is likely to remain In demand on internationaland local markets, but it is available in smallquantities only. It is unlikely that this situa-tion will change. The pharmacological proper-ties deserve research attention

Major references Adjanohoun & Ak6 Assi,1979; Bolza & Keating, 1972; Burki11, 1994;Dudek, F6rster & missenbauer, 1981; Inc.1991; Keay, 1958d; Leonard, 1956; Neuwinger,2000; Savill & Fox, 1967; Voorhoeve, 1979

Other references Aubr6ville, 1959a; Betti,2002; Bouquet & Debray, 1974; Chilima &Namot0, 2008; Chilufya & Tengn6s, 1996; de inMensbruge, 1966; Dery, Otsyina & Ng'atIgwa(Editors), 1999; FAO, 1985; Hubert, undated;Jansen, 1974; Kryn & Fobes, 1959; Lovett ata1. , 2007; Meniaud & Bretonnet, 1926; Nor-mand, 1955; Normand & Paquis, 1976; Rad-cliffe-Smith, 1987b; Ruffo, Birnie & Tengn6. s,2002; Tailfer, 1989; Torelli, Pigkur & Ti^Ier,2003; Vivien & Faure, 1985.

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

Authors L. FA. Oyen

Comoros and Madagascar, and on SOCotra is-land oremen).

Uses The heartwood, often traded as 'iron-wood', is in high demand for flooring, carpentryand panelling, and is widely used for house andbridge construction, counter and table tops,railway sleepers, toolhandles and wagon partsIt produces beautiful furniture, turnery andsliced veneer, and is often used by African art-1sts. It is suitable for Interior trim, sportinggoods, toys, novelties and agricultural imple-merits. In South Africa it has been used tradi-tionally to make assegais. It is also used asfirewood and for charcoal production.The oily fruits are edible and are used insouthern Africa in the preparation of beer andlemonade. In East Africa bark decoctions are

used as an emetic and arithelmintic, and totreat malaria, venereal diseases and femalesterility; bark ash is applied as a dressing towounds. In Swaziland bark decoctions are tak-en to treat peptic ulcers, and in South Africathe bark is used for skin lightening. In south-ern Africa root powder is apphed to fracturesand joint swellings, and leaf infusions to treatinfections of the respiratory tract and painsThe foliage serves as fodder, especially duringthe dry season. The flowers produce nectar forhoney bees. In South Africa 01eo cope, "sis hasbeen used as stock for grafting olive cultivarsfrom the Mediterranean region. The tree isconsidered sacred by the Maasai people and iscommonly used in ceremonies.

Production and international trade Theinternational trade in 01eo copensis wood isvery limited. The volumes sold by auction inSouth Africa in the period 2000-2008 variedfrom 50 in3 to 300 ing/year. In 2008 the price offirst-quality boards of 2.5 cm and 5 cm thickwas Us$ 1225/ina and Us$ 1335/in3 respective-Iy, and in 2009 the price of 2.5 cm thick and 10cm thick boards was Us$ 21/01m3 and Us$2160/ina respectively

Properties The heartwood is pale brown todark brown, often with irregular grey-black oryellowish streaks. It is distinctly demarcatedfrom the yellowish white to grey, 2.5-5 cm widesapwood. The grain is straight to slightly inter-locked, texture fine and even. The wood surfaceshows a nice figure and is slightly oily to thetouch

The wood is heavy, with a density of 860-975(-1170) kg/in3 at 12% moisture content, and hard.It air dries very slowly with a marked tendencyto surface checking, splitting, warping anddistortion. Kiln drying is difficult and should

OLEACAPENSIS L.

Protologue Sp. PI. L 8 (1753)Family OleaceaeVernacular names East African olive,

ironwood olive, Elgon olive, African ironwood,black ironwood (En). 011vier du Cap (Fr).Mushargi, 10/10ndo (Sw)

Origin and geographic distribution 01eocopensis is widespread from Guinea and SierraLeone east to Ethiopia and Kenya, and south toSouth Africa and Swaziland. It also occurs in

OLEA 501

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be done at low temperatures. The shrinkagerates are moderate, from green to oven dry 4.0-4.7% radial and 6.7-7.9% tangential. The driedwood is unstable in service unless properlydried. At 12% moisture content, the modulus ofrupture is (78-)127-174 N/mm2, modulus ofelasticity (8600-)16,500-19,500 N/min2, coin.pression parallel to grain (41-)73-84 N/mm2,compression perpendicular to grain 19 N/min2,shear (16-)22.5-26 N/mm2, cleavage 28-94N/min, Janka side hardness 10,050-13,750 Nand Janka endhardness 9780-14,200 NThe wood Is easy to saw when green, but diffi-cult to saw and work when dried, and bluntssaw teeth and cutting edges rapidly. In work-ing the material should be held firmly. Quar-ter-sawn surfaces have a tendency to pick up inplaning, and a cuttlrig angle of 20' is recoin-mended. The wood finishes with a nice polishwithout the use of a filler. Pre-boring is neces-sary in nailing and screwing. The wood doesnot always glue well with conventional woodglues because of the oily surfaces. It turns fair-Iy well and good-quality veneer has been ob-tained with slicing. The steam bending proper-ties are satisfactory. The wood is moderatelydurable to durable. It is sometimes attacked byambrosia beetles and termites, but is not sus-ceptible to Lyetus beetles. The heartwood issomewhat resistant to Impregnation with pre-servatives, the sapwood Is permeableCoumarin and secoiridoid glucosides have beenisolated from the bark. The Iignans (-)-o11viland (+)-cyc10-o11vilhave also been isolated fromthe bark

Adulterations and substitutes The wood

of wild African olive (01eo europoeo L. subsp.cuspidoto myall. ex G. Don) Cif. ) closely resem-bles that of 01eo copensis and is used for SImi-Iar purposes.

Description Evergreen shrub or small tofairly large tree up to 35(-40) in tall; holebranchless for up to 15 in, straight and cylin-drical but sometimes irregular or fluted, up to90(-150) cm in diameter; bark surface of youngtrees smooth and pale grey, in old trees becoin-ing longitudinalIy fissured and dark grey, in-ner bark thick, hard, cream-coloured to orange-brown or greenish, exuding a blackish gum;crown small and dense with steeply ascendingbranches, or rounded with spreading branches;twigs rounded, glabrous, with scattered whitelenticels. Leaves decussately opposite, simple;stipules absent; petiole 0.5-I(-2) cm long;blade elliptical to oblong-elliptical or ovate-elliptical, 3-Ile16) cm x 1.5-5(-6.5) cm, curie-

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01eo copensis - I, tree hobit, 2, legj:' 3, flower-ing twig, ' 4, fruiting twig.Redrawn grid oddpted by Achingd Sotiri Nur-hornon

ate at base, obtuse to acute or short-acuminateat apex, entire to slightly wavy at margins,leathery, glabrous, glossy green above, palegreen below, pinnately veined with 5-7 pairs ofInconspicuous lateral veins. Inflorescence aterminal panicle 3-8 cm long, glabrous, many-flowered. Flowers bisexual, regular, 4-merous,fragrant; pedicel short; calyx cup-shaped, c. Imm long, with triangularlobes; corolla 2-3 mmlong, white, with short tube and elliptical lobes;stamens 2, inserted on corolla tube, c. 2 minlong; ovary superior, flask-shaped, c. 1.5 minlong, 2-celled, style short. Fruit a globose toellipsoid drupe 0.5-2 cm x 05-I cm, purple-black when ripe; stone with thick and woodywall, usually I-seeded. Seed with copious endo-sperm

Other botanical information 01eo coin-prises 33 species, most of them occurring ineastern and southern Africa and in tropicalAsia.

Three subspecies of 01eo copensis are distin-guished: subsp. copensts restricted to SouthAfrica, subsp. erreruis (Harv. ) I'verd. restrictedto South Africa and Swaziland, and subsp.

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ingcrocorpo (C. H. Wright) I'verd. (synonyms01eo gumeensis Hutch. & Dalzie1, 01eo hoch-stetteri Baker) covering the whole area of dis-tribution of the species. They differ in leafshape and size, and in size of the fruit, subsp.macrocorpo having the largest leaves andfruits. The latter subspecies is often a largertree than the other two, and consequently moreimportant as a timber tree.01eo weiooitschii(Knobl. ) Gilg & Schellenb. hasalso been considered as a subspecies of 01eocopensis, but in most recenttaxonomic pubhca-tions it is regarded again as a distinct species,mainly differing in its usually narrower leaveswith longer petioles. 01eo wei"itschit is wide-spread from Cameroon east to Ethiopia andKenya and south to Zambia, Angola andMozambique, from lowland rainforest to ever-green mountain forest. It is a small to fairlylarge tree up to 35 in tall, with bole up to 100cm in diameter, sometimes with large but-tresses. Its wood resembles that of 01eo copen-sis and is used for similar purposes, e. g. forconstruction, flooring, Joinery, furniture andsliced veneer. However, its density appears tobe slightly lower, 690-820 kg/in3 at 12% inols-ture content, and the wood is less strong butmore stable in service. In Ethiopia the but-tresses are cut to make doors. The wood is alsoused as firewood and for charcoal productionThe bark is applied in traditional medicine forsimilar purposes as that of 01eo copensis.01eo loneeo Lain. is also closely related to 01eocopensis, but differs in its usually narrowerleaves with shorter petioles. It is a shrub orsmall tree up to 6 in tall and occurs in Mada-gascar, Reunion and Mauritius. The wood ('boisde cerF, 'bois d'olive blanc') is used for construc-tion and joinery. Bark decoctions are used totreat infections of the respiratory tract andskin complaints, and leaf infusions as emmen-agogue and aphrodisiac

Anatomy Wood-anatonxical description (IAWAhardwood codes)Growth rings: 2: growth ring boundaries Iridis-tinct or absent. Vessels: 5: wood diffuse-porous;7: vessels in diagonal and/or radial pattern; 10:vessels in radial multiples of 4 or more coin-mon; 13: simple perforation plates; 22: inter-vessel pits alternate; 24: intervessel pits ini-nute (S 4 ILm); 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumina 50-100 pin;49: 40-100 vessels per square minimetre; 58:gums and other deposits in heartwood vessels.

Tracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibrespresent; 70: fibres very thick-walled. Axial pa-renchyma: 78: axial parenchyma scanty para-tracheal; 92: four (3-4) cells per parenchymastrand; (93: eight (5-8) cells per parenchymastrand). Rays: 97: ray width I-3 ceUs; (98:larger rays commonly 4- to 10-senate); 106:body ray cells procumbent with one row of up-right and/or square marginal cells; 1/3: dis-junctive ray parenchyma cell walls present;1/5: 4-12 rays per mm. Mineral inclusions(150: acicular crystals).(E. Uetimane, P. Baas & H. Beeckman)

Growth and development 01eo copensisischaracterized as a shade tolerant species. Ini-tial growth of seedlings may be up to 1.1in/year, but after about 4 years the tree growsslowly. In Tanzania, 31-year-old trees were 12-15 in tallwith a bole diameter of 12.5-14.5 cmIn southern Africa trees flower periodically, atintervals of 2-4(-7) years. Fruits ripen 0.5-Iyear later. They are eaten by birds such asbombills and doves, which may disperse thestones. Stones that have past the digestivesystem of birds germinate after 4-6 monthsStudies of faecal samples of chimpanzees inRwanda showed that these often contained01eo copensis stones. Monkeys, wild pigs, fruitbats and squirrels have also been recorded tofeed on the fruits. Investigation of the rootsrevealed the presence ofvesicular mycorrhizae.

Ecology 01eo copensis is characteristic ofhumid mountain forest. In West Africa it isrestricted to mountains above 600 in altitude,in Central Africa it occurs at 1000-2800 inaltitude, and in East Africa it is mainly foundat 1500-2700(-3200) in altitude. In South Mri-ca 01eo copensis is locally a dominant canopytree, especially in forests along the coast. InEast Africa 01eo copensis can dominate theclimax vegetation in lower and inid-altitudemontane forest, together with 01eo europoeo Lsubsp. cuspidoto (Wall. ex G. Don) Cif. and Po-docorpus lotifolius (Thunb. ) R. Br. ex Mirb. InEthiopia it is often found together with Juni-perus procero Hochst. ex Briar. and Afrocorpus101cutus (Thunb. ) GNPageIn general, 01eo copensis prefers regions withan annual rainfallof800-1500 min and a meanannual temperature of 14-18'0. Young treesprefer well-drained, deep, loamy and fertilesoils. Once established, they are quite droughtresistant and they also grow in poor soils

Propagation and planting Natural regen-eration in mountain forest in south-eastern

OLEA 503

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Ethiopia was reported to be poor, but in Tan-zania it was reported to be abundant. In SouthAfrica seedlings are often abundant in the for-est, but many die off as a result of diseasessuch as damping-offFor sowing, the use of fresh stones is recoin-mended. One kg contains 1500-3300 stonesThe germination rate is about 35% and germi-nation takes 2-9 months, but sometimes up to2 years. Whole fruits or stones can be collectedfrom the ground. Stones can be stored for up to3 months, butthey should be cleaned from pulpby rubbing in running water and subsequentlydried for about 5 days. At a low temperature of3'C they can be stored for longer periods. Be-fore sowing the dry stones should be soaked inwater for 2 days. Wildlings are sometimes alsocollected for planting. Planting should be doneduring the wettest part of the year. Seedlingsof about 180 cm tall with lower leaves strippedoff are often used for planting into the field

Management In Tanzania 01eo copensis isgrown in plantations in clusters of up to 10trees at close spacing (I in x I in) with nursetrees such as Greuil!eo robusto ACUnn. Aninterval of 7-8 in is maintained between theclusters. 01eo copensis does not interfere muchwith crops and has been recommended for ag-roforestry systems. The young tree can bemanaged by lopping, pollarding and coppicingIn South Africa trees showed regeneration bycoppices in 40% of cut stems

Diseases and pests 01eo copensis is oftenbrowsed by goats as well as wild animals suchas elephants and antelopes, but it recoverswell

Harvesting In plantations the holes can beharvested for timber about 75 years afterplanting. In forests in southern South Africa,where 01eo copensis is a dominant canopy tree,trees are harvested under a 10-year fellingcycle and selected according to externalIy visi-ble criteria of maturity

Yield In Tanzania well-established stands of01eo copensis had a standing volume of about20 mama 25-30 years after planting

Handling after harvest It is recommendedto remove logs soon from the forest after fellingbecause the sapwood is prone to fungal andinsect attacks. Freshly harvested logs sink inwater and cannot be transported by river.

