 min long, terminal one 10-22 min long, jointed at the top; leaflets OPPo-site, elliptical to ovate, 4.5-18.5 cm x (1.5-)2-6.5 cm, base usually asymmetric, apex acumi-nate, pinnately veined with 6-12 pairs of Iat-eral veins, markedIy looped. Inflorescence aterminal or sometimes axillary panicle 10-22(-30) cm long, densely rusty hairy, with flowersin fascicles. Flowers unisexual, regular, palecreamy; pedice1 2-4 mm long; sepals 3, broadlyovate, c. 2 mm long, densely hairy outside, gla-brous inside; petals 3, ovate, c. 4 mm long,densely hairy outside, papillate Inside; maleflowers with 6 stamens, filaments c. 1.5 mmlong, ovary much reduced; female flowers withsuperior, hairy ovary and reduced stamensFruit an ovoid drupe c. 2 cm x 1.5 cm, slightlylateralIy flattened, slightly pointed at top, or-ange when ripe, with fleshy, fragrant pulp,stone ellipsoid, c. 2 cm X I cm, I'Seeded. Seed-ling with epigealgerminationDocryodes hidineono grows slowly. In the ever-green forest of ITobo (C6te d'TVoire), the aver-age annual diameter growth is 1.2 min undernatural conditions, and 2.6-3.1 min in partiallythinned locations. Doeryodes hmmeono flowersat the beginning of the dry season, October-November in Liberia, where the fruits are ripein January-February. Natural regeneration byseedlings is abundantDocryodes comprises about 40 species, occur-ring in the miencan, Asian and African trop-ics. In Africa, about 20 species have beenfound, all of section Foehylobus. Gabon seemsrichest with approximately 10 species.

Ecology Doeryodes h!Qineuno mostly occursin the understorey of evergreen, deciduous orriverme forest, sometimes in higher layers ofthe canopy. It is widespread and locally abun-dant.

Management The 1000-seed weightis about1.3 kg. Seeds take 15-30 days to germinate.Young seedlings in the nursery should beshaded. Doeryodes Homeuno is not planted in

C6te d'Ivoire, but trees are retained and pro-tected during clearing for agricultural pur-

Marketable logs are mostly defective,poses

with very large ones often being hollowGenetic resources and breeding In view

of its wide distribution and local abundance,Doeryodes foldineono does not seem muchthreatened, but in C6te d'Ivoire and Cameroonit is considered vulnerable.

Prospects The quality of the wood of Do-cryodes hidineo, IQ is much lower than that ofozigo (Doeryodes buettneri(Engl. ) H. J. Lain),due to the logs often being defective, thesmaller diameter and the wLde sapwood. There-fore, its prospects as a timber tree of coinmer-cialimportance seem limited

Major references Aubr6ville, 1962a;Burki11, 1985; Dudek, F6rster & missenbauer,1981; Takahashi, 1978; Voorhoeve, 1979

Other references Adjanohoun & Me Assi,1979; Aubr6ville, 1959a; de in Mensbruge,1966; Durrieu de Madron at a1. , 1998a; Firmin,1999; myn & Fobes, 1959; Neuwinger, 2000;Normand, 1955; Stoker, Winandy & AChi,1996; Vivien & Faure, 1988a.

Authors M. Brink

DACRYODES 187

DACRYODESNORMANDllAubr6v. & Fellegr

Protologue F1. Gabon 3: 72 (1962).Family BurseraceaeOrigin and geographic distribution Doery-

odes normondiiis probably restricted to Ga-bon.

Uses The wood ('OSsabel') is suitable for lightconstruction, flooring, interior trim, Joinery,ship and boat building, vehicle bodies, furni-ture, cabinet work, toys, novelties, turnery,poles, piles, veneer and plywood. It can also beused for paper making. The cooked fruit Iseaten.

Production and international trade Thewood is very similar to 'ozigo' (wood of Doeryo-des buettneri(Engl. ) H. J. Lain) and 'igaganga'(wood of Doeryodes igogongoAubr6v. & Fellegr. );it is often sold together with the latter

Properties The heartwood is pink-buff withan occasional grey tinge; it is not distinctlydemarcated from the sapwood. The grain Isoften interlocked; texture medium to coarseThe wood is medium-weight, with a density of550-660 kg/ing at 12% moisture content. Sea-soning should be done carefully, to avoid split-ting and twisting. The rates of shrinkage fromgreen to oven dry are 3.8-5.8% radial and

188 TIMBERSl

7.1-9.8% tangential. Once dry, the wood isstable in service. The wood is soft. At 12%

moisture content, the modulus of rupture is106-141 N/min2, modulus of elasticity8700-15,500 N/mm2, compression parallel tograin 42-62 N/mm2, cleavage 11-22 N/mm andChalais-Meudon side hardness 2.2-3.6. Thewood is moderately easy to saw when green. Itworks well with hand and machine tools, butcauses some blunting due to the presence ofsilica. It has good nailing and screwing proper-ties, and it glues, sands, paints and varnisheswell. Peeling characteristics are satisfactoryThe wood is susceptible to termite and marineborer attacks.

Botany Dioecious, medium-sized to fairlylarge tree up to 37 in tall; hole straight, cylin-drical, up to 100 cm in diameter, with low but-tresses; outer bark yellowish, flaking, innerbark pinkish, when cut exuding a fragrantresin becoming amber-coloured on hardening;young branches with caducous rusty stellatehairs. Leaves alternate, in tufts at the ends ofbranches, imparipinnately compound with (6-)8(-9) pairs of leaflets; stipules absent; rachis c.40 cm long, rusty hairy; leaflets opposite, al-most sessile, oblong, c. 16 cm x 5 cm, baseasymmetric, apex acuminate, upper surfaceglabrous, shiny, lower surface densely coveredwith caducous rusty stellate hairs, pinnatelyveined with 8-15 pairs of lateral veins. Inno-rescence a terminal panicle, rusty hairy. Flow-ers unisexual, regular, rusty hairy; sepals 3, c3 mm long, hairy outside; petals 3, c. 5.5 mmlong, hairy outside, covered with glandularhairs inside; stamens 6; ovary hairy. Fruit anobovoid drupe c. 3.5 cm long, stone with roughsurface, I'Seeded.Doeryodes comprises about 40 species, occur-ring in the American, Asian and African trop-

In Africa, about 20 species have beenfound, all of section Pochylobus. Gabon seemsrichest with approximately 10 species.

Ecology Doeryodes normondiioccurs in rain-forest

Genetic resources and breeding In viewof its very limited distribution, there may bedanger of genetic erosion, but so far Doeryodesnormondii has not been included in the IUCNRed list.

Prospects The wood of Doeryodes normondiiis suitable for a wide range of applications, butthe extent of its current use is unclear. Becauseof its limited distribution, exploitation of thisspecies should be done with care and in a sus-tainable way.

Major references Aubr6ville, 1962a; Bolza& Keating, 1972; de Saint-Aubin, 1963; Salle-nave, 1971

Other references Daly & Martinez-Habibe,2002; Normand & Mariaux, 1962; Sallenave,1964; Takahashi, 1978.

Authors M. Brink

DACRYODESPUBESCENS ryermoesen)H. J. Lain

Protologue Bull. Jard. Bot. Bintenzorg, ser. 3,12: 337 (1932).