Genetic resources In many regions Okocopensis is becoming rare due to over-exploitation. This is the case in many moun-tain regions in Ethiopia, Kenya and TanzaniaLocally In South Africa, already by 1890 over-

exploitation of forests where 01eo copensisdominated had made conservation measuresessential. However, in forests in southernSouth Africa, it is stinthe most common cano-py tree. 01eo copensis is legally protected inSouth Africa and harvesting is regulated. Someprovenance testing has been done in Tanzaniawith the aim of selecting the best trees regard-Ing growth performance for larger-scale plant-Ing

Prospects The heavy weight and hardnessof the wood of 01eo copensis are serious draw-backs for many applications. However, thenicely figured wood is attractive for furnitureand sliced veneer. Plantations could producewood with good export prospects, but the rota-tion cycles needed are qulte long. 01eo copensisIs an interesting species for planting in de-graded forests throughout its natural distribu-tion area. Studies to determine the geneticvariation are needed, as well selection of prov.enances with superior hole characteristics fortimber production and investigations on opti-inal SIIvicultural systems. In countries withhigh deforestation rates, immediate action isneeded for in-situ conservation of remainingstands

Major references Bekele-Tesemma, 2007;Bolza & Keating, 1972; Coates Palgrave, 1983;Friis, 1992; Green, 2002; Hines & ECkman,1993; Maundu & Tengnds (Editors), 2005;Palmer & Pitman, 1972-1974; Takahashi,1978; van Wyk & van Wyk, 1997.

Other references Baldoni at a1. , 2002;Barnuamba at a1. , 2008; Beentje, 1994; Burkill,1997; Bussmann, 2001; Chikamaiet al. , undat-ed; Grace at a1. , 2002a; Green, 2003; Gunb-Fakim & Brendler, 2004; Hawthorne & Jong-kind, 2006; Katende, Birnie & Tengnas, 1995;Liben, 1973; Maundu at a1. , 2001; Muthaura ata1. , 2007; Neuwinger, 2000; Scott, 1981; Shan-gall at a1. , 2004; Sommerlatte & Sommerlatte,1990; Tsukamoto, Hisada & Nishibe, 1985; vanWyk & Gencke, 2000

Sources of illustration Bekele-Tesemma,2007; Hawthorne & Jongkind, 2006;Breitenbach, 1974.

Authors R. Aerts

ONCOBABREVIPES Stapf

Protologue Journ. Linn. SOC. , Bot. 37: 84(1905)

Family F1acourtiaceae (APG: AChariaceae)Chromosome number2n=24

von

Synonyms Coloncobo breujpes (Stapfj Gilg(1908).

Origin and geographic distribution Oneobobreuipes occurs from Gulnea east to westernC6te d'TVoire.

Uses The wood of Oneob0 67euipes is usedfor poles in house building, fence posts, toolhandles and sticks. The seed anIs are edible.

The inner bark and the leaves are used medic-

many in Liberia against headache, either in apoultice or in decoction by draught. The seedoil or a paste from pulverized seeds is used fortreating skin diseases, river blindness andscrofula. The bark yields a fish poison.

Properties The sapwood is greyish, theheartwood pale brown to reddish brown. Thewood is medium-weight and moderately hardand strong. The grain is generally straight,texture fine. The wood is easy to work and fin-ishes fairly smoothly. There is no informationon its durability.

Botany Deciduous small tree up to 15 intall; hole up to 25 cm in diameter; bark surfacesmooth, brown to blackish, inner bark thin,red; twigs covered with scales or resinousglands. Leaves alternate, simple and entire;stipules absent; petiole 1.5-3(-4) cm long;blade obovate-oblong to oblanceolate, 12-20(-26) cm x 4.5-7(-10) cm, base curieate, apexabruptly short-acuminate, papery to thin-leathery, glabrous, pinnately veined with 7-9pairs of lateral veins. Flowers solitary or some-times 2-3 together in axils of leaves on youngtwigs, bisexual or male, regular; pedice1 3.5-4cm long, up to 8 cm in fruit; sepals 3, 0bovate-oblong, c. 3 cm x 2 cm, greenish with whitemargins, with glandular dots outside; petals 9-12, 0blanceolate, c. 6 cm x 3 cm, tapering to-

wards the base, white; stamens numerous,yellow, but arithers becoming brown; ovarysuperior, oblong-ovoid, I-celled, style c. I cmlong, at apex divided into 5-6 stigmatic lobes.Fruit an ellipsoid to ovoid capsule 6-8 cm x 3-5 cm, with faint ridges, many-seeded. Seedsangular, c. 5 min long, enveloped by an anISeedhng with epigeal germination; hypocotyl2-3 cm long, epicotyl c. 1.5 cm long; cotyledonsleafy, oblong-elliptical, c. 1.5 cm long; firstleaves alternate.

In C6te d'TVoire, trees flower in September-October, and young fruits have been collectedin MayOneobo comprises about 35 species, most ofthem in tropical Africa and 6 in tropical Amen-ca. Several genera, including Coloncobo, havebeen merged into Oneobo in 1997. However, in2002 a phylogenetic analysis based on DNAsequences resulted in a position of Calorieoboin the family AChorioceoe, whereas Oneobospinoso Forssk. , the type of Oneobo, was placedin Suitcoceue. Currently research is done toelucidate the status of the species within On-cobo sensu Iato

Oncobo gi!gtono Sprague (synonym: Coloncobogilgiono (Sprague) GIIg) is a shrub or smalltree up to 15 in tall, occurring from Guineaeastward to Garnero0n. Its pale brown andhard wood is used in house construction and

for inlay and cabinet work, and also as fire-wood. The seed anIs are edible. It has showyflowers and may have prospects as ornamental

Ecology Oncobo breuipes occurs in the un-derstorey of evergreen rainforest, also inbrushwood and forest regrowth, often inswampy and occasionally flooded localities, upto 700 in altitude.

Management There are about 4000 seedsper kg. The germination rate of seeds is high,within 8-12 days after sowing

Genetic resources and breeding Oneobobreujpes does not seem to be selectively har-vested for timber or other products and has notbeen classified as endangered or threatened.However, it has a limited area of distributionand its habitatis shrinking

Prospects The wood is likely to remain oflocal importance for posts and poles, and bark,leaves and seeds for medicinal purposes. Theornamental value of Oneob0 67euipes with itsshowy flowers deserves attention

Major references Aubr6ville, 1959c; Cooper& Record, 1931; HUI & Breteler, 1997; Kryn &Fobes, 1959; SIeumer, 1974

Other references Barrier & Muller, 1943;

ONCOBA 505

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Chase at a1. , 2002; de in Mensbruge, 1966;Hawthorne & Jongkind, 2006; Keay, 1954b;Kew Science Directory, undated; Neuwinger,2000; Normand, 1960; T6r6, 2004

Authors L. P. A. Oyen

ONGOKEAGORE (Hua) Pierre

Protologue Bull. Mens. SOC. Linn. Paris 2:1314 (1897).

Family 01acaceaeSynonyms Origoheo Mumeono Pierre (1897),

Origoheu homerunensis Eng1. (1909).Vernacular names ingueuk, boleko, isano

(En). Angueuk, holeko, origok6a (Fr). Nsanu(P0). 1<11eku, ntuli, o1eko (Sw).

Origin and geographic distribution Origo-heu gore occurs from Sierra Leone eastward toeastern DR Congo and southward to Angola.

Uses The wood of Origokeo gore, called 'an-gueuk'in trade, is used mostly locally in heavyconstruction, for railway sleepers and vehicleframes, In Interior and exterior carpentry, forflooring, containers and boxes, turnery and

It Is well suited for interior joineryprovided it is perfectly dry to avoid defor-mation

The seed oil, called 'holeko oil' or 'isano oil', ismedible but can be used as additive to linseed

onin the manufacture of paints, varnishes andlinoleum and to oilfor moulding cores in metalfoundry. It can also be used to protect metaland wooden surfaces. Polymerization at inod-erately high temperatures yields a film withremarkable properties: strong, flexible andinsoluble in acid and alkaline solvents. This

makes it suitable for manufacturing brake

pads and linings. In association with linseed oilthe oil can be made into a standoil(a heat-polymerized oil, very thick and strongly adhe-SIve, but slowly drying; used as a final coat inoil painting) of superior qualities. Boilingholeko oil with copal gives this resin a veryhigh heat resistance. The oil can be used tomake de-emulsifying products for the crude oilextraction industry and for the prevention oficing-up of airplane wings. It can also be vul-canized to yield highly resistant synthetic-rubber products. Ozonolytic cleavage can yieldsaturated double acids, which are used in thesynthesis of polyamides. The use of fatty acidsfrom holeko oil in the manufacturing of sill-cones and of isolating glue for lithium-basedbatteries has been patented. The o111s usedtraditionally to ano1nt the skin.The pulp of the fruitis edible. The bark is laxa-tive; in Congo fresh bark is rubbed on thebreasts of Iactating mothers to purge their ba-bies; similarly, in Gabon a decoction of the barkis used as a wash for babies or they are given apinch of pounded bark mixed with a little saltThe sap is used as styptic and the bark to treatsplenomegaly in DR Congo. The seeds are usedas bait for small rodents and the fruits as

spinning tops for children.Production and international trade The

wood of Origoheo gore is of little importance ininternational trade and is mostly included instatistics under 'miscellaneous timbers'. Few

accurate data are available: Equatorial Guineaexported 400 in31year between 1963 and 1968,while Cameroon exported 500 ing/year in 1997and 1998. In the Central African Republic thetotal extractible volume has been estimated at

3.7 million ina, of which 2.2 million ing is quali-ty class I and 2Boleko oil has been traded in small amounts.

At the end of the 1950s less than 100 t/yearwere exported, although France and Belgiumhad high hopes to develop the use of the oil inthe paintindustry. Potential production at thattime was estimated at 30,000 t/year for DRCongo alone. No information is available on thecurrent production and trade of holeko oil

Properties The heartwood of Origoheo goreis pale yellow to pale brown and darkens onexposure to light. It is indistinctly demarcatedfrom the 6-10 cm thick sapwood. The grain isstraight, sometimes finely interlocked or wavy,texture fine and even. Quarter sawn surfacesare sometimes finely inottled or banded andslightly lustrous. The wood is heavy, with adensity of 840-910 kg/ina at 12% moisture con-

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tent. The rates of shrinkage on drying are high,from green to oven dry 4.0% radial and 10.7%tangential. The wood should be dried slowly,and there is a high risk of distortion and aslight risk of checking. Logs should be quartersawn before drying to avoid warpingAt 12% moisture content the modulus of rup-ture is 94-143 N/mm2, modulus of elasticity10,000-16,135 N/mm2, compression parallel tograin 53-74 N/min2, shear 9-11 N/mm2, cleav-age 20-33.5 N/min and Chalais-Meudon sidehardness 3.0-7.5

Once dry, the wood is easy to work, saw andplane with little blunting of tools. It is easy tofinish, sand and polish. It can be painted, var-nished, waxed and glued without difficulty. Fornailing pre-boring is often required. It can besriced into veneer, but requires much forceThe heartwood is durable; in a test in Japan itwas little affected by decay fungi or termitesand was resistant to marine borers and in atest in Ghana it was little affected in a 3-yearwood graveyard test. The sapwood is sensitiveto blue-stain and to dry-wood borers. Theheartwood is extremely resistant to impregna-tion, whereas the sapwood is moderately re-slstant

The dry seed contains about 63% oil. The seedoil differs from other vegetable oils in its fattyacid composition. Boleko oil has a high iodinenumber, but it does not dry when exposed in athin film such as linseed oil or tung oil. Whenheated to 250'0 a strongly exothermic sponta-neous polymerization reaction starts, whichmay lead to a further increase in temperatureto more than 400'C and to an explosion. Di-acetylenic fatty acids and hydroxy-diacetylenicfatty acids characterize the oil; It consistsmainly of is antc acid and holekic acid (together30-50%) and of isano11c acid (15-35%). It fur-ther contains saturated and unsaturated fattyacids of which linoleic acid is the most jin-

portant one. Isantc acid is an unbranched Ci8-fatty acid with a single ethylene bond and 2conjugated acetylene bonds; its formula is 17-octadecene-9.11-diynoic acid. Bolekic acid is 13-octadecene-9.11-diynoic acid, isano11c acid 17-octadecene-8-hydroxy-9.11-diynoic acid. Theunsaponifiable matter of the oil contains acrystalline dialcohol with molecular formulaC2sH4402.

The pulp of the fresh fruit contains 67% ino1s-ture; its smellis reminiscent of apple, its tasteis sweet but slightly astringent. The root andstem bark of Origoheo gore contain cyclohexa-noid protaflavanones named origokeins; they

are related to sakuranetin and are character-

ized by a non-aromatic CG-ring molety that isotherwise only known from certain ferns.

Description Medium-sized to large, gla-brous tree up to 40 in tall; hole straight andcylindrical, branchless for up to 20 in, 100(-150) cm in diameter, without buttresses butsometimes with heavy root swellings; barksurface grey to dark brown or black, finely fis-sured and peeling off in fine Irregular scales,inner bark I-2 cm thick, softly granular, yel-low-brown; crown pyramidal, rather open, withfew heavy branches; twigs lateralIy coin-pressed. Leaves alternate, simple and entire,without stipules; petiole thin, 0.5-I cm long,grooved above, decurrent into 2 fine ridgesalong the branch; blade elliptical, 4-12 cm x 2-5 cm, base rounded to curieate, apex short-acuminate, papery, glabrous, pinnately veinedwith 6-10 pairs of lateral veins. Inflorescencean axillary panicle up to 15 cm long, consistingof densely flowered, umbel-shaped cymesFlowers bisexual or functionally unisexual,regular, 4-merous, greenish; pedicel slender, c.6 min long; calyx shallowly cup-shaped, c. Imm in diameter; petals strap-shaped, 3-4 mm

ONGOKEA 507

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Origoheo gore - I, 60se of 601e, ' 2, floweringtwig, . 3, flower, ' 4, fruit, ' 5, fruit stone.Redrawn und odopted by Ishoh Syomsudin

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long, recurved; disk 4-10bed; stamens unitedinto a tube c. 3 min long; ovary superior, ses-SIIe, I-celled, style hardly exserted from thestaminaltube. Fruit a globose drupe 2-4 cm indiameter, enclosed by the enlarged calyx exceptfor apical part, slightly acuminate, I-seeded.Seed globose, c. 1.5 cm in diameter. Seedlingwith epigeal germination; hypocotylvery short,epicotyl c. 18 cm long; first pair of leaves OPPo-site.

Other botanical information Origoheocomprises a single species. It is closely relatedto Aptondru, a genus with about 4 species intropical America and one species in tropicalAfrica, Aptondro zenheri Engl. , which differsfrom Origoheo gore in its raceme-like inflores-cences and large, collar-shaped, pinkish calyxsurrounding the fruit

Anatomy Wood-anatomical description (IAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(7: vessels in diagonal and/or radial pattern); 9vessels exclusively solitary (90% or more); 13simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); 25: intervessel pits small (4-7 pin); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; 31: vessel-ray pits with much re-duced borders to apparently simple: pitsrounded or angular; 33: vessel-ray pits of twodistinct sizes or types in the same ray cell; 42mean tangential diameter of vessellumina100-200 I'm; 47: 5-20 vessels per square inn-limetre. Tracheids and fibres: 62: fibres withdistinctly bordered pits; 63: fibre pits commonin both radial and tangential walls; 66: non-septate fibres present; 69: fibres thin- to thick-walled; 70: fibres very thick-walled. Axial pa-renchyma: 76: axial parenchyma diffuse; 77axial parenchyma diffuse-in-aggregates; 78axial parenchyma scanty paratrachea1; (86:axial parenchyma in narrow bands or lines upto three cells wide); 92: four (3-4) cells per pa-renchyma strand. Rays: 97: ray width I-3 cells;(104: all ray cells procumbent); 106: body raycells procumbent with one row of uprightand/or square marginal cells; 1/6: z: 12 rays permm. Mineral Inclusions: 136: prismatic crystalspresent; 138: prismatic crystals in procumbentray cells; 139: prismatic crystals in radialalignment in procumbent ray cells; 140: pris-matic crystals in chambered upright and/orsquare ray cells; 142: prismatic crystals inchambered axial parenchyma cells.

(E. Uetimane, P. Baas & H. Beeckman)Growth and development In C6te d'TVoire

Origoheo gore flowers from January to Juneand fruits from May to July; in DR Congo fruit-ing is abundant in September, in Gabon inDecember and January. The fruits are eaten bymany animals and the seeds are dispersed e. g.by monkeys.

Ecology Origoheo gore is found scattered indense evergreen forest and in moist semi-deciduous forest. It occurs on dry ground andin periodically inundated localities. In Gabon itoften occurs in forest dominated by SOCoglottisgobonensis (Baill. ) Urb. and AucoumeoMumeono Pierre.

Propagation and planting Germination isslow and may take several months and evenmore than one year. Because of its slow andirregular germination, Origoheo gore is notgrown In nurseries

Management Large trees of Origoheo goreoccur scattered in the forest. In Liberia I tree

with a hole diameter over 60 cm has been re-ported per 43 ha for evergreen forest, and Itree per 7.5 ha for moist semi-deciduous forest.

Harvesting Fruits of Origokeo gore are col-leeted from the wild and mostly the pulp isallowed to rot away before the fruit stones arecollected from the soil.

Handling after harvest Fresh logs sink inwater and cannot be transported by river.Depulping of fruits can be done by passing thefruits between rubber rollers and washingthem with cold water. Boleko oilis produced byhydraulic pressing, but this is hampered by thehigh viscosity of the oil. During pressing thetemperature can rise to 80"C which can alterthe properties of the oil. The press cake con-tains considerable amounts of polymerized oilThe cake is unsuitable as cattle feed, but canbe used as manure. The oilcan also be extract-ed by solvents after the kernels have beenground and subjected to treatment with coldmethanol

Genetic resources Origokeo gore is wide-spread and does not seem to be in danger ofgenetic erosion. No germplasm collections areknown to exist.

Prospects Origoheo gore is likely to remainimportant in its region of origin. There are noindications that it will become a commodity ininternational trade, but its volume in miscella-neous timber lots is likely to increase. Demandfor the o111s likely to remain low except if localpaint industries develop or if new applicationsfor its unique fatty acids are found

Major references anonymous, 1957a; Au-br6viUe, 1959b; Chudnoff, 1980; CIRAD Forest-ry Department, 2003; CTFT, undated; Miller ata1. , 1977; Normand, 1950a; Fouliquen, 1959;Vieux & Taratibu, 1968; Voorhoeve, 1979

Other references Burki11, 1997; De Borger,1960; De Vries, 1956; De Vries, 1957; Heckel,1902; Jerz, Warnel & Achenbach, 2005; Keay,1989; Libouga, Womeni & Bitjoka, 2002; Mag-110cca, 1998; Mangala, 1999; Normand &Paquis, 1976; Pauwels, 1993; Raponda-Walker& Sillans, 1961; Sallenave, 1955; Saunders &Hall, 1968; Tsunoda, 1990; Viniers, 1973b; vonMikusch, 1963; von Mikusch, 1964; Wilks &TSSemb6, 2000

Sources of illustration Pauwels, 1993;Voorhoeve, 1979; Wilks & TSSemb6, 2000

Authors D. Louppe

up to 30(-40) in tall; hole usually straight andlong, up to 90(-120) cm in diameter, with steepand flat buttresses up to 4 in high; bark surfacefairly smooth, peeling off in thin flakes, yellow-ish green to pale brown or pale grey, Innerbark thin, brittle, yellow-orange; crown round-ed, rather open, with spreading branches;twigs slightly short-hairy or glabrous. Leavesalternate, simple and entire; stipules minute,caducous; petiole c. I cm long; blade elliptical,7-15 cm x 2.5-6 cm, curieate to obtuse at base,acuminate at apex, papery to thin-leathery,glabrous, with I-2 pairs of lateral veins fromnear the base of leaf blade and c. 3 pairs ofadditional lateral veins. Inflorescence a termi-

nal panicle c. 20 cm long, with long, slenderbranches, short-hairy, many-flowered. Flowersbisexual or male, regular, small, creamy togreenish white, scented; pedicel c. I mm long;sepals 5, free, ovate, c. 2 mm long; petals ab-sent; stamens 5-6, c. 2.5 mm long; disk withhairy lobes alternating with stamens; ovarysuperior, ovoid, hairy, I-celled, style thick, with3 spreading branches. Fruit an ellipsoid cap-SUIe up to 1.5 cm x I cm, pointed, short-hairy,pinkish to reddish, dehiscent with 3 valves, 3-6-seeded. Seeds with small, lobed anI. Seedlingwith epigeal germination; hypocoty1 3-3.5 cmlong, epicotyl c. 2 cm long, short-hairy; cotyle-dons leafy, rounded, c. 1.5 cm long; first leavesalternate, slightly toothed.In C6te d'TVoire trees have been recorded to

flower in June and fruits have been collected in

August-October.Ophiobotrys comprises a single species. It isrelated to Coseorio and several genera re-stricted to tropical America and tropical Asia

Ecology Ophiobotrys genheri occurs mainlyin lowland semi-deciduous forest

Management There are about 25,000 seedsper kg. The seeds germinate within 8-12 daysafter sowing, but the germination rate is re-portedly low, 15-25%. Larger trees are oftendifficult to fell because they have large but-tresses. Freshly harvested logs sink in waterand cannot be transported by river

Genetic resources and breeding AlthoughOphiobotrys zenheri is quite widespread, itseems to be uncommon or even rare In manyregions within its distribution area. From somecountries very few herbarium collections areavailable, e. g. from Cote d'TVoire and Gabon.

Prospects Very little is known about Ophi-o60trys zenkeri. It seems to be rarely exploitedfor timber, probably because it is uncommonand has hard wood and large buttresses.

OFHIOBOTRYSZENKERIGilg

Protologue Bot. Jahrb. Syst. 40(4): 516(1908).

Family F1acourtiaceae (APG: Salicaceae)Origin and geographic distribution Ophio-

botrys zenheri occurs from Liberia east to theCentral African Republic and south to Gabonand western DR Congo.

Uses The wood is used locally, but there islittle technical information on its uses. In Gha-na it has been used for utensils including inor-tars.

Properties The pale brownish yenow woodis moderately heavy and hard. It has someresemblance to the wood of Scottellio hmmeonoPierre.

Botany Shrub or smallto medium-sized tree

OFHiOBOTRYS 509

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Ophtobotrys zenheri- wild

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Major references Aubr6ville, 1959c; Burkill,1994; Normand, 1960; Normand & Paquis,1976; SIeumer & Bamps, 1976.

Other references de to Mensbruge, 1966;Hawthorne, 1995; Hawthorne & Jongkind,2006; HUI, 1995; InsideWood, undated; Irvine,1961; Keay, 1954b; Keay, 1989; Miller, 1975;Tailfer, 1989

Authors R. H. M. J. Lemmens

OUBANGUIAAFRICANA Baill.

Protologue Bull. Mens. SOC. Linn. Paris a869 (1890).

Family Scytopetalaceae (APG: Lecythida-ceae)

Synonyms Oubongttio denticuloto Tiegh.(1905), 0,160nguio IOUrentii Bail1. (1908).

Origin and geographic distribution Oubon-guto of neono is found in Cameroon, CentralAfrican Republic, Equatorial Guinea, Gabon,CongoandDR Congo

Uses The wood of Oubonguio ofricono isused in Gabon to make packing cases. In DRCongo the bark is powdered and taken orally tocure backache and haemorrhoids.

Properties The wood ofOubonguio of neonois greyish pink, heavy, and similar to that ofthe commercially more important timber treeScytopeto!urn tinghemii (Achev. ) Hutch. &Dalziel.

Botany Smallto medium-sized tree up to 20in tall; hole often short, up to 50(-80) cm indiameter; bark surface longitudinalIy fissured,pinkish, inner bark fibrous, reddish; crownsmall; twigs angular. Leaves alternate, crowd-ed towards the ends of twigs, simple; stipules

very small, caducous; petiole I-3 mm long;blade ovate to lanceolate, sometimes elliptical,

base curieate and8-16 cm x 3.5-6.5 cm,

asymmetrical, apex long-acuminate, marginentire to finely toothed, leathery, glabrous,pinnately veined with 6-8 pairs of lateralveins. Inflorescence an axillary or terminalpanicle up to 15 cm long. Flowers bisexual,regular; pedice1 5-6(-8) mm long; calyx saucer-shaped to cup-shaped, 3-4 mm in diameter,margin entire or incised; petals 6-8, 6-8 mmlong, recurved, white; stamens numerous, c. 5mm long; ovary superior, globular, 3-4(-5)-celled, style slender, c. 5 mm long, stigmasmall. Fruit a globose to obovoid capsule, 1.5-2cm x I-1.5 cm, violet, usually I-seeded. Seedoblong, 9-11 mm x 6-7 mm.Oubonguio comprises 3 species. OubonguioIOUrifo!to (Pierre) Tiegh. is a rare species re-stricted to south-eastern Nigeria, Garnero0nand Gabon. It is a smalltree up to 15 in tallwith a hole diameter up to 60 cm. In Cameroonthe wood is used to make oars.

Oubongui0 o10to Baker f. is also found insouth-eastern Nigeria, Garnero0n and Gabon,but nowhere further from the coast than 100

kin. It is a smallto medium-sized tree up to 20in tallwith a low-branching hole up to 50 cm indiameter. The wood is probably used for SImi-Iar purposes at the other Oubonguio sppFruits are produced in abundance in someyears and the seeds are often eaten by chil-dren. The seeds are hard, low in oilcontent andthey taste like coconut but have a bitter after-taste

Ecology Oubo, Iguio of neono is found inswamp forest and periodically inundated for-est, and along rivers

Genetic resources and breeding Oubon-gum ofricoito is fairly widespread and occurslocally abundantly, although in general consid-ered to be rather uncommon. At present, thereis no reason to consider this species as threat-ened

Prospects 01160ngui0 o17icono and otherOubonguio spp. will probably remain of inter-est for local use only.

Major references numbe Bayeli, 2006;Letouzey, 1978a; Raponda-Walker & Sillans,1961.

Other references Breteler, 2002; Breteler,2005; Germain, 1963; Lebrun & Stork, 2003;Moyersoen, Alexander & Fitter, 1998; Nor-inarid & Paquis, 1976; Thomas at a1. , 1989;Vivien & Faure, 1985; Vivien & Faure, 1996.

Authors C. H. BOSch

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Oubo, Igutoof neono- wild

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OXYAl{THUS ZANGUEBARICUS (Hiern) Bridson

Protologue Kew Bull. 34(I): 1/9 (1979)Family RubiaceaeVernacular names Mfupapu (Sw)Origin and geographic distribution Oryon-

thus 20ngueboricus is distributed in Somalia,Kenya, Tanzania and Mozambique.

Uses In Kenya the stems are used as polesin building. In Mozambique the wood is madeinto dishes, spoons and other utensils

Properties The wood is durableBotany Shrub or smalltree up to 8.5 in tall,

much-branched; twigs somewhat grooved, usu-ally glabrous. Leaves opposite, simple and en-tire; stipules triangular, 5-11 min long, acu-inmate at apex, persistent; petiole 3-10 mmlong; blade elliptical to narrowly ovate, 5-15.5(-19) cm x 2-6(-7) cm, base obtuse torounded, apex acute to short-acuminate, thin-leathery, usually glabrous, pinnately veinedwith 8-9 main pairs of lateral veins. Inno-rescence an axillary, compact, narrowly pyram-Idal panicle, nearly sessile, (3-)5-12-110wered;bracts lanceolate or linear-lanceolate, (7-)10-12 mm long. Flowers bisexual, regular, 5-meTous; pedicel up to 5 min long; calyx gla-brous, tube 4-5 min long, lobes linear-subulate,(4-)5-8 mm long; corolla white, tube 7-11.5 cmlong, lobes oblong-lanceolate, I-2 cm long,acuminate; stamens inserted on corolla throat,alternating with lobes, arithers nearly sessile;ovary inferior, I-2-celled, style slender, at apexwith an oblong-ellipsoid pollen presenter 2-2.5mm long. Fruit a pear-shaped berry 3.5-7 cm x2.5-3 cm, greenish flecked with white. Seedscompressed-ellipsoid, c. 6 mm long, blackishbrown, strongly striated

Oryonthus comprises about 35 species and isconfined to mainland Africa. The wood of sev-

eral species is used in tropical Africa for SImi-Iar purposes as that of Oryonthus gonguebori-cus, but most of these are more important asornamental shrub or tree.

Ecology Oryonthus gongueboricus occurs inopen forest including coastal and riverme for-est, woodland and bushland, from sea-level upto 500 in altitude

Genetic resources and breeding Oxyan-thus gongueboricus has a limited distribution,butts fairly common and seems notthreatenedwith genetic erosion.

Prospects Oxyanthus gongueburicus Is auseful local source of wood for construction

poles and utensils. Information on its woodproperties Is lacking, but in view of its smallsize its importance is unlikely to increase

Major references Beentje, 1994; Bridson &Verdcourt, 2003; Verdcourt & Bridson, 1991.

Other references Bridson, Thulin & De-greef, 2006; d'OliveIra Feij50, 1960.