Family BUTSeraceaeSynonyms Pochylobuspubescens Vermoesen

(1923)Origin and geographic distribution Doery-

odes pubescerts is distributed in Congo, DRCongo and northern Angola, possibly also insouthern Gabon

Uses The wood (trade name: safukala) isrecommended for furniture, screens, door andwindow frames, stairs and other light carpen-try, flooring, construction, vehicle bodies, mineprops, poles and piles. The wood is also consid-ered suitable for ship and boat building, toolhandles, agricultural implements, joinery, vats,toys, novelties, turnery, railway sleepers, ve-neer, core stock, hardboard and particle boardThe cooked fruit is eaten. The resin has beenused for torches.

Production and international trade Thewood is difficult to distinguish from the moreimportant 'ozigo' (Doeryodes huettneri(Engl. )H. J. Lain), but is traded separately as 'safu-kala', together with the wood of Doeryodes he-terotricho (Fellegr. ) H. J. Lain. In 2004 about1000 ina of 'safukala' logs were exported fromCongo

Properties The sapwood is pale pink, yellowor grey, and very wide, often extending overhalf the radius of the log; the heartwood issomewhat darker. The grain is straight or in-terlocked; texture medium. The wood is Ius-trous, and ribbon-like markings and smallstripes give it an attractive figure.The wood has a density of 610-690 kg/ina at12% moisture content. Seasoning should bedone carefully, to avoid checking. The rates ofshrinkage from green to oven dry are 4.6-5.7%radial and 6.2-9.5% tangential. Once dried, thewood is stable in service

At 12% moisture content, the modulus of rup-ture is 1/1-163 N/mm2, modulus of elasticity10,500-12,700(-16,000) N/mm2, compression

ICS

parallel to grain 53-62 N/min2, cleavage 15-25N/mm and Chalais-Meudon side hardness 3.1-

6.3

The wood saws slowly and with some difficulty,with a blunting effect on tools; the use of stel-lite-tipped saw teeth is recommended. It worksfairly well with hand and machine tools, butrough patches may be produced on planing. Itdoes not split on nailing and holds nails andscrews well. Peeling characteristics are good,and gluing, boring and sanding properties sat-isfactory. The wood paints and varnishes wellThe wood is not very durable, being susceptibleto attack by fungi; reports on its liability toattack by insects are conflicting. The heart-wood is resistant to impregnation, the sapwoodmoderately resistant.

Botany Dioecious, evergreen, medium-sizedto fairly large tree up to 37 in tall; hole up to150 cm in diameter, often sinuous; outer barkcream-pink coloured, flaking, inner bark palepink, exuding a colourless fragrant resin when

voluminous, hemispherical,cut;

branches long and spreading; young branchesrusty stellate hairy. Leaves alternate, imparip-innateIy compound with (4-)5-8 pairs of leaf-lets, up to 40 cm long, persistently rusty stel-late hairy on petiole, rachis and petiolules;stipules absent; petiole 5-10 cm long; lateralpetiolules up to 5 mm long, terminal one 15-20min; leaflets opposite, oblong to oblong-lanceolate or oblong-obovate, 5-20 cm x 2-7cm, base rounded, sometimes asymmetric, apexlong-acumlnate, upper surface glabrous excepton inIdvein, lower surface with persistent rustystellate hairs, pinnately veined with 10-18pairs of lateral veins. Inflorescence a panicleup to 30 cm x 4 cm. Flowers unisexual, regular;pedice1 I-4 mm long; calyx 3-10bed, 3-4 mm x2-3 mm, segments ovate; petals 3, ovate, 4-5min x 3 mm; stamens 6, filaments c. 2 minlong; ovary ovoid. Fruit an oblong-ellipsoiddrupe 2-3 cm x I-1.5 cm, violet, glabrous,stone I-seeded

Doeryodes comprises about 40 species, occur-ring in the American, Asian and African trop-ics. In Africa, about 20 species have beenfound, all of section Foehylobus. Gabon seemsrichest with approximately 10 speciesThe wood of Docryodes heterotricho (Penegr. )H. J. Lain is very similar to that of Doeryodespubescerts and is also known as 'safukala', al-though it has also been called 'mouganga'. Do-cryodes heterotricho is a medium-sized tree upto 25 in tall, with a bole up to 100 cm in diame-ter, distributed in Gabon and Congo. The wood

has a density of 610-760 kg/ms at 12% ino1s-ture content. The rates of shrinkage from greento oven dry are 3.0-6.1% radial and 7.5-8.9%tangential. At 12% moisture content, themodulus of rupture is 1/9-201 N/mm2,modulus of elasticity 10,500-15,400 N/min2,compression parallel to grain 56-74 N/mm2,shear 10-14 N/mm2, cleavage 25-33 N/mm andChalais-Meudon side hardness 4.1-7.4. Thefruit is edible. A decoction of the leaf is takento treat whooping cough

Ecology Doeryodes pubescerts occurs in low-land rainforest and along rivers, sometimesunder inundated conditions.

Management In an inventory in Cabinda(Angola) in the 1960s, an average of 11.8 treesper ha was found in an area of 16 ha, with 71%having a hole diameter of less than 50 cm, and7% having a diameter of over 80 cm.

Genetic resources and breeding In viewof its limited distribution, there may be somedanger of genetic erosion, but Doeryodespubes-certs is notincluded in the TUGN Red list

Prospects It is unclear to what extent andfor which purposes the wood of Doeryodes PM-beseens is currently used, but it seems suitablefor a wide range of applications.

Major references Bolza & Keating, 1972;Fouarge & G6rard, 1964; Liberato at a1. , 2002;Normand & Mariaux, 1962; Sallenave, 1971.

Other references Anonymous, 1966; ATIBT,1986; ATIBT, 2004; ATIBT, 2005; Aubr6ville,1962a; CIRAD Forestry Department, 2003;Daly & Martinez-Habibe, 2002; de Saint-Aubin,1963; Normand & Paquis, 1976; Takahashi,1978; Troupin, 1958

Authors M. Brink

crown

DAI, BERGIA 189

DALBERGIABARONllBaker

Protologue Journ. Linn. SOC. , Bot. 21: 337(1884).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Vernacular names Voamboana, panssandrebrun, panssandre de Madagascar, panssandrerouge des marais (Fr)

Origin and geographic distribution Do16er-gio boroniiis endemic to eastern Madagascar.It has occasionally been planted elsewhere, e. g.in Tanzania

Uses The wood of Do16ergio boroniiis usu-ally not distinguished from that of some otherDaibergio species, notably Do!bergio monticoloBosser & R. Rabev. The latter was not distin-

190 TIMBERSl

.

Do16ergio boronii- wild

gutshed from Do!bergio boronii until recentlyThe wood is one of the so-called rosewoods('Madagascar rosewood', 'pansander'), whichare much in demand for cabinet making, furni-ture, marquetry and parquet flooring. It is oneof the favoured woods for musical instruments,especially guitars, not only because of its beau-tiful colour and venation, but also because ofits clearness of tone. It is also suitable for shin-

gles, exterior and Tritenor trim, Joinery, carpen-try and framing, ship and boat building, vehi-cle bodies, precision equipment, carvings, toysand novelties, turnery, pattern making, veneerand plywood. It is used for carving traditionalart by the Mahafale people. In the past thewood was used exclusively for the constructionof houses for royal people.

Production and international trade The

wood is still traded on the international mar-

ket, usually in small amounts and at highprices, for special applications such as musicalinstruments. In recent years it has replacedBrazilian rosewood (from Do!bergio wigroryell. ) Benth. ) because this South-Americanspecies has been included in Appendix I ofCITES as an endangered species. It is oftentraded as quarter-sawn pieces of comparativelysmall dimensions. In 1999 Madagascar offi-cially exported about 1500 ina of pansanderfrom different Daibergio spp. , but according toother estimations about 3200 ina was exported.Sculptures made of Daibergio boronii wood arefound on local art markets in Madagascar

Properties The wood of Daibergio boroniiissimilar to that of Do16ergio monticolo and thetwo are not distinguished in trade. The follow-ing description refers to both species. The

heartwood is greyish yellow-brown to reddishbrown or dark brown, often with darkerstripes, and distinctly demarcated from thesapwood. The grain is generally straight, tex-ture fine and even. Fresh wood has a sweetishsmell.

The wood is moderately heavy to heavy, with adensity of 620-950 kg/ing at 12% moisture con-tent. It air-dries satisfactorily but slowly;turned pieces used for precision equipment ormusical instruments should be dried thor-

oughly to avoid later distortion. The rates ofshrinkage are moderate, from green to ovendry about 4.1% radial and 7.6% tangentialOnce dry, the wood is very stable in service.At 12% moisture content, the modulus of rup-ture is 132-221 N/mm2, compression paralleltograin 58-86 N/mm2, cleavage 14-20 N/mm andChalais-Meudon side hardness 2.9-7.8

The wood works well, both with hand tools andmachine tools. It finishes well, taking a beauti-ful polish. The nailing properties are moderateand pre-boring is needed. Finishing with oil-based paint gives moderate results, and thegluing properties are variable. The wood is suit-able for sliced veneer. It is moderately durable,and resistant to termites. The heartwood is

very resistant to treatment with preservativesAdulterations and substitutes The wood

of several other Do16ergio species from Mada-gascar is traded as Madagascar rosewood orpansander

Description Deciduous medium-sized treeup to 25(-30) in tall; hole usually short, branch-less for up to 6(-20) in, up to 100(-140) cm indiameter; bark whitish to pale brown, cracked;young branches short-hairy, brown, olderbranches glabrous, longitudinalIy striped,blackish brown. Leaves arranged spiralIy, jin-paripinnately compound with 17-25 leaflets;stipules small, caducous; petiole and rachisdensely bristly hairy; petiolules c. I mm long;leaflets alternate, obovate to elliptical or ob-long, 5-17(-20) min x 2.5-8(-11) min, leathery,yellowish hairy below. Inflorescence an axillarypanicle 1.5-4 cm long, hairy. Flowers bisexual,papilionaceous, 4-5 mm long; pedice1 0.5-1.5min long; calyx campanulate, 2.5-3 min long,lobes shorter than tube, lower lobe longest,upper lobes fused; corolla whitish, with obovateto violin-shaped standard and clawed wingsand keel; stamens 10, fused into a tube, butfree in upper part; ovary superlor, with distinctstipe at base, style short. Fruit a flat, obovateto oblong pod 1.5-7 cm x I-2 cm, with stipe I-2 mm long, glabrous, pale brown, indistinctly

L,

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I

2

Madagascar, which also resembles Do16ergioboronii, but differs in its leaves with fewer andslightly larger leaflets. It is classified as en-dangered in the IUCN Red list

Growth and development The roots formnodules with nitrogen-fixing bacteria; Ago-rhizobium, Bradyrhizobium and Mesoyhizo-binm strains have been isolated from the rootnodules.

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 prn); 27:intervessel pits large (Z 10 prn); 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 F1m; (45:vessels of two distinct diameter classes, woodnot ring-porous); 46: :S 5 vessels per squareminimetre; 58: gums and other deposTts inheartwood vessels. Tracheids and fibres: 61:fibres with simple to minutely bordered pits;66: non-septate fibres present; 69: fibres thin-to thick-walled. Axial parenchyma: 80: axialparenchyma anform; 82: axial parenchymawinged-allform; 85: axial parenchyma bandsmore than three cells wide; 86: axial paren-chyma in narrow bands or lines up to threecells wide; (89: axial parenchyma in marginalor in seemingly marginal bands); 90: fusiformparenchyma cells; 91: two cells per paren-chyma strand. Rays: (96: rays exclusivelyuniseriate); (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 min. Storiedstructure: 1/8: all rays stoned; 120: axial pa-renchyma and/or vessel elements stoned. Min-eralinclusions: 136: prismatic crystals present;142: prismatic crystals in chambered axial pa-Tenchyma cells.(P. Derienne & P. E. Gasson)

Ecology Do16ergio boroniioccurs in lowlandevergreen, humid rainforest, from sea-level upto 150(-600) in altitude. It is often found alongwatercourses, but also in swamp forest and onthe land margin of mangrove vegetation. Itusually occurs on sandy soils, which are some-times saline, rarely on ferrallitic soils at higheraltitudes

Genetic resources Although Do16ergi0 60-

:.* . .,\

A ' \>, '

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*

Daibergio boronii- I, flowering brunch, ' 2, fruit-ing byoneh, 3, seedRedrawn Grid odopted by Ishoh Syomsudin

veined, indehiscent, I-3-seeded. Seeds kidney-shaped, c. 9 min x 5 min, dark red-brown

Other botanical information Daibergio isa large pantropicalgenus comprising about 250species. Tropical Asia and tropical Americahave about 70 species each, continental Africaabout 50 and Madagascar slightly over 40. InMadagascar many Do16ergio species producehigh-quahty wood. Some of them resembleDo16ergio boronii. DoIbergio doDidit Bosser &R. Rabev. is similar in its leaflets, but differs inits glabrous rachis and inflorescence structureThis species is only known from a single areain western Madagascar and is classlfled asendangered in the IDCN Red list of threatenedspecies; it is selectively felled for Its valuabletimber. Do!bergio pseudoburonit R. Vig. resem-bles Do16ergio boronii in its leaves and flow-ers, but differs in its larger I'Seeded fruits.This species, classified as vulnerable in theIDCN Red list, is restricted to northern Mada-gascar. It is selectively felled for its high-quality timber. This is also the case for Do16er-gio tsarotononensis Bosser & R. Rabev. , a spe-cies from the Tsaratanana Massif in northern

C>

DALEERGIA 191

3

.

192 TIMBERSl

ronii is fairly widely distributed in lowlandrainforest along the east coast of Madagascar,its habitat has been much reduced. Moreover,it is selectively felled and large trees of Do16er-gio boronii have become rare. It is included inthe IUCN Red list of threatened species, inwhich it is classified as vulnerable. Studies onthe genetic vanability of Do16ergio boronit areongoing.

Prospects Do!be78io boroniiis overexploited,and will soon disappear from the timber mar-ket as stands have largely been depleted. Pro-tection of remaining stands is badly needed,and Do!bergio boronii will only have a role ascommercial timber in the future if plantationsbecome successful, or if the timber is sustaina-bly harvested from natural forest. This willprobably allow only very low yield levels be-cause trees presumably grow slowly. ResearchInto propagation techniques and proper man-agement methods seems worthwhile in thelight of the excellent properties of the wood.