Authors M. Brink

PARAMACROLOBIUM 511

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FARM^CROLOBIUMCOERULEUM (Taub. )J. Leonard

Protologue Bull. Jard. Bot. Etat 24(4): 348(1954).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Chromosome number2n=24

Synonyms Mocrolobium coeruleotdes DeWild. (1907), Mocro!obium coeruleum (Taub. )Harms (1915), Macro!obium dowei Hutch. &Dalzie1(1928)

Vernacular names Mkwe (Sw).Origin and geographic distribution Ford-

inocrolobium coeruleum is widespread, fromGuinea east to Kenya and south to DR Congo,Tanzania and northern Angola. However, thedistribution area is notably discontinuous.

Uses In DR Congo the wood is used for join-ery, doors, frames of doors, furniture, railwaysleepers and gongs. It is suitable for heavyflooring, Interior trim, toys, novelties, turnery,carving, veneer and plywood. Poromocrolobi-urn coeruleum has been used as shade tree incocoa plantations in Sierra Leone. In Kenyathe bark is used as rough fibre. The seeds areused in games.

Properties The heartwood is pale brown toyellowish brown or pinkish brown with darkerbrown streaks on quarter-sawn surfaces, andOxyonthus 20ngueboricus - wild

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Purgingcrolobium coeruleum - wild

usually indistinctly demarcated from the sap-wood. The grain is straight to interlocked, tex-ture moderately coarse and even. The wood ismoderately heavy, with a density of about 700kg/in3 at 12% moisture content, and is moder-ately hard. The rates of shrinkage during dry-ing are high and the wood may develop severesurface splits and cupping, whereas it is sus-ceptible to blue stain attack during drying. Thewood saws and works well, is easy to plane andtakes a good finish. The gluing and bendingproperties are satisfactory. The wood is moder-ately durable, being liable to termite, Lyctusand marine borer attacks

Several long-chain fatty acids have been iso1at-ed from root bark extracts; these acetylenicacids showed inhibitory activity of 3-hydroxy-3-methylglutaryl coenzyme A reductase, the en-zyme responsible for the formation of mevalo-nate in the rate-determining step of cholesterolbiosynthesis

Botany Evergreen, medium-sized to fairlylarge tree up to 35(-40) in tall; hole branchlessfor up to 20 in but often low-branching,straight and cylindrical or irregular, up to 90(-100) cm in diameter, with buttresses; barksurface nearly smooth, but with fine longitudi-nal grooves and lenticels, becoming flaky, grey-brown, inner bark fibrous, brownish to reddish;crown often long and narrow, often with droop-Ing branches; twigs usually glabrous, with Ien-ticels. Leaves alternate, paripinnately coin-pound with 2-5 pairs of leaflets; stipules fused,up to 1.5 cm long, clasping the twigs, persis-tent; petiole and rachis together 5-22 cm long,grooved; petiolules I-4 min long, twisted; leaf-lets opposite, elliptical to oblong or lanceolate,

2-15 cm x I-6 cm, asymmetrical at base, acu-inmate at apex, thinly leathery, glabrous, pin-nately veined with 10-15 pairs of indistinctlateral veins. Inflorescence a terminal, nat-topped panicle 4-8 cm long, usually glabrousFlowers bisexual, zygomorphic, with 2 bracte-o1es up to 3.5 cm long at base; pedice1 1.5-3 cmlong; sepals 4, I-1.5 cm long, I broader thanthe other 3 and 2-toothed; petals 5, free, une-qual, bluish, upper one largest, up to 4.5 cm x2.5 cm, lateral 2 up to 3 cm x 2.5 min, lower 2minute; stamens 9, fused at base, usually 3fertile, large, up to 3.5 cm long and 6 rudimen-tary; ovary superlor, linear, I-1.5 cm long,hairy, with long stipe, I-celled, style 2-2.5 cmlong. Fruit an oblong, flattened pod 8.5-20 cmx 2.5-6 cm, glabrous, dehiscing with 2 spiral-ling woody valves, 3-8-seeded. Seeds rectangu-Iar, flattened, 1.5-2.5 cm x I-2 cm, glossy darkbrown, seed coat hard. Seedling with epigealgermination; hypocoty1 5-12 cm long, epicotyl4-7 cm long; first 2 leaves opposite, with 2-3pairs of leaflets.In the nursery seedlings are about 30 cm tallafter 4 months and 60 cm tall after 15 months.In Guinea 6-years-old saplings showed a meanannual height growth of 85-130 cm, but in DRCongo planted trees were only 4.5 in tall after11 years and had a high mortality. Fruits areeaten by monkeys and large rodents, whichmay serve as seed dispersers.Paromocrolobtum comprises a single speciesand seemsto be related to Cryptosepolum

Ecology Purgingcrolobium coeruleum occursin lowland rainforest up to 450 in altitude, butalso in gallery forest in savanna areas and inwooded savanna

Management Seeds can be stored for atleast 18 months, when the viahinty is stillabout 40%. Pre-treatment of seeds is not need-ed, but soaking in cold water for I-2 days or inboiling water for a few minutes acceleratesgermination. Seedlings should be planted outin the full sun and in fertile soils to obtain goodgrowth. They are planted at a spacing of 3 in x3 in in pure stands or together with other inod-erately fast growing timber species.

Genetic resources and breeding AlthoughFordingcrolobium coeruleum occupies a largedistribution area, it is absent in large regionswithin that area and thus shows a disjunctdistribution pattern. It is common in manyregions, and there is no reason to consider itthreatened by genetic erosion

Prospects Poromocro!obium coeruleum willremain a locally Importanttimber tree, but it is

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unlikely that it will gain importance for coin-mercial exploitation because in areas where itis common the logs are usually too small andtoo poorly shaped. However, surprisingIy littleis known about this widespread tree that mayhave prospects in agroforestry systems or astimber tree in plantations. Research on propa-gation and growth rates is recommended.

Major references Bolza & Keating, 1972;Brenan, 1967; Chikamai at al. , undated; Patilat a1. , 1989; Savill & Fox, 1967.

Other references Beentje, 1994; Burkill,1995; GIIbert & Bellefontaine, 1973; Haw-thorne & Jongkind, 2006; Hubert, undated;Lewis at a1. , 2005; Normand & Paquis, 1976;Pauwels, 1993; Wilczek at a1. , 1952; Wimbush,1957

Authors R. H. M. J. Lemmens

mountains of East Africa and in the northern

part of the Zambezian region. It occurs fromSenegal eastward to Sudan and Uganda, andsouthward to Mozambique and Angola. InSouth America it is found from Bolivia, Peruand Colombia to eastern Venezuela, the GUI-arias and Braziland northward to Costa Rica

Uses The wood, known in trade as 'sougu6''kokodi' or 'African greenheart', is traditionallyused for house posts. Tenda people from Sene-gal, Guinea Bissau and Guinea use it to makebarrels, drums and mortars. In Zambia it isused similarly and to make dug-out canoes.The wood is suitable for heavy construction,also for hydraulic works in sea water, heavy-duty flooring, poles, piles, joinery, interior trim,mine props, vehicle bodies, furniture, ladders,sporting goods, agricultural Implements, toolhandles, veneer, plywood and block-board. Ithas been used for railway sleepers after treat-merit with preservatives. It is in demand asfirewood and for charcoal production.The pulp of the fruit is eaten, but is not as fla-vourful as that of Portnori eurotellifoliuPlanch. ex Benth. The fruit pulp is also boiledwith groundnuts into a sauce, or is fermentedinto an alcoholic drink. Seeds are oily and eat-en after roasting. Roasted bark is added topalm wine to improve its flavour.Poringri excelso has numerous uses in tradi-

tional medicine in Africa. In the Central Mri-

can Republic an extract of the bark is takenagainst gall bladder complaints, and a macer-ate of the bark is taken in wine as a sexual

stimulant. In central DR Congo a decoction ofthe bark is drunk against chest pain. In C6ted'TVoire bark decoctions are taken by pregnantwomen against anaemia and, in combinationwith a plaster of the leaves, against rheumaticpains. In Tanzania a decoction of the bark en-ters into medicines against malaria, fever andparasites such as hookworm. In Senegal andSierra Leone bark decoctions are taken againststomach problems including dysentery. In Sen-egal and DR Congo a plaster made of the barkis applied to circumcision woundsIn Senegal and the Central African Republic adecoction of the leaves is applied as a wash orits vapour Is inhaled to treat gingivitis andtoothache. In Senegal women take root decoc-tions against headache, stomach pain and in-fertility. A maceration of the roots is applied towounds as antiseptic and haemostatic;it is alsodrunk and applied as a bath against pain ofvarious causes and amnesia. An infusion of the

fruit pulp is taken against diarrhoea

PARINARIEXCELSA Sabine

Protologue Trans. HOTt. SOC. London 5: 451(1824).

Family ChrysobalanaceaeChromosome number2n=20

Synonyms Portnoriholstii Eng1. (1895).Vernacular names Mubura, grey plum,

Guinea plum, rough-skinned plum (En). Pan-de Gum6enari, manguier sauvage, prunLer

(Fr). Farmari, mainpataz (Po). unula, inbura(Sw)

Origin and geographic distribution Port-nonexcelso is extremely widespread, occurringon both sides of the Atlantic Ocean. In Africa it

occurs in nearly all forest types of the Gumeo-Congolian region, but is absent from the wet-test parts; it is also widely distributed on the

PARTNERi 513

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Poringtiercelso - wild

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The tree is used for shade in Cincho, ,o and

coffee plantations. The nectar in the flowers iscollected by honey bees. Leafy twigs are usedas fodder. A dye obtained from the fruit pulpand fruit stones is traditionany used to colourleather. In Tanzania the bark has been used in

tanning leather. The fibre of the fruit stones isused as tinder to start fire

Production and international trade For-

merly, the wood of Poringri excelso was ex-PIOited commercially, e. g. in Sierra Leonewhere it was logged until 1964; later it wasconsidered undesirable because it is too hard,difficult to work and perishable. Like otherPoringri spp. , it has now no commercial valuein international timber markets, and even inthe countries where it grows its value as tim-ber is limited. It cannot be used economicalIywhen resawing and machining of dried woodare required, and may only be used on a largerscale for construction purposes as posts or inlarge sizes, sawn when still green.In local markets in Guinea fruits were sold in

2002 at Us$ 0.15-0.25 per kgProperties The heartwood is yenowish brown,

darkening upon exposure to deep red or choco-late brown, and not clearly demarcated fromthe yellowish white sapwood. The grain iswavy to interlocked, texture moderately coarseto coarse. Fresh sapwood has a smellofhoney.The wood is heavy, with a density of (530-)730-920 kg/ina at 12% moisture content, andhard. It dries slowly with a tendency to distort,check and split. Air drying prior to kiln dryingand mild kiln drying schedules are recoin-mended. Boards of 2.5 cm thick were dried in

Liberia to 20% moisture content in 3 months.

The shrinkage rates are high, from green tooven dry 5.0-7.6% radial and 8.3-11.4(-12.8)%tangential. Once dry, the wood is moderatelystable to unstable in service.

At 12% moisture content, the modulus of rup-ture is 1/4-204 N/mm2, modulus of elasticity10,900-18,050(-23,900) N/min2, compressionparallel to grain 42-88 N/mm2, shear 10-16N/mm2, cleavage 15-26 N/mm, Janka sidehardness 7640-8880 N, Chalais-Meudon sidehardness (2.9-)4.5-7.8 and Janka end hard-ness 8220-10,890 N.The wood is difficult to saw and work with

hand and machine tools, and has a strongblunting effect on saw teeth and tool edges dueto its high silica content (usually more thanI%). It is best sawn with stellite-tipped bladesand when still green. In planing picking up ofgrain may occur due to the presence of inter-

locked grain. The wood finishes wellif a filleris used. Pre-boring is required for nailing andscrewing, but holding properties are good.Steam bending properties are moderate. Thegluing properties are moderate. The wood istechnicalIy suitable for plywood or veneer onlywhen processed green. It is not suitable forturning or moulding. The wood is reslstant tomarine borers, but susceptible to attacks bytermites and boring insects. Its susceptibilityto fungal decay is moderate; in tests in Tanza-nia untreated wood lasted for less than I yearin contact with the ground. The sapwood ispermeable to preservatives, the heartwoodmoderately resistant but easily treatable underpressure. The wood is not suitable for pulpingThe yellowish fruit pulp of Portnoriexcelso hasa taste like avocado, is very nutritive and con-tains 38-40% carbohydrates. The seed yieldsabout 14% oil. The oilis edible, but dries quick-Iy. It contains or-elaeostearic acid. In spite ofthe numerous uses of most parts of the tree intraditional medicine, little research has beendone into its pharmacological compounds orproperties

Description Evergreen large tree up to 45(-

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Portnori excels0 - I, 60se of 601e, ' 2, floweringtwig, . 3, flower, . 4, fruit.Redrown Grid odopted by J. M. de Vries

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50) tall; bole branchless for up to 20(-25) in,straight or somewhat sinuous, cylindrical, upto 150 cm in diameter, with buttresses up to 3in high; bark surface slightly rough with nu-merous warty lenticels or with deep longitudi-nal fissures, yellowish grey to brownish grey,scaling off in flakes, inner bark hard, granularand brittle, red or pale reddish brown withwhitish spots, smelling sour, with a resinousexudate; crown rounded or flattened, cauliflow-er-shaped, dense, golden brown from below;twigs sparsely hairy. Leaves alternate, simpleand entire; stipules up to 2 mm long, caducous;petiole 3-8 min long, terete, with 2 glands be-coming inconspicuous with age; blade ovate tooblong-elliptical, 3-10 cm x 1.5-5 cm, basecurieate to rounded, apex acuminate, leathery,glabrous above, densely brownish or greyishhairy below, pinnately veined with 13-24 pairsof lateral veins. Inflorescence a terminal or

axillary panicle, up to 12 cm long, brownishhairy. Flowers bisexual, slightly zygomorphic,5-meTous; pedice1 I-2 min long; receptacle bell-shaped to top-shaped, 2-3 min long, hairy; sep-als narrowly ovate to triangular, 2-3 min long,hairy; petals free, obovate to narrowly ellipti-cal, 2-2.5 mm long, pinkish white; stamens 7(-8), c. 2 mm long, on I side of flower, with 7-8rudimentary stamens on opposite side; ovarysuperior, Inserted in upper part of receptacle,hairy, 2-celled, style originating from base ofovary, 2.5-3.5 min long. Fruit an ellipsoid tonearly globose drupe 2.5-6 cm x 2-4 cm, warty,yellowish to reddish brown when ripe, pulpfleshy, yellowish; stone granular and fibrous,slightly ridged, I(-2)-seeded. Seedling withhypogeal germination; epicoty1 5-10 cm long,hairy; cotyledons remaining within seed coat;first c. 8 leaves strongly reduced, first normalleaves alternate.

Other botanical information Portnori

comprises about 40 species and is pantropicalwith most species in tropical America and trop-ical Asia and 6 in Africa. In general, the spe-cies are remarkably similar despite the verylarge distribution area of the genus. Poringriexcelso is quite variable, particularly in indu-meritum colour, shape of inflorescence andflower size.