Major references Bolza & Keating, 1972;CTFT, 1962f; du Puy at a1. , 2002; Takahashi,1978

Other references Boiteau, Boiteau & A1-10rge-Bolteau, 1999; du Puy, 1998a; Gu6neau,1971; Gu6neau, Bedel & Thie1, 1970-1975;National Academy of Sciences, 1979; Nelson,1993; Rasolomampianina at a1. , 2005.

Sources of illustration du Puy at a1. , 2002.Authors R. H. M. J. Lemmens

wood is traded in small amounts in local and

international markets, often mixed with thewood of other Daibergio spp.

Properties The heartwood is red to purplishgrey with darker streaks, distinctly demar-cated from the reddish grey sapwood. The tex-ture is medium. The wood is moderately heavy,slightly lighter in weight than the wood of DoI-bergio boronii Baker, hard and elastic. Shrink-age during drying is moderate to high. Thewood is suitable for sliced veneer. It is moder-

ately durable.The presence of navonoids, tannins, triter-penes, steroids, and arithra-coumarins

cenosides has been recorded for Do!bergiochope!ieri

Botany Deciduousshrub or smallto medium-sized tree up to 15(-18) in tall; hole up to 60 cmin diameter; outer bark thin, with lenticels,whitish to greyish, inner bark brownish; youngbranches glabrous. Leaves arranged spiralIy,imparipinnately compound with (7-)11-15(-19) leaflets; stipules small, caducous; petioleand rachis usually glabrous; petiolules 1.5-4min long; leaflets alternate, obovate to ellipti-cal or oblong-elliptical, 2-6 cm x I-3 cm, Ieath-ery, glabrous. Inflorescence a terminal or axil-Iary panicle 5-10 cm long, short-hairy. Flowersbisexual, papilionaceous, 8-12 mm long; pedi-ce1 1.5-4(-5) min long; calyx campanulate, 6-9min long, reddish to purple, lobes longer thantube, upper lobes fused; corolla whitish, oftenreddish tinged, with broadly elliptical standardand clawed wings and keel; stamens 10, fusedinto a tube, but free in upper part; ovary supe-nor, with distinct stipe at base, style slender, c.2 mm long. Fruit a flat, elliptical to oblong pod5-8(-13) cm x 1.5-3 cm, with stipe 3-5 mmlong, glabrous, reddish brown, finely reticu-lately veined, indehiscent, I-3-seededDo16ergio chopelieritrees flower when they areleafless or forming new foliage, from August toApril. Nitrogen-fixing root nodules have beenreported for Daibergio chopelieri; Brodyrhizo-bium and Mesorhizobium strains have beenisolated from the root nodules

Do16ergio is a large pantropical genus coinpris-ing about 250 species. Tropical Asia and tropi-cal America have about 70 species each, conti-nental Africa about 50 and Madagascarslightly over 40. In Madagascar many Do16er-gio species produce high-quality woodDo!bergio peruillei Vatke from western Mada-gascar is very similar to Do16ergio chopelieri,but it can be distinguished by its usually short-hairy petiole and rachis and slightly smaller

DALBERGIACHAPELIERiBaill.

Protologue Bull. Mens. SOC. Linn. Paris I:436 (1884)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Synonyms Daibergio prerocurpi/lord Baker(1890)

Vernacular names Hazovola a grandesfeuilles (ET).

Origin and geographic distribution Daiber-gio chope!tort is endemic to eastern Madagas-

Uses The wood is used for construction, car-pentry and furniture. In the past, it has beenused for railway sleepers. It is also used intraditional medicine to treat parasitic diseasesincluding bilharzia, intestinal complaints in-cluding diarrhoea and dysentery, and to facili-tate childbirth. The bark is sometimes collected

and used for tanning hides and dyeing.Production and international trade The

car

fruits. The wood of Daibergto perui!leiis alsoused for carpentry and furniture, and the red-dish exudate is used in local medicine to treat

laryngitis. Do16ergio peruillei is classified inthe lower risk category in the IUCN Red listDaibergio tricolor Drake is related to Daibergioperuillei; it is selectively felled for its timber,and its bark yields a black dye and its wood ared one. It is classified as vulnerable in theIUCN Red list. Daibergio tsiondolono R. Vig. isanother somewhat SImlIar species from west-ern Madagascar; it can be distinguished by itsmore numerous and smaller leaflets. Althoughit is a small and rare tree, it is reportedIy se-Iectively felled for its good-quality rosewood,although confusion with other species may bethe basis of this statement. Do16ergio tsion-dojono is classified as endangered in the IUCNRed list. Do16ergio globerrimo Bosser &R. Rabev. also resembles Do!bergio chopelieri,especially in its flowers, but differs in fewerleaflets per leaf and its inflorescence being araceme. It is classified as vulnerable in theIUCN Red list. The wood of Do16ergio glober-rimo is used in western Madagascar for con-struction and furniture.

Ecology Daibergio chopelieri occurs in ever-green humid forest, up to 1000 in altitude. Itcan be found in humid valleys as well as ondrier crests, and even may survive as a shrubafter resprouting in secondary vegetation.

Genetic resources and breeding AlthoughDo16ergio chopelieriis widespread in easternMadagascar, from Maroantsetra in the north toT61atiaro in the south, it occurs mainly in low-land forest, a habitat that is under much pres-sure because of growing human populationsand the resulting demand for agricultural landMoreover, Do16ergio chapelieriis locally selec-tively felled for its valued timber. It is includedin the IUCN Red list as vulnerable.

Prospects Very little information is availableon Do16ergio chopeliert, and much research isstill needed to judge its prospects. However,these do not seem bright because of the oftensmall size of the tree and its declining num-hers

Major references Boiteau, Bolteau & A1-10rge-Boiteau, 1999; du Puy at a1. , 2002; Gu6-neau, Bedel & Thie1, 1970-1975; Rasoanirina,2002; Razafindrasoa, 2003

Other references du Puy, 1998b; Rasolo-mainpianina at a1. , 2005.

Authors R. H. M. J. Lemmens

DAI, BERGIACHLOROCARPAR. Vig.

Protologue Notul. Syst. (Paris) 14(3): 183(1951)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution Do16er-gio chlorocorpo is endemLc to western Mada-gascar

Uses The wood is used for construction andcabinet making, and also as fuelwood.

Production and international trade Thewood is traded in small amounts in local andinternational markets, often mixed with thewood of other Do!bergio spp.