Poringri hypochryseo Mildbr. ex Letouzey &Funiite closely resembles Poringriexcelso, butdiffers in its rounded to cordate leafbases. It is

locally abundant in a narrow band of evergreencoastal rainforest in Nigeria, GamerDon, Equa-tonal Guinea and Gabon, and its wood is POS-sthly used for similar purposes as that of

Formariexcelso.

Portnorz congensis Didr. is an evergreen medi-urn-sized tree up to 30 in tall with often short,tortuous hole up to 100 cm in diameter, charac-tenstic of swamp forest and periodically flood-ed forest from Guinea and Maneastward to the

Central African Republic and southward to DRCongo. Its orange-brown and hard wood is usedfor rafters and for posts and poles in construc-tion, and is probably suitable for other purpos-us similar to Poringtiercelso. In DR Congo thebark is used as purgative and enters into medi-cines against leprosy, in Congo it is usedagainst dysentery. The fruits are eaten andused as fish bait. The seeds are also edible

Anatomy Wood-anatointcal description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;9: vessels exclusively solitary (90% or more);13: simple perforation plates; (19: reticulate,foraminate, and/or other types of multiple per-foration plates); 22: intervessel pits alternate;25: intervessel pits small(4-7 prn); (30: vessel-ray pits with distinct borders; similar to inter-vessel pits in size and shape throughout theray ceU); (31: vessel-ray pits with much reducedborders to apparently simple: pits rounded orangular); 32: vessel-ray pits with much reducedborders to apparently simple: pits horizontal(scalariform, gash-like) to vertical (palisade);43: mean tangential diameter of vesselluminaZ 200 F1m; 46: S 5 vessels per square minimetre;(47: 5-20 vessels per square minimetre). Tra-cheids and fibres: 62: fibres with distinctlybordered pits; 63: fibre pits common in bothradial and tangential walls; 66: non-septatefibres present; 69: fibres thin- to thick-walled;70: fibres very thick-walled. Axial parenchyma86: axial parenchyma in narrow bands or linesup to three cells wide; 93: eight (5-8) cells perparenchyma strand; 94: over eight cells perparenchyma strand. Rays: (96: rays exclusivelyuniseriate); 97: ray width I-3 cells; 104: all raycells procumbent; (106: body ray cells procum-bent with one row of upright and/or squaremarginal cells); 1/6: z 12 rays per min. Mineralinclusions: 159: silica bodies present; 160: siftcabodies in ray cells(E. Ebanyenle, P. E. Gasson & E. A. Wheeler)

Growth and development Growth followsthe tree architectural model of Troll, in whichall axes grow horizontally, butthe basal part ofeach axis can assume vertical growth after leafloss to become the new main axis. In Liberia

and C6te d'TVoire trees flower in January-

PARTNER1 515

516 TIMBERS2

April(June) and fruit in October-January (-March); in Nigeria flowering occurs in Janu-ary-May and October-November, and fruitsmature in November-December and March; ineastern Tanzania flowering occurs in August-March, and fruit maturation peaks in Novem-her-December; in Zambia flowering occurs inJuly-October, and fruiting in May-NovemberSeed dispersal is mainly by elephants, whicheat the fruits, but some studies indicate thatdispersal is very limited. Regeneration in ina-ture forest is poor, except in clearings or alongroads. In secondary vegetation gregariousstands of all ages may develop from seedlingsand root suckers, and this may eventually re-SUIt in a single-dominant Portnoriexcelso highforest

Ecology Poringri excelso Is a characteristiccanopy tree occurring throughout the Africanrainforest, but also in drier forest types such asdry evergreen forest and well-drained fringingforest, in areas with average annual tempera-tures of 10-27"C, up to 2100 in altitude. Some-times it persists in grassland after the destruc-tion of forest, and then may act as centre forthe re-establishment offorest

Propagation and planting Germination offruit stones takes 2 months to 3 years, and thegermination rate is reported to be low, alt-bough natural regeneration may be abundantunder optimal circumstances. Poringri excelsocan also be propagated by wildlings and rootsuckers.

Management Densities of up to 08 treesper hectare with a hole diameter of more than60 cm have been recorded in Liberia. The high-est density observed in Sierra Leone was 7.5treesA1a with a hole diameter of 20-60 cm, de-creasing to 0.1 trees/ha with a hole diameter ofmore than 120 cm. Portnori excelso can be

managed by lopping, coppicing and pollardingSeedlings planted into the field should be pro-tected against herbivores.In logged-over forest where Portnori excelsotrees have not been removed during loggingoperations, Its numbers may increase as youngtrees continue to grow and regeneration can beabundant. Under such circumstances, Poringriexcelso may become difficult to control, sup-pressing more valuable tree species.

Harvesting In the Central African Republicand Gabon, the minimum bole diameter al-lowed for harvesting is 70 cm.

Handling after harvest Freshly harvestedlogs sink in water and cannot be transportedby river

Genetic resources Poringri excelso is ex-

tremely widespread and locally common, alsoin disturbed forest. Although there are reportsthat it is locally becoming threatened, there isno general risk of genetic erosion

Prospects Although the wood of Poringriexcelso is difficult to work, it is available inlarge sizes and fairly large amounts and maygain importance as better tools have becomeavailable. In its area of distribution it will re-

main important for construction purposes,firewood and charcoal production, and as asource of edible fruits and products for tradi-tional medicine. It may be promising for agro-forestry systems, but more research is neededon suitable propagation methods and SIIvicul-turalpractices

Major references Aubr6ville, 1959c; Bolza& Keating, 1972; Burki11, 1985; Burki11, 2000;Graham, 1960; Letouzey & White, 1978a;Phongphaew, 2003; Prance & Sothers, 2003a;Takahashi, 1978; Vivien & Faure, 1985.

Other references Baerts & Lehmann,2012; Chilufya & Tengnas, 1996; Coradin,Giannasi & Prance, 1985; de in Mensbruge,1966; de Saint-Aubin, 1963; Edema, Oderoha &Daro-Idol10, 2009; FAO, 1994; Gross-Camp,Mulindahabi & Kaplin, 2009; Keay, 1958i; Kryn& Fobes, 1959; Lovett at a1. , 2007; 0gunka-Nnoka & Mepba, 2008; Poorter at a1. , 2004;Prance & White, 1988; Ruffo, Birnie & Teng-rias, 2002; Vivien & Faure, 1996; Voorhoeve,1965; White, 1976; White, 1978a; White & Ab-ernethy, 1997.

Sources of illustration Keay, 1958i; Vivien& Faure, 1985; Voorhoeve, 1965

Authors L. FA. Oyen

FAUSINYSTALLAJOHIMBE (K. Schum. ) Pierreex Beille

Protologue Act. SOC. Linn. Bordeaux 61:130 (1906).

Family RubiaceaeVernacular names Johimbe, yohimbe (En).

Yohimbe, d6marreur (Fr). Pau de Cabinda (Po)Origin and geographic distribution Pous-

mysterner johimbe is distributed from southernNigeria to Cabinda (Angola). Its occurrence inDR Congo needs to be confirmed.

Uses Young stems are used as poles in con-struction. The wood is used as fuelwood. The

bark is used in house construction, and theinner bark is utilized for making straps forhunting panniers. Young saplings are used as

I,

Puttsinystoliojohtmbe - wild

snare-trap mechanismsBark preparations are widely used as aphrodis-Iacs and stimulants. In Cameroon, for instance,the bark is popularly known as 'the Africanviagra'. In Gabon small quantities of the barkare chewed to dispel sleep, while as an aphro-disiac it is used in larger quantities or morefrequently. The bark is also used as a localanaesthetic, a hallucinogen, for the treatmentof angina, against constipation and Intestinalworms, as a performance enhancer for athletes,and to increase the clarity of singers' voices. InCongo a bark decoction is drunk for the treat-merit of pelvic pain. In Gabon the bark is usedas a fish poison.The bark is the standard source of the alkaloid

yohimbine, which is included in various Euro-pean pharmacopoeias for Its sympatholytic,hypotensive and local anaesthetic activity. Yo-himbine-based products are widely used asaphrodisiacs in the Western World, eitherthrough pharmaceutical channels or herbalmedicine markets. The most common use of

yohimbine-based drugs in western medicine isin the treatment of diabetes-related male impo-tence. Yohimbine-based products are also usedas food supplements for body weight reductionIn the United States they are recorded to beused as food supplements to substitute ariabol-IC steroids and improve athletic performanceHowever, in a study investigating the alkaloidcontent of 26 yohimbe products sold in retailhealth food outlets throughout the UnitedStates, 9 of the products appeared not to con-tain any yohimbine, 8 only contained traceamounts, and the product with the highestyohimbine content contained only 0.05%. In the

United Kingdom yohimbine-based drugs havebecome fashionable as 'herbal highs'

Production and international trade The

bark is extensively traded in local markets andexported. Most of the bark entering commercialtrade comes from Garnero0n. In 1985-1991the

annual exports of bark from Cameroon amount-ed to 286 t, of which 65% to the Netherlands,18% to Germany, 11% to Belgium/Luxembourgand 6% to France. The bark exports from Cam-eroon in the season 1997-1998 amounted to

715 t, with a value of about Us$ 600,000. How-ever, a large proportion of the exported bark isprobably not fromPousinystoliojohimbe but fromPuttsinystolio mereroceros (K. Schum. ) Pierre,which has a much lower yohimbine content.

Properties The heartwood is ochre-yellowand not clearly demarcated from the thin, yel-low sapwood. The grain is fine. The wood ismoderately heavy, with a density of 650-750kg/ms at 12% moisture content, and moderatelyhard.

The bast fibre cells are 0.7-1.6 min long, with adiameter of 22-29 prn. The bark of Pousiny-sternojohimbe contains up to 6% alkaloids. Thealkaloid content is highest in the root bark andthe bark of the hole, lower in the bark of thebranches, and very low in the leaves. Withinthe hole, the alkaloid content increases fromthe base upwards. The alkaloids present inPerusinystolio johimbe include yohimbine (10-15%), mesoyohimbine, yohimbinine, corynan-thine, alloyohimbine and ajamalicineYohimbine (also known as aphrodine, quebra-chine or corynine) is a selective inhibitor of u-2-adrenergenic receptors. At low doses it hashypertensive activity, while at higher doses itis hypotensive, through vasodilation of penph-eral vessels. Yohimbine raises heart rate and

norepinephrine levels. The use ofyohimbine asan aphrodisiac is attributed to its dilating ef-rect on the blood vessels of the sexual organs,thus increasing blood supply, while it also pro-vides an enhancement of the reflexes involved

in the control of ejaculation. Tests have shownthat treatment with yohimbine indeed resultsin increased libido and easier ejaculation. Theeffects ofyohimbine on smooth muscles favourstonus and movement of the intestine. It also

acts on o-2-adrenergenic receptors of adjpo-cytes, resulting in increased lipolysis. However,double-blind trials on the effectiveness of yo-himbine for body-weight reduction gave con-flicting results, and it is therefore unclearwhether yohimbine is effective in reducingbody weight. A wide range of adverse effects of

L.

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PAUSINYSTALIA 517

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518 TIMBERS2

the use of yohimbine have been recorded, in-cluding hypertension, mania, bronchospasm,anxiety, agitation, hallucinations, vertigo,stomach problems, headache and weakness. Itmay also Interact with antidepressants. ESPe-clany people with hypertension, prostate prob-Iems or heart diseases are warned against us-ing yohimbine-based productsAjamalicine also has vasodilating activity.Corynanthine closely resembles yohimbine, butis more active as a sympatholytic agent andless toxic. The bark also contains tannins

Adulterations and substitutes Pousiny-stolio johimbe bark is frequently adulteratedwith the bark of other Poustnystolio and Gory-nonthe spp. , which have a lower yohimbinecontent

Description Evergreen, medium-sized treeup to 30(-35) in tall; hole straight, up to 50(-60) cm in diameter, without buttresses butgrooved at base; bark easy to peel off, bitter-tasting, bark surface longitudinalIy fissured,with transverse cracks, grey to reddish brown,inner bark fibrous, pinkish, turning reddishbrown on exposure; crown compact, withbranches in whorls. Leaves in whorls of 3, SIm-

PIe; stipules 1.5-2 cm long, glabrous, caducous;petiole up to 5(-8) mm long; blade obovate oroblanceolate, (11-)13-47 cm x 5-17.5(-19) cm,base cordate, curieate or rounded, apex short-acuminate, margin often wavy, glabrous, pin-nately veined with (8-)10-20 pairs of lateralveins. Inflorescence a terminal or ax111ary pan-ICle 5-21(-30) cm x 9-15 cm, branched inwhorls of 3, with flowers in clusters at the endsof branches, main axes glabrous; peduncle 0.5-5 cm long, with 3 ridges. Flowers bisexual, reg-ular, (4-)5(-6)-meTous, fragrant; calyx consist-ing with short tube and triangular or roundedlobes c. 0.5 min long, hairy; corolla white,sometimes yellowish, red or purple, with tubeconsisting of a basal narrow part c. 0.5 minlong and an apneal bladder 1.5-2.5 min long,hairy inside, glabrous outside, lobes erect,shortly toothed, each with a linear appendage8-20(-25) mm long; stamens attached at thebase of the corolla bladder, alternating withthe lobes, sessile; ovary inferior, 2(-3)-celled,style I-2 min long, stigma 2-lobed. Fruit anoblong, compressed capsule 10-16 mm x 5-7mm, dehiscing with 4 valves, many-seeded.Seeds oblong, 8-12 min x 1.5-2.5 mm, coin-pressed, winged.

Other botanical information Perusinysto-lid comprises 5 species, and occurs in West andCentral Africa. It is closely related to Conynon-the

Puttsinystolio johimbe can be distinguishedfrom Purrsinysto!to ingcroceros by its largerleaves with shorter or no petioles. Further-more, the bark of Puttsinystoliojohimbe is tast-ing extremely bitter and is easy to peel off,whereas that of Pousinystolio mereroceros isless bitter and difficult to peel off. Differencesin slash characteristics, often mentioned asbeing distinguishing, are not reliable

Growth and development Light is neces-sary for germination and good seedling growth,and the survival rate of seedlings under aclosed canopy is very low. Growth is usuallyfast, but the hole diameter never becomes Iarg-er than 50-60 cm. Throughout its range,Puttsinystolio johimbe usually flowers in Au-

' gust-February, and bears fruits in September-March. Pollination is probably by insects.Large amounts of seeds are produced each year.The seeds are dispersed by wind. Pousinystoliojohimbe coppices wellfrom stumps, producingstrong, highly phototropic shoots

Ecology Pousinystolio johimbe occurs inevergreen lowland forest, in primary as well assecondary forest, at relatively low densities.

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Propagation and planting Vegetativepropagation trials with single-node leafy cut-tings have given good results. Propagation withseeds is also done, but seeds are difficult tocorrect. Furthermore, the survival rate of seed-lings was very low in experiments. Wildlingscollected from the forest showed poor survivaland low growth rates

Management Although Pousinystolio jo-him be is often stated to be common, inventoriesin Gamero0n and Equatorial Guinea in the1990s indicated that there are on average only15 plants/ha with a diameter over I cm and 4trees/ha with a diameter over 10 cm.

The World Agroforestry Centre (ICRAF) hasinitiated a research programme to Investlgatethe domestication potential of Puttsirtysto!tojohimbe and its possible inclusion in agrofor-estry systems. Initial results have shown thatit will readily grow in association with annualfood crops.

Diseases and pests Removal of large barksurfaces can lead to stem borers penetratingthe unprotected stem and killing the tree.

Harvesting Trees are harvested when theyhave reached a hole diameter of about 10 cm

As yohimbine levels are highest in the rainyseason, harvesting is a seasonal activity. Thebark is usually stripped after the tree has beenfelled for this purpose, making exploitationunsustainable. It would be better to harvest

bark strips from standing trees, which willstayalive and renew the bark in 2 years, However,harvesters often claim that the tree will die

anyway after removal of part of the bark, be-cause of stem borer attacks. Although a permitis required for harvesting the bark in Came-roon, it is often done by local people who arepaid for delivery to contractors. The bark mayalso be harvested by employees of logging coin-panies after the felling of timber trees in an

Handling after harvest The bark is mar-keted in dried flat or curved pieces up to I inlong, or ground.

Genetic resources The high demand formedicines based on the bark of Puttsinystoliojohimbe has led to over-exploitation and maylead to local scarcity of the species, if not long-term endangermerit.

Prospects The wood is of good quality, butthe relatively small hole diameters limit itsusability. Nevertheless, it has been suggestedas a suitable substitute for nayo iuorensisA. Chev. Probably, however, Pousinystolio jo-him be will remain more Important as a source

of yohimbine and local medicine than as asource of timber

The high demand for medicines based on thebark has led to over-exploitation of naturalpopulations. The tree is easily vegetativeIypropagated, however, and may have potentialfor inclusion in agroforestry systems, whichcould help to relieve the pressure on naturalpopulations. For this purpose, it has been rec-ommended to carry out studies on the breedingsystem (the presence of self-incompatibility) ofPousinystolio johimbe, and on seed dormancy,seed germination and seedling survival.Care should be taken with the use of yohim-bine-based products for medicinal purposes, asadverse side effects have been recorded. As the

content ofyohimbine and other alkaloids in thebark may vary widely, the use of crude prod-ucts is even more dangerous than that of prod-ucts with a known amount of yohimbineTherefore, yohimbine-based products should betaken cautiously and only under supervision ofa specialist, and not as self-medication.

Major references Burki11, 1997; Ernst &Pittler, 1998; Hall6, 1966; Hostettmann at al. ,2000; Pousset, 2004; Stoffelen, Robbrecht &Sinets, 1996; Sunderland at a1. , 1999; Sunder-land at a1. , 2004; Tchoundjeu at a1. , 2004;Vivien & Faure, 1985.

Other references Betz & White, 1995;Cunningham, 1997; de Saint-Anbin, 1963;Goetz, 2006; Hepper & Keay, 1963; Keay, 1989;Keay, Onochie & Stanfield, 1964; MissouriBotanical Garden, undated; Musa, 2005; Neu-winger, 2000; Orwa at a1. , 2009; Paris &Letouzey, 1960; Pittler & ETnst, 2003; Pittler,Schmidt & ETnst, 2005; Raponda-Walker &Sillans, 1961; Razafimandimbison & Bremer,2002; Sunderland, Tchoundjeu & Ngo-Mpeck,2000; Tharakan & Manyam, 2005; Vani &Giardina, 2002; Zanolari at a1. , 2003

Sources of illustration Hall6, 1966; White& Ahemethy, 1997; Wilks & Issemb6, 2000.

Authors R. B. Jiofack Tafokou

PADSiwSTALIA 519

area

FAUSllTYSTALIALANE-POOLEl(Hutch. )Hutch. ex Lane-Poole

Protologue A list of trees, shrubs, herbsand climbers of Sierra Leone: 74 (1916)

Family RubiaceaeOrigin and geographic distribution Putts-

mystolio lonerpoolei occurs from Sierra Leoneto Gabon, eastern DR Congo and Rwanda.

Uses The wood is used for house posts and

520 TIMBERS2

I.

PuusLnystolio lone-poolei- wild

Interior joinery. It is suitable for furniture andveneer. The strong and flexible branches areused for making traps. In traditional medicinein Liberia, the pounded bark is used as a poul-tice for the treatment ofyaws, scabies and itchThe rootis an ingredient of arrow poison

Properties The heartwood is pale brown topinkish, turning pale olive on exposure, and isnot clearly demarcated from the sapwood. Thegrain is straight to irregular, texture fine. Thewood is moderately heavy, with a density ofabout 720 kg/in" at 12% moisture content. It ismoderately hard. At 12% moisture content, themodulus of rupture is 93 N/mm2 modulus ofelasticity 13,900 N/mm2, compression parallelto grain 65 N/min2, Janka side hardness 6540N and Janka end hardness 7560 N. The woodworks and finishes well. It is recorded to bedurable, but it has also been reported to besusceptible to Insect attacks

Botany Evergreen, medium-sized to fairlylarge tree up to 40 in tall; hole branchless forup to 15 in, straight, cylindrical, up to 120 cmin diameter, with short buttresses or withoutbuttresses; bark surface scaly, the scales Ieav-ing rounded pits, grey-brown, inner bark soft,fibrous, pinkish yellow, darkening on exposure,bitter-tasting. Leaves decussately opposite,simple and entire; stipules triangular, cadu-cous; petiole up to 3 cm long; blade elliptical,oblong or obovate, 7-20(-24) cm x 3-9.5 cm,base curieate to rounded, apex short-acumi-nate, leathery, glabrous, pinnately veined with9-14 pairs of lateral veins. Inflorescence a ter-minal panicle, 10-25 cm x 15-24 cm, withflowers in clusters at the ends of branchesFlowers bisexual, regular, (4-)5-merous, fra-

grant; calyx with short tube and lobes, hairyinside and outside; corolla white, sometimespinkish, tube with narrow basal part up to Imm long and apical bladder 2-2.5 mm long,hairy inside, glabrous outside, each lobe with alinear appendage 7-15 mm long; stamens at-tached at base of the corolla bladder, alternat-ing with the lobes, sessile; ovary inferior, 2-celled, style c. 1.5 min long, stigma 2-lobed.Fruit an oblong, compressed capsule, 8-20 minlong, dehiscent with 4 valves, many-seeded.Seeds 5-9 min x I-2.5 min, wingedIn Liberia Poustnysto!to lonerpoolei flowers inthe rainy seasonPousinystolio comprises 5 species, and occursin West and Central Africa. It is closely relatedto Gorynonthe. POLLStnystolio lonerpoolei has astrikingly disjunct area of distribution. Twosubspecies are distinguished. Subsp. lone-poolei occurs from Sierra Leone to Ghana.Subsp. itunense (De Wild. ) Stoffelen & Robbr.(synonym: Perusinystolio itunense De Wild. )has slightly shorter corolla lobe appendagesand slightly smaller fruits and seeds and isfound in eastern DR Congo and Rwanda, withone collection from Gabon

Ecology Pousinysto!to lone. poolei occurs inevergreen forest, usually at altitudes above 500in, up to 2000 in altitude.

Genetic resources and breeding It isunclear to what extent Puttstrtystolio lone-pooleL Is threatened with genetic erosion. InSierra Leone and Liberia it is recorded to becommon, but in Ghana it is rare

Prospects Few reports exist on the use ofthe timber of Pawsinystolio lone-poolei, alt-hough research indicated it is of fairly goodquality. The timber is unlikely to become moreimportant in the future.

Major references Abbiw, 1990; Burkill,1997; Kryn & Fobes, 1959; Stoffelen, Robbrecht& Sinets, 1996; Takahashi, 1978

Other references Hall6, 1966; Hawthorne,1995; Hawthorne & Gyakari, 2006; Hepper &Keay, 1963; Irvine, 1961; Neuwinger, 1998a;Neuwinger, 2000; Razafimandimbison & Bre-mer, 2002; Savill& Fox, 1967; Voorhoeve, 1979

Authors M. Brink

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FAUSINYSTALIAMACROCERAS (K. Schum. )Pierre ex Beille

Protologue Act. SOC. Linn. Bordeaux 61:130 (1906).

Family Rubiaceae

Vernacular names Falsejohimbe (En).Origin and geographic distribution Pous-

mystolio ingcroceros is distributed from Nige-ria eastward to DR Congo and southward toCabinda (MgOla).

Uses The wood is valued for furniture andcabinet work. In Cabinda (Angola)it is used forhouse construction and flooring. The wood issuitable for joinery, interior trim, ship build-ing, vehicle bodies, ladders, sporting goods,toys, novelties, agriculturallinPIements, han-dles, musical instruments, boxes, crates, preci-SIon equipment, matches, carvings, turnery,pattern making, veneer and plywood. In Nige-ria it is used as fuelwood

The bark is widely used as an aphrodisiac andstimulant. In Cameroon, for instance, the pul-verized bark is eaten against sexual weaknessIn Congo the bark is used to stave off sleep,and sap from the bark is applied in an enemaas a vermifuge and topicaUy for the treatmentof ringworm and other fungal skin Infections.

Properties The heartwood is pale yellow topale pink, with irregular stripes occasionallypresent; it is not clearly demarcated from thesapwood. The grain is straight, texture fineand even. The wood is lustrous and contains areddish brown gumThe wood is medium-weight, with a density ofabout 700 kg/in' at 12% moisture content. It airdries rapidly without serious deformation. Therates of shrinkage are rather high, from greento oven dry 4.7% radial and 10.7% tangentialOnce dry, the wood is moderately stable inservice. At 12% moisture content, the modulusof rupture is 151 N/min2, modulus of elasticity10,900 N/mm2, compression parallel to grain 65N/mm2 shear 8 N/min2, cleavage 12.5 N/min

and Chalais-Meudon side hardness 3.8

The wood saws well, works easily with antools,planes to a smooth surface, and finishes well. Ittakes and holds nails and screws well, andglues satisfactorily. The peeling properties aregood, but splitting of veneer may occur duringdrying. For slicing, radial sections are recoin-mended to reduce defects caused by the ratherhigh shrinkage ratesThe durability of the wood is classified as inod-erate, although it is not susceptible to attacksby termites and marine borers. The sapwood isnot susceptible to Lyctus borer attack. The heart-wood is resistant to impregnation with pre-servatives, the sapwood moderately resistantThe bark contains the indole alkaloid yohim-bine, but in smaller quantities than in the barkof FOMstrtysto!to johimbe (K. Schum. ) Pierre exBeille, which has led to Puttsinystolio ina-croceros sometimes being called 'false yohim-be'. The bark has been recorded to contain only0.02-0.32% indole alkaloids and more of theinactive alkaloid yohimbinine than of yohim-bine. However, bark from Congo was found tocontain 4% alkaloids, of which 60-65% yohim-bine. Other alkaloids isolated from this bark

were corynanthidine (u-yohimbine), p-yohim-bine, aimalicine, corynanthine, and the Ievo-rotatory isomer of calycanthineThe bark also contains saponins and tannins.Methanolic extracts of the stem bark haveshown in-vivo anti-ulcerogenic, antidiarrhoeal,analgesic and anti-inflammatory effects in rats.

Botany Evergreen, medium-sized to fairlylarge tree up to 40 in tall; hole branchless forup to 18 in, straight, cylindrical, up to 90 cm indiameter, without buttresses, but with groovedbase; bark surface often fissured, grey tobrown, inner bark fibrous, creamy white, turn-ing reddish brown upon exposure. Leaves de-cussately opposite, simple and entire; stipulescaducous; petiole (3-)7-25 min long; blade obo-vate to elliptical, 6.5-47 cm x 2.5-17.5 cm, basecurieate to rounded, apex short-acuminate,papery or thinly leathery, glabrous, pinnatelyveined with 5-16 pairs of lateral veins. Inno-rescence an axillary panicle, 9-15 cm x 5-14

with flowers in clusters at the ends ofcm

branches. Flowers bisexual, regular, (4-)5-6-merous, fragrant; pedicel up to 0.5 mm long;calyx with short tube up to 0.5 min long andtriangular to rounded lobes 05-I mm long,acute, glabrous or somewhat hairy outside,densely hairy inside; corolla white, sometimespink, glabrous outside, hairy inside, tube withbasal narrow part 0.5-I mm long and apical

PAUSINYSTAi, IA 521

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FOMsinystolio ingcroceros - wild

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522 TIMBERS2

bladder I-2 mm long, lobes erect, each with alinear appendage 6-12.5 min long; stamensattached at base of the corolla bladder, alter-nating with the lobes, sessile; ovary inferior, 2-celled, glabrous, style I-2 mm long, stigma 2-lobed. Fruit an oblong, compressed capsule,10-20 min x 5-7 min, 4-valved, many-seeded.Seeds 5-14 mm x I-2 min, winged.In Nigeria Pousinysto!to inocroceros flowers inMay-September, and fruits in January-March.Puttsinystolio comprises 5 species, and occursin West and Central Africa. It is closely relatedto Gory, tonthe. Puttsirtystolio inderoceros canbe distinguished from Puttstnystu!adjohtmbe byits smaller leaves with longer petioles. Fur-thermore, the bark of Pousinystolio johimbe istasting extremely bitter and is easy to peel off,whereas that of Pousinystolio macroceros isless bitter and difficult to peel off. Differencesin slash characteristics, often mentioned asbeing distinguishing, are not reliable.

Ecology Pousinystolio ingcroceros occurs inlowland evergreen forest, primary as well assecondary, and also in gallery forest

Genetic resources and breeding In viewof its fairly wide distribution and local abun-dance, it is unlikely that Pousinysto!to ina-croceros Is threatened with genetic erosion

Prospects Although the wood is recorded tobe suitable for a wide range of purposes, evi-dence of its use is scarce. Its role as a timber isunlikely to become more important in the fu-ture. The bark of Pousinystolio macroceros isused as an aphrodisiac and stimulant as a sub-stitute or adulterant of that of Perusinystoliojohimbe, but contains lower amounts of themain active compound 6, objmbine).

Major references Bolza & Keating, 1972;Burki11, 1997; Leboeuf at a1. , 1981; Stoffelen,Robbrecht & Sinets, 1996; Sunderland at al. ,2004.