Botany Deciduous small to medlurn-sizedtree up to 15(-20) in tall; bark greyish, longitu-dinally fissured; young branches short-hairyLeaves arranged spiralIy, Tinparipinnatelycompound with (20-)30-35(-39) leaflets; stip-ules small, caducous; petiole and rachis hairy;petiolules 05-1.5 mm long; leaflets alternate,ovate to elliptical or oblong, 14 cm x 0.5-I cm,thinly leathery, hairy below. Inflorescence aterminal or axillary panicle 10-30 cm long,with short and slightly coiled final divisions,hairy. Flowers bisexual, papilionaceous, 5-6mm long, almost sesslIe; calyx campanulate, 3-4.5 min long, violet-brown, lobes longer thantube, lower lobe slightly longer, upper lobesfree; corolla whitish, with broadly obovate toviolin-shaped standard and clawed wings andkeel; stamens 10, fused into a tube, but free inupper part; ovary superior, with distinct stipeat base, style c. I mm long. Fruit a flat, ellipti-cal to ovate pod 5-11.5 cm x 2-3.5 cm, withstipe 7-12 min long, glabrous, pale brown, re-ticulately veined, indehiscent, I-2-seeded.Seeds kidney-shaped, reddish brown.Daibergio chlorocorpo trees flower from Marchto June. The trees were found to be prolific seedbearers with abundant natural regenerationDoIbergiu is a large pantropical genus coinpris-ing about 250 species. Tropical Asia and tropi-cal America have about 70 species each, conti-nental Africa about 50 and Madagascarslightly over 40. In Madagascar many Daiber-gio species produce high-quality wood. Severalspecies more or less resembling Do!bergiochiorocorpo also have recorded timber usesDaibergio gloucocorpo Bosser & R. Rabev. fromnorth-western Madagascar has been confusedwith DoIbergio chiorocorpo, but differs in itslarger flowers, calyx characteristics and fruitslacking the reticulate venation; it is selectivelyfelled for its timber that is used locally. It is

DAI, BERGiA 193

194 TIMBERSl

classified as endangered in the IUCN Red list.Do16ergi0 o6rohomii Bosser & R. Rabev. fromnorthern Madagascar differs from Do16ergiochiorocorpo in having fewer leaflets per leafand longitudinalIy veined fruits. Although it isclassified as an endangered species in theIUCN Red list, it is selectively felled for itsreddish and nicely streaked wood. Do16ergiourschii Bosser & R. Rabev. from the same re-gion is also listed as endangered and is selec-tively felled, the wood being used for construc-tion and carpentry. Do16ergio defyhtnensisBosser & R. Rabev. , which is restricted tosouth-eastern Madagascar, is another endan-gered species selectively felled for its timber; itis characterized by few (7-11) leaflets andshort inflorescences. DoIbergio orientolisBosser & R. Rabev. from eastern Madagascarhas similar flowers to Do16ergio defyhinensis,but differs in its leaves and fruits; its wood isused in cabinet making and carpentry. It isclassified as vulnerable in the IUCN Red list

Ecology Do16ergio chiorocorpo occurs scat-tered in deciduous, seasonalIy dry forest andwoodland, up to 400 in altitude, mainly onsandy soils

Genetic resources and breeding AlthoughDo16ergio chiorocorpo is fairly widespread inwestern Madagascar, it is selectively felled forits valued timber. It has been included in theTUGN Red list as vulnerable

Prospects Very little information is avail-able on Do16ergio chiorocorpo, and much re-search Is stillneeded to judge its prospects as atimber tree of future importance. Its decliningnumbers warrant protection of the remainingstands, which are fortunately partly in pro-tected areas

Major references du Puy at a1. , 2002Other references Deleporte, Randrianasolo

& Rakotonirina, 1996; du Puy, 1998cAuthors R. H. M. J. Lemmens

.

Do16ergiogreueono - wild

Uses The wood is one of the so-called rose-

woods ('Madagascar rosewood', 'pansander'),which are much in demand for cabinet making,furniture, marquetry and parquetflooring. It isone of the favoured woods for musical instru-merits, not only because of its beautiful colourand venation, but also because of its clearnessof tone. It is also suitable for interior trim,joinery, ship and boat building, vehicle bodies,poles and piles, precision equipment, carvings,toys and novelties, sporting goods, handles,ladders, turnery, pattern making, veneer andplywood. The wood is used locally in Madagas-car, e. g. for construction and paddles and asfirewood. The Mikea people of south-westernMadagascar rub pieces of branches on stoneswith water to produce a paste which is appliedto the face as a medicine against various ail-merits; Do!bergio greueono is a sacred tree forthese people.

Production and international trade For-merly Do16ergio greueo, IQ timber representedthe bulk of timber exports from western Mada-gascar, and is still exported in limitedamounts. The wood is traded on the interna-tionalmarket, usually in smallamounts and athigh prices, for special applications such asornaments, turnery and musical instruments.In recent years it has replaced Brazilian rose-wood (from Do16ergio itigro ryell. ) Benth. ) be-cause this South-American species has beenincluded in Appendix I of CITES as an endan-gered species. It Is often traded as quarter-sawn pieces of comparatively smalldimensions.

Properties The heartwood is purplish brown,often with darker stripes, and distinctly de-marcated from the lighter sapwood. The grain

DALBERGIAGREVEANA Baill.

..

o

,^

Protologue Bull. Mens. SOC. Linn. Paris I:436 (1884).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Vernacular names French rosewood, Mada-gascar rosewood (En). Panssandre violet, pans-sandre de Madagascar (Fr)

Origin and geographic distribution Do16er-gio greueono is endemic to Madagascar, whereit is widespread in the western part

,.

..

is generally straight, texture fine to moderatelyfine and even.

The wood is heavy, with a density of about1080 kg/ing at 12% moisture content; It Is veryhard. It should be air-dried with great carebecause it is liable to checking and splittingBefore air-drying logs should preferably bequartersawn. Turned pieces used for precisionequipment or musical instruments should bedried thoroughly to avoid distortion. However,the reported rates of shrinkage are moderate,from green to oven dry about 3.3% radial and5.4% tangential. Once dry, the wood is verystable in service

At 12% moisture content, the modulus of rup-ture is 181-226 N/mm2, compression paralleltograin 98 N/min2, cleavage 21.5 N/min, Jankaside hardness 13,350 N and Chalais-Meudonside hardness 18.6

The wood works well, both with hand tools andmachine tools, but it blunts sawteeth rapidly.For nailing and screwing pre-boring is neededPainting and varnishing gives moderate re-SUIts because of the oily surface of the wood,and the gluing properties are moderate. Thewood is suitable for sliced veneer. It is durable,

being resistant to termite and Lyetus attacks,but only moderately resistantto marine borers.The heartwood is very resistant to treatmentwith preservativesDichloromethane and methanolic extracts ofDo!bergio greueono bark showed activityagainst gram-positive bacteria

Adulterations and substitutes The woodof several other DoIbergio species from Mada-gascar is also traded as Madagascar rosewoodor pansander.

Description Deciduous small to medium-sized tree up to 15(-20) in tall; bole often shortand crooked, up to 50 cm in diameter; barkwhitish to blackish grey, smooth to rough;young branches glabrous. Leaves arrangedspiralIy, imparipinnately compound with 7-9(-11) leaflets; stipules small, caducous; petioleand rachis slightly hairy to glabrous; petiolules3-8 min long; leaflets alternate, ovate to errip-tical or almost circular, (2-)2.5-6 cm x (0.5-)1.5-3 cm, thinly leathery, minutely hairy toglabrous below. Inflorescence a terminal or

long, withaxillary panicle 5-15(-20)slightly coiled final divisions, hairy; bracts per-SIStent. Flowers bisexual, papilionaceous, 3-4.5mm long; pedicel c. 0.5(-1.5) mm long; calyxcampanulate, 2-3 mm long, lobes shorter thantube, lower lobe slightly longer, upper lobesfused; corona whitish becoinlrig cream-coloured,

with broadly obovate to violin-shaped standardand clawed wings and keel; stamens 10, fusedinto a tube, but free in upper part; ovary supe-nor, with distinct stipe at base, style shortFruit a flat, elhptical to Thornbic pod 3-6.5 cm xI-2.5 cm, with short stipe 4-5 mm long, yenow-ish brown, indehiscent, usually I-seeded. Seedkidney-shaped, c. 8 mm x 4 mm, reddish brown.