Other references Hall6, 1966; Thenyen,Okoegwale & Mensah, 2009;'Keay, 1989; Neu-winger, 2000; Nwafor, Jacks & Ekanem, 2007;Nwafor at a1. , 2005; Sallenave, 1964; Sunder-land at a1. , 1999; Takahashi, 1978; Vivien &Faure, 1985.

Authors M. Brink

Origin and geographic distribution Pous-mystolio taibotii occurs in Nigeria and Came-

Uses The wood is suitable for construction,flooring, joinery, Interior trim, mine props, shipbuilding, vehicle bodies, furniture, cabinetwork, ladders, sporting goods, agricultural jin-PIements, handles, musical instruments, toys,novelties, precision equipment, boxes, crates,matches, carvings, turnery, draining boards,pattern making, veneer and plywoodIn traditional medicine a bark extract is drunkfor the treatment of fever. In Cameroon thepulverized bark is eaten against sexual asthe-nia. The fruit is eaten to cure dysentery andapplied as a topical treatment for tumours.

Properties The heartwood is pale yellow,turning yellow-brown or olive brown upon ex-POSure; it is not clearly demarcated from thesapwood. The grain is usually straight andregular, texture very fine. The wood is Ius-trous.

The wood is medium-weight, with a density ofabout 680 kg/in" at 12% moisture content. It airdries rapidly with no serious degrade. Therates of shrinkage are moderate, from green tooven dry 2.1-3.0% radial and 3.6-5.0% tangen-tial. At 12% moisture content, the modulus ofrupture Is 1/8 N/min2, modulus of elasticity14,100 N/min2, compression parallelto grain 57N/min2, cleavage 19.5 N/min and Chalais-Meudon side hardness 4.7

The wood is easy to saw and works well withboth hand and machine tools. It planes to alustrous surface. It glues well, takes nails andscrews satisfactorily and has good peeling andslicing propertiesThe wood is durable, being resistant to attacks

roon.

FAUSINYSTALIATALBOTllWernham

Protologue Rendle, Cat. PI. Oban: 40 (1913)Family RubiaceaeSynonyms Pousinysto!to sunkeyi Hutch. &

Dalzie1(1931).

I.

Pousinystolio to160tii- wild

L. .

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by termites and marine borers. The sapwood isnot susceptible to Lyetus borer attack. Theheartwood is resistant to Impregnation withpreservatives, the sapwood moderately re-SIStant

Description Evergreen, small to medium-sized tree up to 30 in tall; hole straight andcylindrical, up to 90 cm in diameter, with but-tresses; bark surface smooth, grey, Inner barksoft, cream or pink, turning brownish uponexposure. Leaves usually decussately opposite,simple and entire; stipules triangular, cadu-cous; petiole I-3 cm long; blade elliptical toovate, 9-25 cm X 4-11.5 cm, base curieate, apexshort-acuminate, leathery, glabrous, pinnatelyveined with 8-13 pairs of lateral veins. Inno-rescence an axillary panicle, (5-)9-15(-20) cmlong, with flowers in clusters at the ends ofbranches, axes more or less hairy. Flowers bi-sexual, regular, (4-)5-merous, almost sessile;calyx with short tube and triangular lobes,densely hairy outside and inside; corolla whiteto red, glabrous outside, densely hairy inside,tube with a basal narrow part c. 0.5 mm longand an apical bladder c. 1.5 mm long, lobeseach with a linear appendage (4-)5-8 mm long;stamens inserted in throat of the corolla tube,alternating with the lobes; ovary inferior, 2-celled, style c. 1.5 mm long, stigma 2-lobedFruit an oblong, compressed capsule, 7-23 mmlong, dehiscing with 4 valves, many-seeded.Seeds narrowly ellipsoid, 9-11 min x 2-2.5mm, winged

Other botanical information Pousinysto-lid comprises 5 species, and occurs in West andCentral Africa. It is closely related to Gorynon-the.

Puttsinystolio to160tii may have been confusedin the literature with other POWsinystolio spp. ,and therefore the information on wood proper-ties may partly refer to other species.

Anatomy Wood-anatointcal description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: (4: wood semi-ring-porous); 5: wood diffuse-porous; (9: vessels ex-clusively solitary (90% or more)); (12: solitaryvessel outline angular); 13: simple perforationplates; 22: intervessel pits alternate; 23: shapeof alternate pits polygonal; 24: intervessel pitsminute (S 4 prn); 25: intervessel pits small(4-7urn); 29: vestured pits; 30: vessel-ray pits withdistinct borders; similar to intervessel pits insize and shape throughout the ray cell; 41:mean tangential diameter of vessellumina 50-100 pin; 49: 40-100 vessels per square minime-

tre. Tracheids and fibres: 62: fibres with dis-

tinctly bordered pits; 63: fibre pits common inboth radial and tangential walls; 66: non-septate fibres present; 69: fibres thin- to thick-walled; (70: fibres very thick-walled). Axialparenchyma: 76: axial parenchyma diffuse; 77:axial parenchyma diffuse-in-aggregates; 78:axial parenchyma scanty paratrachea1; 93:eight (5-8) cells per parenchyma strand. Rays:97: ray width I-3 cells; (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; 1/6: 2 12 rays per

PELLEGRiNIODENDRON 523

min

(S. N'Danikou, P. Baas & H. Beeckman)Growth and development In Nigeria flow-

ering is in March-April, and fruiting in De-cember-April.

Ecology Puttsinysto!to to160tii occurs inrainforest, but also in secondary vegetation

Genetic resources Pousinystolio tatbotiihas a limited distribution area, but does notseem to be threatened by genetic erosion.

Prospects Although the wood of FOMsinysto-jig to160tiiis of good quality and durable, andthe range of potential uses is wide, It seems notmuch used at present. Too little information isavailable on the prevalence, growth character-1stics and management aspects of the speciesfor a good assessment of its prospects

Major references Bolza & Keating, 1972;Burki11, 1997; Stoffelen, Robbrecht & Sinets,1996; Takahashi, 1978

Other references GBIF, 2011; Hepper &Keay, 1963; Keay, 1989; Keay, Onochie & Stan-field, 1964; Missouri Botanical Garden, undat-ed; Neuwinger, 2000; Razafimandimbison &Bremer, 2002

Authors E. E. Ewudzie, J. R. Cobbinah, SBritwumAcquah & E. A. Obeng

FELLEGRINIODENDRONDIPHYLLUM (Harms)J. Leonard

Protologue Bull. Jard. Bot. Etat 25(2): 203(1955)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Synonyms Mocro!objum diphyllum Harms(1901).

Vernacular names Faux copalier(Fr)Origin and geographic distribution Pelle-

griniode, Idron diphyllum occurs in Liberia,C6te d'TVoire, Ghana, Garnero0n, Equatorial

524 TIMBERS2

I.

Penegriniodendron diphyllum - wild

Guinea and Gabon

Uses In Ghana the wood is used for door

frames and in Gabon for joinery. It is suitablefor cabinet work.

Properties The heartwood is brown to red-dish brown, often marbled. The texture is inod-erately fine. The wood is quite heavy and hard.

Botany Smallto medium-sized tree up to 25in tall; hole often low branching, up to at least30 cm in diameter, often with adventitiousshoots at base; bark surface smooth, finely fis-sured, with lenticels, greyish brown, inner barkfibrous, reddish; crown rounded, with droopingbranches. Leaves alternate, panpinnatelycompound with I pair of leaflets; stipules tri-angular, c. 3 min long; petiole 3-5 min long;petiolules 2-3 min long, with short stipels atbase; leaflets elliptical to oblong or obovate,10-24 cm x 3-10 cm, asymmetrical at base,short-acuminate at apex with acumen oftencontorted or rolled up, leathery, glabrous, withsome glands near margins, pinnately veinedwith c. 10 pairs of lateral veins. Inflorescence aterminal, drooping panicle up to 10 cm long,with raceme-like, densely flowered sidebranches 2-3 cm long, with conspicuous scarsof fallen flowers and bracts, usually glabrous.Flowers bisexual, zygomorphic, with 2 pinkishbracteoles up to I cm long at base; pedice1 6-8mm long; sepals 5, lanceolate, 3-4.5 mm long,slightly fused at base; petals 5, free, unequal,upper one up to 1.5 cm long, 2-lobed, whitish,other ones resembling the sepals; stamens 3,free, I-2 cm long, reddish, with a collar ofsmall rudimentary stamens at base; ovary su-perlor, oblong, hairy at margins, I-celled, stylec. 1.5 cm long. Fruit an elliptical to obovoid,

flattened pod c. 10 cm x 4 cm, pointed at apex,smooth, glabrous, reddish brown, dehiscingwith 2 spiralling thinly woody valves, I-3-seeded. Seeds rectangular to elliptical, flat-tened, up to 3 cm x 2 cm, glossy brown, seedcoat thin. Seedling with epigeal germination;hypocoty1 1.5-4 cm long, epicoty1 3.5-7 cmlong; cotyledons thick and fleshy, rectangular-elliptical, 2-2.5 cm long; first 2 leaves stronglyreduced to opposite scales, subsequent leavesalternate, with I pair of leaflets.Young leaves are often bright pink. In WestAfrica Penegriniodendron diphy!Ium flowers inSeptember-December and fruits mature about5 months after flowering. The presence of ec-tomycorrhizae has been recorded.Fellegriniodendron comprises a single speciesand seems to be related to Gilbertiodendron.

Ecology Penegriniodendron diphyllum usu-ally occurs in the understorey of lowland rain-forest, often on humid localities along streamsand in moist gullies

Management Natural regeneration is oftenabundant, and the seedlings are shade toIer-ant. Germination starts 1.5-4 weeks after sow-Ing, with a germination rate of 70-80%. Ma-ture trees are often found in small, densegroups.

Genetic resources and breeding Fellegri-ittodendron diphy!him is not threatened be-cause it Is fairly wldespread and locally coin-mon with abundant regeneration. However, inGhana it has been rated as a gold star species,meaning that there Is some inescapable re-sponsibility for maintaining this species be-cause it has a very restricted distribution inGhana.

Prospects Penegriitiodendron diphy!!urn winremain of little importance as a timber treebecause the logs are usually too small and toopoorly shaped.

Major references Aubr6ville, 1959b; Aubr6-vine, 1970; de Koning, 1983; ITvine, 1961;Raponda-Walker & Sillans, 1961

Other references Aubr6ville, 1968; BurkiU,1995; Diabate at a1. , 2005; Hawthorne, 1995;Hawthorne & Jongkind, 2006; Lewis at al. ,2005; Normand & Paquis, 1976; Voorhoeve,1979; White & Ahemethy, 1997.

Authors R. H. M. J. Lemmens

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PEMPHiSACiDULA JR. Forst. & G. Forst.

Protologue Char. gen. PI. , ed. L 34 (1775)Family LythraceaeChromosome number2n= 32

Vernacular names Small-leaved mangrove(En). Bois matelot (Fr). inIalamba kike (Sw)

Origin and geographic distribution Pem-phis octdulo is found along the east coast ofAfrica from Somalia to Mozambique and on theIndian Ocean islands. It is further found on

shores of tropical Asia and Australia, and onislands of the Pacific Ocean.

Uses The hard and durable wood ofPemphisoctdulo is used for house posts, fence posts,tool handles, walking-sticks, domestic imple-merits, turnery, anchors, boat nails and pestlesIt is also used for firewood and charcoal pro-duction. Along the East African coast the barkhas been used for tanning.In Asia Pemphts acidulo is occasionally plant-ed as an ornamental, whereas the acid tastingleaves are eaten raw or boiled as a vegetable.In Vanuatu a filtered infusion of the bark is

used as an abortifacient, and in Indonesia thebark is used to treat stomatitis.

Production and international trade Be-

cause of the smallsize and the poor form of thebole of Pemphis octdulo, as well as its limitedsupply, the use of the wood is on a small andlocal scale only. There is significant trade inseeds and live plants, especially in Asia forbonsai.

Properties The heartwood of Feinphis oci-du!a is reddish brown to dark reddish brown,turning dark brown upon exposure, and isclearly differentiated from the pale, about I cmwide sapwood. The grain is interlocked, texture

very fine. The wood Is lustrous.The wood is very heavy, with a density of1100-1210 kg/ina at 15% moisture content. Itair dries well with little checking and warping.The rates of shrinkage upon air drying aremoderate, from green to oven dry 5.4% radialand 8.5% tangential for wood from the Philip-pines. The wood is very hard and very strongIt is very difficult to work, but with care ittakes a high finish. It is very durable, beingresistant to dry-wood termite and Lyetus at-tacks.

The bark contains 19-43% tannin. In-vitro

tests of the bark showed an increased activityon the amplitude and frequency of uterine con-tractions, which confirms the traditional use inVanuata as an abortifacient. Bark extracts

were found to have antibacterial, antioxidantand topoisomerase I inhibitor activities. Fourgalloyl navonol glycosides with antioxidantactivity have been isolated from leafextracts

Botany Shrub to smalltree up to 11 in tall;hole usually gnarled and much branched; barksurface dark grey, rough, Teticulately flaking;twigs angular, densely hairy, nodes thickenedwith conspicuous leaf scars. Leaves opposite,simple and entire, sessile; stipules absent;blade obovate-oblanceolate to linear-lanceolate,I-3.5 cm x 3-13 min, curieate at base, acute toobtuse at apex, leathery to fleshy, silky-haired,with only inIdrib distinct. Flowers solitary inleaf axils, bisexual, regular, 6-merous, hetero-stylous, sweet-scented; pedice1 5-13 mm long;calyx bell-shaped, greenish, 12-ribbed, hairy,with short lobes; petals inserted on calyx tube,c. 5 min x 4 mm, white or pink, early caducous;stamens 12, inserted slightly below the middleof the calyx tube, 6 longer ones alternatingwith 6 shorter or all 12 nearly equal; ovarysuperior, globose, c. 2 min long, 3-celled, styleshort or elongate. Fruit a globose capsule 4-6min long, purplish green, the upper part open-ing by a lid, many-seeded. Seeds c. 3 mm x 2mm, with thick marginal wing, reddishPemphis comprises a single species since Pem-phis inoddgoscoriensis (Baker) Koehne, en-demicto Madagascar, has been transferred to aseparate genus Koehnerio. The development ofthe tree corresponds to Attims' architecturalmodel, characterized by axes with continuousgrowth, differentiated into a monopodialtrunkand equivalent branches. Flowers show beter-odistyly, with stamens and styles in 2 lengthclasses. Flowers are self-incompatible

Ecology Pemphis octdulu occurs at thelandward side of mangroves. It may also be

PEMPHiS 525

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Pemphis acidulo- coild

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526 TIMBERS2

found exposed to the ocean, but then oftenpartly sheltered by rocky shore, and it is occa-SIonally even found below the high-watermark

Genetic resources and breeding AlthoughPemphis octdulo is extremely widespread, it isuncommon along the east coast of mainlandAfrica, and the most important threat is to itshabitat, i. e. mangrove vegetation, of which thearea is rapidly diminishing

Prospects Increase of the use of Pemphisoddu!a wood is unlikely in tropical Africa, butthe wood will remain of interest for specialtyuses

Major references Graham, Tobe & Baas,1986; Furnobasuki & Irawan, 1998; Tomlinson,1986; Verdeourt, 1994.