Other botanical information Do16ergio Isa large pantropicalgenus comprising about 250species. Tropical Asia and-tropical Americahave about 70 species each, continental Africaabout 50 and Madagascar slightly over 40. InMadagascar many Do16ergio species producehigh-quality wood. Do16ergio humbertii R. Vig.may resemble Do16ergio greueono, but differsin more numerous leaflets (11-15) and largerfruits. Do16ergio humbertiiis mainly found inthe inkarana Massif in northern Madagascar,and yields a good-quality rosewood. Do16ergiosuoresensis Baill. is also close to Do16ergio gre-ueono, and has the same number of leaflets asDo16ergio humbertii, but sinaUer fruits thanthe latter species. Daibergiu suoresensis re re-stricted to the area around hatsiranana innorthern Madagascar, and the wood is used incabinet making. Do16ergio hi, in bertii and DoI-bergio suoresensis are both classified as en-dangered in the IDCN Redlist

Anatomy Wood-anatomical 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); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to Intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;43: mean tangential diameter of vesselluminaZ 200 prn; 45: vessels of two distinct diameterclasses, wood not ring-porous; 47: 5-20 vesselsper square minimetre; 58: gums and other de-POSits in heartwood vessels. Tracheids andfibres: 61: fibres with simple to inlnutely bor-dered pits; 66: non-septate fibres present. Axialparenchyma: 77: axial parenchyma diffuse-in-aggregates; 80: axial parenchyma allform; 82axial parenchyma winged-allform; 84: axialparenchyma unilateral paratrachea1; (86: axialparenchyma in narrow bands or lines up tothree cells wide); (89: axial parenchyma inmarginal or in seemingly marginal bands); 90fusiform parenchyma cells; 91: two cells per

DanBERGiA 195

cm

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196 TIMBERSl

parenchyma strand. Rays: (96: rays exclusivelyuniseriate); (97: ray width I-3 cells); 104: allray cells procumbent; 1/5: 4-12 rays per min.Stoned structure: (1/8: all rays stoned); (1/9low rays stoned, high rays non-stoned); 120:axial parenchyma and/or vessel elements sto-ried; (122: rays and/or axial elements irregu-Iarly stoned). Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells.(P. 06tienne & P. E. Gasson)

Growth and development Do16ergio gre-ueono flowersfrom October to April

Ecology Do16ergio greueonu occurs in deci-duous, seasonalIy dry forest and woodland,sometimes as a shrub in grassland, up to 800in altitude. It is found on various soils, fromsandy to limestone-derived and ferrallitic.

Propagation and planting Natural regen-eration of Daibergio greueono appears to bepoor in comparison to other Daibergio sppfrom western Madagascar, although in Moron-dava region the trees were found to be prolificseed bearers with abundant natural regenera-tion

Genetic resources Although DoIbergio gre-ueono Is still widely distributed in westernMadagascar, its population has much declinedIt Is selectively felled for its valuable timberand its habitat is under pressure. It is includedin the IUCN Red list of threatened species,where it is classified as a lower-risk species,but close to vulnerable.

Prospects Do!bergio greueono seems to beoverexploited, and may soon disappear fromthe timber market because of stand depletionProtection of remaining stands is needed, andDo16ergio greueono may only have a role astimber of importance in the future if planta-tions are successful, or if the timber is sustain-ably harvested from natural forest. This willprobably allow only very low yield levels be-cause trees presumably grow slowly. However,research into propagation techniques andproper management methods seems worth-while in the light of the excellent properties ofthe wood. The comparatively small size andoften poor shape of the hole is a drawback

Major references Bolza & Keating, 1972;du Puy at a1. , 2002; Sanda, 2004; Takahashi,1978

Other references Bolteau, Bolteau & A1-10rge-Boiteau, 1999; Deleporte, Randrianasolo& Rakotonirina, 1996; du Puy, 1998d; Inside-Wood, undated; Stiles, 1998

Authors R. H. M. J. Lemmens

DAI, ~BERGIAHILDEBRANDTllVatke

Protologue Linnaea 43: 101 (1881)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Origin and geographic distribution Do16er-

gio hitdebrondttiis endemic to northern andwestern Madagascar.

Uses The wood is valued for cabinet making;it is reportedIy a good-quality Madagascarrosewood.

Botany Smalltree up to 10 in tall, occasion-ally larger; young branches glabrescent,brownish to yellowish. Leaves arranged spi-rally, Imparipinnately compound with 7-13leaflets; stipules small, caducous; petiole andrachis glabrous or sometimes slightly hairy;petiolules 2.54.5 mm long; leaflets alternate,ovate to elliptical or oblong, 2-6.5 cm x I-2.5cm, thinly leathery, short-hairy below. Inno-rescence a terminal or axillary panicle up to 30cm long, with slightly coiled final divisions,short-hairy. Flowers bisexual, papilionaceous,3.5-4 mm long; pedice1 0.5-I min long; calyxcampanulate, 2-2.5 mm long, dark purple,lobes about as long as tube, lower lobe slightlylonger, upper lobes fused; corolla whitish, withbroadly obovate to violin-shaped standard andclawed wings and keel; stamens 10, fused intoa tube, but free in upper part; ovary superior,with distinct stipe at base, style short. Fruit aflat, elliptical to oblong pod 2-6 cm x 0.5-I cm,with stipe 3-5 mm long, glabrous, reddishbrown, Teticulately veined, indehiscent, I-3-seeded.

Do16ergio hildebrondtii flowers from March toMay.Do!bergio is a large pantropical genus coinpris-ing about 250 species. Tropical Asia and tropi-cal America have about 70 species each, conti-nental Africa about 50 and Madagascarslightly over 40. In Madagascar many Do16er-gio species produce high-quality wood. Twospecies which are probably related to Daibergiohitdebrondtii also have recorded timber uses.Do16ergio emirnensis Benth. from the southernhalf of Madagascar has similar flowers to DoI-bergio hildebrundtii, indicating an affinity, butdiffers in leaf and fruit characteristics; its woodis used for beams and toolhandles, as firewoodand for charcoal production. Daibergio emir-itensis Is classified in the lower risk category inthe IUCN Red list. Do16ergio neoperrieriBosser & R. Rabev. from western Madagascarresembles Do16ergiu hitdebrondtii, but canalso be distinguished by its leaves and fruits; it

^ I

is selectively felled for its timber. Do16ergioneoperrieri is classified as vulnerable in theIUCN Red list.

Ecology Daibergio hildebrondtii occurs indeciduous, seasonalIy dry forest and woodland,up to 600 in altitude, on sandy, clayey or rockysoils.

Genetic resources and breeding AlthoughDo16ergio hildebrondtiiis fairly widespread, itis uncommon and selectively felled for its tim-her. It has been included in the IUCN Red listas vulnerable

Prospects Very tittle information is availableon DoIbergio hildebrondtii, but it does notseem to have good prospects as a timber tree offuture importance because of its decliningnumbers and often smallsize.

Major references du Puy at a1. , 2002.Other references du Puy, 1998e.Authors R. H. M. J. Lemmens

most popular woods for carving and engravingIt is suitable for turnery and is excellent forhigh-class bentwood furniture, walking sticks,umbrella handles and other bentwood articlesIt is also used for making musical instrumentsand sports equipment.Do16ergio lotifo!to is used as a shade tree inagroforestry in India and Indonesia, for affore-station of eroded soils, and as a soilimproverfixing nitrogen and providing mulch. It is alsoplanted as a roadside tree and shade tree incoffee plantations. The foliage is used as rod-der. Do16ergio lotifolio yields a dark amber,strong flavoured honey. The bark is used intraditional medicine in India, to treat diar-rhoea, indigestion and leprosy, and as a vermi-fuge.

Production and international trade In In-dia Do16ergio lotifolio timber is one of the mostvaluable timbers, with an average price higherthan that of teak. In Java the price is coinpa-Table with that of teak. Plywood is exported.

Properties The heartwood is golden brownto dark purplish brown with very dark brownto black streaks, and distinctly demarcatedfrom the 3-5 cm thick whitish to yellowishsapwood. The grain is straight, sometimeswavy or interlocked, texture moderately fine torather coarse. The density of the wood is 750-880 kg/ina at 12% moisture content. Air dryingshould be done carefully and slowly becausethe wood splits easily at the ends during dry-ing. The rates of shrinkage from green to ovendry are 2.3-2.9% radial and 5.6-6.4% tangen-tial. Once dry, the wood is exceptionally stableIn service.

At 12% moisture content, the modulus of rup-ture is 92-121 N/mm2, modulus of elasticity10,000-11,900 N/mm2, compression parallel tograin 48-65 N/min2, shear 8-9 N/mm2, cleav-age 85 N/mm radial and 91 N/min tangential,Janka side hardness 6970 N and Janka endhardness 8015 NThe wood is rather difficult to work with handtools, but it is quite easy to machine. It can beplaned to a smooth surface. Turning, screwing,polishing and gluing give good results, and thewood can be peeled or sliced to make decorativeveneer and plywood. The heartwood is durable,being resistant to dry-wood termites and wood-rotting fungi; it is difficult to treat with pre-servatives. The sapwood is perishable but read-11y treatable. The wood can cause allergic con-tact dermatitis in people working with it

Botany Deciduous or evergreen medium-sizedto large tree up to 40 in tall; hole straight or

DALBERGIALATIFOLIARoxb

Protologue PI. Coromande12: 7, t. 1/3 (1799)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Chromosome number2n= 20

Vernacular names Indian rosewood, EastIndian rosewood, Bombay blackwood (En). Pal-issandre de 1'1nde, panssandre d'Asie (Fr).

Origin and geographic distribution Do16er-gio lotifolio is native to tropical Asia, fromNepal to India and in Java (Indonesia). It isplanted in tropical Asia, and locally also intropical Africa, e. g. in Nigeria, Kenya, Ugandaand Tanzania, and also in Reunion and Mauri-tms. However, in tropical Africa it is onlyplanted on a small scale and usually as an or-namental plant, especially in botanical gar-dens

Uses The wood of Do16ergio lotifolio fromtropical Asia is well known for its applicationsin high-class furniture, cabinet making and asa decorative wood used, for example, in pas-senger ships and for instrument cases. It issuitable for high-grade plywood and, owing toits beautiful colour and figure, for decorativeveneer. Because of its strength and durability,it is suitable for all kinds of constructionalwork, doors, window frames and wagon build-ing. It is also used for handles of heavy-dutyhammers and axes and for agricultural imple-merits such as ploughs, harrows and rollers. Incart and carriage building, it is used for wheelrims, spokes, poles and shafts. It is one of the

DALBERGIA 197

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taproot and are practically devoid of any sec-ondary roots when young. Initial growth ofseedlings is slow. Nodules which are the resultof symbiosis with nitrogen-fixing bacteria arealready found on the roots of seedlings. Youngtrees are also relatively slow growing, but re-ported growth rates differ considerably. InJava (Indonesia) an annual height growth of 2in and an annual volumeincrementof 15 mama

have been recorded for young plantations onfavourable sites, but in India 10-year-oldstands had an average height of 6 in with ahole diameter of 4-5 cm. In India the averageage of reaching a diameter of 60 cm has beenestimated at no less than 240 years! Do16ergiolotifolio trees are often surrounded by numer-ous root suckers. Do16ergio lotifolio is a inod-erate light demander and seedlings can with-stand moderate shade. In too open locations ittends to become crooked and branchy. In Nige-ria trees flower in September-October andJanuary-February. The thin pods are dis-persed by windDo!bergiu is a large pantropical genus coinpris-ing about 250 species. Tropical Asia and tropi-cal America have about 70 species each, conti-nental Africa about 50 and Madagascarslightly over 40

Ecology In its native area of distribution intropical Asia Do16ergio lotifoliu occurs scat-tered in deciduous forest in periodically verydry localities, up to 900(-1500) in altitude. It issuccessfully cultivated as high as 1000 in alti-tude. Older trees are very drought resistantDaibergio lotifolio thrives in areas with up to 6dry months with mean monthly rainfall of lessthan 40 min. It tolerates minimum tempera-tures of 0-6'C. It is only moderately fire resis-tant. It grows well on deep, permanently moistbut well-drained soils, but also attains largedimensions on vertisols

Management The weight of 1000 seeds is25-55 g. Seeds remain viable for up to 6

dormancy. Pre-months. They have nO

treatment of seeds is not necessary, althoughsoaking in water for 12-24 hours acceleratesgermination. Germination of fresh seed takes7-25 days, and the germination rate is 45-80%. Under natural conditions regeneration isgenerally satisfactory, with seeds germinatingat the beginning of the rainy season. However,seedlings should be protected against fire andgrazing

For planting, seeds are sown in well-raisedseedbeds of porous sandy loam or in polythenebags. However, Do16ergio lotifo!to is often

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Do16ergio lotifolio - I, tree hubit, 2, floweringtwig, . 3, leaflet, 4, fruit.Source. . PROSEA

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slightly twisted, branchless for up to 12(-24)in, up to 80(-150) cm in diameter, often withprominent buttresses; bark surface whitish togrey, thin, becoming flaking; crown rounded todome-shaped. Leaves arranged spiralIy, jin-panpinnately compound with (3-)5-7(-9) leaf-lets; stipules small, caducous; petiole and ra-chis glabrous; petiolules up to I cm long; leaf-lets alternate, broadly obovate to elliptical-oblong, 4-12 cm x 2.5-9 cm, obtuse, rounded ornotched at apex, papery or thinly leathery,glabrous. Inflorescence a terminal or axillarypanicle 5-15 cm long, laxly branched, almostglabrous, many-flowered. Flowers bisexual,papilionaceous, 6-8 mm long, distinctly pedi-cellate; calyx campanulate, c. 4 mm long, lobesshorter than tube, lower lobe longest, upperlobes fused; corolla whitish, with obovate stan.dard and clawed wings and keel; stamens usu-ally 9, fused into a tube, but free in upper part;ovary superior, with distinct stipe at base, styleshort. Fruit a flat, elliptical to oblong, paperypod 4-10 cm x 1.5-2.5 cm, with stipe up to Icm long, glabrous, reticulately veined, indehis-cent, I-3(-4)-seeded. Seeds kidney-shaped, 7-10 min longSeedlings of Do!bergio lotifolio have a strong

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propagated from root suckers of I-2.5 cm di-ameter. Root and stem cuttings can also beused. The buds of root suckers and stem cut-

tings start to sprout about 9 days after plant-ing, and those of root cuttings about 15 daysafter planting, but after 2 months all youngplants have more or less the same heightStump planting is very successful uslrig

stumps from 2-3-year-old seedlings with a col-Iar diameter of 5-15 min, a rootlength of about15 cm and a shoot length of about 5 cm. In In-dia successful methods of tissue culture havebeen developed.Do16ergio lotifolio is generally grown in purestands, usually managed by clear felling fol-lowed by artificial regeneration, but sometimesit is mixed with other timber species such asmahogany (Swigtertio sp. ). In agroforestry sys-terns it is intercropped with annual crops orfruit trees. In pure stands, spacing is 1.2-2.5 inx 1.0-1.8 in, but in India an initial spacing of 5in x 5 in, followed by thinning of alternatetrees, is recommended. Regular weeding isnecessary for several years. Pruning and thin-rimg are recommended at 5-10 years afterplanting. Trees can be coppiced and pollardedIn Java FMSorium soloni has caused wide-

spread damage to plantations over 15 yearsold. The symptoms are Inward rolling of youngleaves, dieback and discoloration of otherleaves, and red streaks formed on outer layersof the sapwood. Root suckers of affected treesshould not be used for propagation. Seedlingsoften suffer seriously from damping-off; themortality rate may be up to 60%. Up to 12years old, Daibergio lotifolio is susceptible toPhytophthoro attack. The trees are attacked byvarious insects such as leaf-miners, defoliatorsand stem-borers, but this causes no real prob-Iems for trees grown under favourable condi-tions

Genetic resources and breeding In tropi-cal Asia natural stands of Do16ergio lotifoliohave been depleted considerably, and in manyareas large trees have become rare. The speciesis included in the IUCN Red list as vulnerable.

A germplasm bank has been established at theKerala Forest Research Institute (India)

Prospects Do16ergio lotifolio is suitable forincorporation in agroforestry systems, but toobtain straight boles close spacing is desirable,and for timber production the establishment ofmonoculture plantations is recommended. A1-though the timber is much in demand and ex-pensive, Do16ergio lotifolio is too slow-growingto be a preferred timber plantation species for

tropical Africa at present. Breeding pro-grammes aimed at fast-growing trees withstraight boles may result in profitable planta-tions in the future, but for the time being itseems more rational to focus on proper man-agement methods of the African Do16ergio spe-cies producing valuable timber. An advantageof Daibergio lotifo!to over most DoIbergio sppindigenous to Africa is the larger bale size ofmature trees

Major references CAB International, 2005;Prawirohatmodjo at a1. , 1993; Rainesh Rao &Purkayastha, 1972; Tewari, 1995; World Agro-forestry Centre, undated

Other references Asian Regional Work-shop, 1998b; Athavale at a1. , 2003; Burkill,1995; CSIR, 1952; Farmer, 1972; Keay, 1989;National Academy of Sciences, 1979; Filetta ata1. , 1996; Polhi11, 1990; Ravishankar Rai &Jagadish Chandra, 1988

Sources of illustration Prawirohatmodjoat a1. , 1993.

Authors R. H. M. J. Lemmens

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

DAI. BERGIA 199

DALBERGIALOIAJELllR. Vig.

Protologue Notul. Syst. (Paris) 14(3): 184(1951)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Vernacular names Volombodipona derandesreuiUes (Fr)

Origin and geographic distribution Do16er-gio IOUueliiis endemic to eastern Madagascar,where it occurs from Maroantsetra in the north

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to Mariakara in the south.

Uses The wood of Do16ergio IOUuelitis one ofthe so-called rosewoods ('Madagascar rosewood','pansander') much in demand for cabinet inak-Ing, furniture, marquetry and parquet flooringIt is one of the favoured woods for musical in-

struments. In Madagascar it is in high demandfor carving and turning, and it has been usedtraditionally for tombs. The heartwood is usedin traditional medicine to treat bilharzia andmalaria.

Production and international trade Mada-

gascar rosewood is still traded on the interna-tional market, usually in small volumes and athigh prices, for special applications such asmusical instruments. In recent years it hasreplaced Brazilian rosewood (from Do16ergioitigro ryell. ) Benth. ) because this South-American species has been included in Appen-dix I of CITES. It is often traded as quarter-sawn pieces of comparatively small dimen-SIons. However, larger trees of Daibergio IOU-Dent have become so rare that the share of this

species in the total export is probably verysmallor practically zero

Properties The heartwood is purplish red,becoming purplish black upon drying. The tex-ture is fine and even, and the wood has a beau-tifulpolish. It is very heavy and very hard. Thewood has a density of 800-900 kg/ina at 12%moisture content. The rates of shrinkage arerather low, from green to oven dry 3.4-4.6%radial and 5.8-7.6% tangential.Several navonoids have been isolated from theheartwood, some of which showed antiplasmo-dial actrvity in vitro

Adulterations and substitutes The wood

of several other Do16ergio species from Mada-gascar is traded as Madagascar rosewood orpansander

Description Deciduous medium-sized treeup to 20 in tall; bark greyish; young branchesshort-hairy. Leaves arranged spiralIy, imparip-InnateIy compound with 9-15 leaflets; stipulessmall, caducous; petiole and rachis bristlyhairy but glabrescent; petiolules I-2.5 minlong; leaflets alternate, ovate to elliptical, (I-)2-4 cm x 0.5-1.5 cm, thinly leathery, usuallyhairy on both surfaces. Inflorescence an axil-Iary raceme shorter than the leaves, hairyFlowers bisexual, papilionaceous, 12-15(-18)mm long; pedice1 2-10 min long, jointed; calyxcampanulate, 7-10 mm long, lobes about aslong as tube, lower lobe longer, upper lobesfused; corolla whitish, with almost circularstandard and clawed wings and keel; stamens

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Do16ergio IOUuelii - flowering und fruitingbrunch

Redrown und udopted by Achingd Sotiri Nur-human

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10, fused into a tube, but free in upper part;ovary superior, with distinct stipe at base, styleslender, 2.5-3.5 min long. Fruit a flat, oblong-elliptical pod 48 cm x c. 1.5 cm, with slenderstipe I-1.5 cm long, reddish brown, slightlyTeticulately veined, indehiscent, I-2-seeded.Seeds kidney-shaped, c. 13 min x 6 min, red-dish brown.

Other botanical information Do16ergio isa large pantropicalgenus comprising about 250species. Tropical Asia and tropical Americahave about 70 species each, continental Africaabout 50 and Madagascar slightly over 40. InMadagascar many Daibergio species producehigh-quality wood. Do16ergio IOUuelit is distin-guished by its comparatively large flowers.Daibergio normondiiBosser & R. Rabev. , a rarespecies from eastern Madagascar, has similarflowers, but fewer leaflets per leaf; it provides aMadagascar rosewood of excellent quality,which is used in cabinet making. It is classifiedas endangered in the IDCN Red list. The flow-ers of Do16ergio moritimo R. Vig. also resemblethose of Do16ergio IOUue!it, but they are smaller