Other references Bourdy at a1. , 1996;Coode, 1993; GIIbert & Thulin, 1993; Gill &Kyauka, 1977; Greenway, 1941; Hardjit0, 2007;Masuda at a1. , 2001; MMG, undated

Authors C. H. BOSch

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

I.

PETERSIANTHUSMACROCARPUS (P. Beauv. )Liben

Protologue Bull. Jard. Bot. Etat 38: 207(1968)

Family LecythidaceaeChromosome number2n= 52

Synonyms Combretodendron o17icunum(WeIw. ex Benth. ) Exe11(1930), Combretoden-dron mererocorpum (P. Beauv. ) Keay (1958)

Vernacular names Stinkwood tree, soaptree (En).

Origin and geographic distribution Peter-signtht, s distributed frominOCrOCOrpuS Is

Guinea eastward to the Central African Repub.11c and southward to DR Congo and northernAngola

Uses The wood of Petersionthus ingcrocor-

pus is used for construction, carpentry, furni-ture, canoes, mortars, tool handles, sliced ve-neer and plywood. It is suitable for flooring,mine props, vehicle bodies, railway sleepers,sporting goods, toys, novelties, agriculturalimplements and draining boards. It is valuedas firewood and for charcoal production.In C6te d'TVoire the bark is used as a purgativeand laxative and is considered abortifacient. In

Ghana bark decoctions are taken as expecto-rant, and in DR Congo as cholagogue and as acure for stomach pain, pneumonia and jaun-

Petersionthus inderocorpus - wild

dice. In Equatorial Guinea the bark is adminis-tered as arithelminthic and to cure cough. Barkdecoctions are widely used to clean wounds andto promote wound healing. Hot bark is appliedto the skin against muscle soreness. In Gabonleaf decoctions have been used orally and byenema for the treatment of haemorrhoids, con-stipation, paralysis and ulcerative wounds. InCameroon the leaves are used as a medicine for

dysentery. Edible caterpillars, which feed onthe leaves, are collected and eaten after roast.ing or boiling

Production and international trade Wood

of Petersionthus inderocorpus, known as 'essia','esia', 'aba16', 'abing' and 'owewe' is mainlyused locally. Ghana has been exporting consid-erable volumes, predominantly as sliced ve-neer, but in 1998 still about 2350 in3 of logswere exported.

Properties The heartwood is reddish brown,darkening upon exposure and often speckledwith darker streaks, and is distinctly demar-cated from the 4-10 cm thick, yenowish whitesapwood. The grain is straight to interlocked,texture moderately coarse to fine. Quarter-sawn wood shows a nice ray figure. The greenwood produces an extremely unpleasant smellwhen cut, butthis disappears upon drying.The wood is medium-weight to heavy, with adensity of 630-920 kg/in3 at 12% moisture con-tent, and moderately hard. Drying needs to bedone slowly to avoid splitting, checking, warp-ing or even collapse. The rates of shrinkage arehigh, from green to oven dry 4.3-6.8% radialand 9.1-11.6% tangential. It is recommendedto quarter-saw logs before drying. Once dry,the wood is moderately stable to unstable in

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service

At 12% moisture content, the modulus of rup-ture is (76-)112-187 N/min2, modulus of elas-ticity 7940-19,300 N/mm2, compression paral-Iel to grain 43-75 N/min2, shear 8.5-14.5N/min2, cleavage 13-37 N/min, Janka sidehardness 6360-9690 N, Janka end hardness7070-10,400 N and Chalais-Meudon sidehardness 2.9-60.

The wood usually saws well, but sometimeswith some difficulty because of high densityand the presence of interlocked grain resultingin a tendency of over-heating saw blades,thereby charring sawn surfaces. A reducedcutting angle of 20' or less is recommended inplaning to avoid picking up of grain at surfac-es. The wood has a tendency to char in boringand chiselling operations. It may split uponnailing and screwing, and pre-boring is ad-vised. It stains and polishes well when a filleris used. The bending properties are poor. Theheartwood is moderately to fairly durable, withcontradictory reports on resistance to termltesand borers; it is quite resistant to fungal at-tacks. The sapwood is fairly resistant to Lyctusattack, but susceptible to blue stain. Theheartwood is resistant to impregnation by pre-setvatives, butthe sapwood is permeableThe wood contains 39.5-40.5% cellulose, 29-30% jignin, 14.5-15.5% pentosan, 0.4-0.6% ashand little silica. The solubility is 6.2-9.6% inalcohol-benzene, 2.1-3.3% in hot water and18.3% in a I% NaOH solution

The bark contains high levels of sterols, tan-rims and saponosides, and traces of navonoidsBark extracts have shown strong filaricidaleffect against Log 100. High concentrations ofbark extract act on smooth muscles, circula-tion, heart-muscles and interferes with theoestrus cycle, conception and pregnancy. Anethanolic leaf extract showed antiproliferativeactivity on human colon cancer cells (IC50 = 17F1g/in"

Adulterations and substitutes In Ghana

Petersiunthus inderocorpus wood has been sug-gested as a good substitute for the woods ofDopoco gumeensis Mtill. Arg. , negheme!IQheeke!it (Achev. ) Roberty and Diospyros kg-merunensis GIIrke which have become scarce

as a result of exploitation.Description Deciduous medium-sized to

large tree up to 45 in tall; hole branchless forup to 25 in, usually straight and cylindrical, upto 130 cm in diameter, thickened and slightlyfluted at base or with small buttresses; barksurface longitudinalIy fissured and becoming

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PETERSiANTHUS 527

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Petersionthus inderocorpus - I, 60se of 601e, ' 2,flowering twig, . 3, flower, 4, fruits.Redrown grid odopted by W Wessel-ByGrid

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scaly, medium to dark brown, inner bark fi-brous, cream-coloured to yellow-orange or pink-brown, with an unpleasant smell; crownrounded, fairly dense; twigs finely hairy, be-coming glabrous. Leaves arranged spiralIy,crowded towards the ends of twigs, simple;stipules absent; petiole 0.5-2.5 cm long, nar-rowly winged; blade elliptical or obovate, 6-16cm x 47 cm, base curieate, apex acute to acu-inmate, margin entire to slightly wavy orslightly toothed, papery, nearly glabrous, pin-nately veined with 6-12 pairs of lateral veinsInflorescence a terminal panicle or raceme upto 10 cm long, short-hairy. Flowers bisexual,regular, 4-merous; pedice1 1.5-2 mm long,jointed below the middle; sepals broadly ovate,c. 2 min x 2 mm, attached to winged receptacle;petals broadly elliptical, c. 7 min x 7 min, whiteto pale green, soon caducous; stamens numer-ous, fused at base, c. I cm long, early caducous;ovary inferior, 2-celled, style straight, c. I cmlong. Fruit a spindle-shaped nut, with 4 paperywings up to 7 cm x 3.5 cm, indehiscent, I-seeded. Seeds spindle-shaped, I-1.5 cm long.Seedling with epigeal germination; hypocotylc.

,, I",'*, a, ,*~;-, L'

528 TIMBERS2

5 cm long, epicotyl very short; cotyledons leafy,elliptical to ovate, I-1.5 cm long, erect; leavesarranged spiralIy, nearly sessile, margins fine-Iy toothed

Other botanical information Petersion-thus comprises only 2 species, one in Africa andthe other endemic to the Philippines.

Anatomy Wood-anatointcal description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 27: intervessel pits large G 10Urn); 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; 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); 33: vessel-ray pits of twodistinct sizes or types in the same ray cell; 42mean tangential diameter of vessellumina100-200 pin; 47: 5-20 vessels per square inn-limetre; 56: tyloses common. Tracheids andfibres: 61: fibres with simple to minutely bor-dered pits; 66: non-septate fibres present; 70:fibres very thick-walled. Axial parenchyma:(76: axial parenchyma diffuse); 79: axial paren-chyma vasicentric; 80: axial parenchyma all-form; 82: axial parenchyma winged-anform; 83axial parenchyma confluent; 92: four (3-4) cellsper parenchyma strand; 93: eight (5-8) cellsper parenchyma 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 min); 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; 110: sheathcells present; 113: disjunctive ray parenchymacell walls present; 1/5: 4/2 rays per mm.Mineral Inclusions: 136: prismatic crystalspresent; (137: prismatic crystals in uprightand/or square ray cells); 138: prismatic crystalsin procumbent ray cells.(L. Awoyemi, A. A. Oteng-Amoako & P. Baas)

Growth and development Growth of seed-tings Is slow; after 9 months they reach about11 cm tall. When planted in full sun, seedlingswere only 75 cm tall after 5 years, but in inod-erate shade they reached about I in tall4 yearsafter plantingAlthough Petersionthus inocrocorpus is consid-ered to be an indicator of disturbances in the

forest, seedlings have been reported to toleratesome shade and are most common in small

forest gaps; saplings can be found in small aswellas larger gapsTrees are leaness for a short period towardsthe end of the dry season. Leaves turn red be-fore shedding. In Liberia and C6te d'Ivoireflowering is irregular but peaks around De-cember and May. During flowering the groundunder the tree is covered with fallen petals andstamens with a penetrant and unpleasantsmell. It has been reported that abundantfruiting occurs twice a year. The fruits are dis-persed by wind. In forests where elephants arepresent, Petersionthus inderocorpt, s trees havestrongly swollen bases as a reaction to regulardebarking. After debarking the bark growsback not only from the edge of the injury butalso from pores in the wood which speeds uprecovery and results in reduced rates of infoc-tion.

Ecology In Liberia PetersionthL, s mererocor-pus ls most common in moist semi-deciduousforest and more uncommon in evergreen forest,but in C6te d'TVoire and Ghana it seems to be

more abundant in evergreen forest and transi-tional zones between and moistevergreen

semi-deciduous forest. It apparently does nottolerate waterlogging for longer periods. InCentral Africa it is reported to be characteristicfor secondary forest. In southern Cameroon itis frequently found in agroforestry plantationsof cocoa. Petersionthus ingcrocorpus prefersregions with an annual rainfall of about 2000min

Propagation and planting In Guinea andC6te d'Ivoire seeds are best collected in Janu-

ary-February(-April) and August. There areabout 4300 seeds per kg. The germination rateis only 15-25%, and many fruits do not developa viable seed or are attacked by insects. Ger-mination starts 3.5-7(-10) weeks after sowing.Seedlings are ready for planting after I yearIn planting tests in Guinea, mortality wasquite high, particularly when seedlings hadbeen planted in fullsun.

Management In forest in southern Came-roon, the average density of Petersionthusingcrocorpus trees with a hole diameter ofmore than 60 cm is 0.3-04 trees per ha, with amean wood volume of 1.5-3 ms per ha. In Ga-bon the mean wood volume has been reportedas 0.2 matha. Trees can be coppiced

Harvesting In Ghana, Central African Re-public and Gabon, the minimum hole diameterallowed for felling is 70 cm. In 2001 it was 50

cm in Cameroon.

Yield A hole felled in south-western DR

Congo branchless for 20.5 in and with a diame-ter of 85 cm yielded 7.7 ing of wood.

Handling after harvest It has been record-ed that freshly felled logs were subject to pin-hole borer attack, but in general logs do notdeteriorate rapidly when left in the forest afterfelling, although deep splits may develop aftera longer time. Freshly felled logs are too heavyto be transported by river

Genetic resources Petersiunthus macro-

corpus is fairly widespread in West and Cen-tral Africa and occurs commonly in secondaryforest. Export volumes are low, certainly sofrom Ghana. It is rarely harvested for the in-ternationaltimber market and not commonlycut for local use because of the unpleasantsmell and the hardness of the wood. Hence, atpresentthe species is notthreatened

Prospects Peterstonthus ingcrocorpus woodhas been suggested as a substitute of othermore durable woods which have been over-

exploited. However, little information is avail-able on growth rates, propagation and SIIvicul-tural aspects, but the data available suggestthat low rates of regeneration may hampersustainable exploitation from natural forest.More research is justified in the light of its fairwood quality, good hole shape and size and itspreference for disturbed forest.

Major references CIRAD Forestry De-partment, 2008; de Koning, 1983; Fouarge &G6rard, 1964; Irvine, 1961; Liben, 1971; Siepel,Poorter & Hawthorne, 2004; Tailfer, 1989;Takahashi, 1978; Voorhoeve, 1965; White &Abernethy, 1997.

Other references Addae-Mensah & Ayar-kwa, 1998; Aubr6ville, 1959c; de in Mensbruge,1966; Fouarge, Quoilin & Roosen, 1970; Haw-thorne, 1995; Hawthorne & Jongkind, 2006;Keay, 1954c; Kouitcheu-Maheku, Kuiate & Es-same, 2011; Mangenot & Mangenot, 1962;Mengome at a1. , 2009; Mengome at a1. , 2010;Nkeoua & Boundzanga, 1999; Normand &Paquis, 1976; 0teng-Amoako (Editor), 2006;Sallenave, 1955; Sallenave, 1964; Sallenave,1971; Senterre & Leioly, 2001; Terashima &Ichikawa, 2003; Wilks & Issemb6, 2000.

Sources of illustration Liben, 1971; Voor-hoeve, 1965; Wilks & Issemb6, 2000

Authors F. W. Owusu

PHYLLARTHRONMADAGASCARIENSE

K. Schum.

Protologue Engl. & Prantl, Nat. Pflanzen-fom. IV, 3b: 250, t. 93H (1895).

Family BignoniaceaeOrigin and geographic distribution Phyl-

107thron inoddgoscoriense is endemic to Mada-gascar, where it is widespread in the easternand central parts of the island. It is commonlyplanted in Madagascar.

Uses The tough wood, known as 'zahana', istraditionally used for the handles of weapons,especially of assegais. It is also in demand forbridges, fence posts, mine props and railwaysleepers because of its durability. It is suitablefor heavy flooring and framework, joinery, in-tenor trim, vehicle bodies, sporting goods, toys,novelties, agricultural Implements and turn-ery

The fruits are edible although not very palata-ble, with a consistency and taste of dried hana-na. The stem bark is used in traditional medi-cine to treat diarrhoea, inflammations, woundsand cough. The leaves are applied externalIy totreat sores, whereas leaf decoctions are admin-istered against blennorrhoea, and as antineu-ralgic and relaxant. Men drink tea made fromthe leaves to treat impotence

Properties The heartwood is yellowishbrown and distinctly demarcated from thesapwood. The grain is straight, sometimesslightly interlocked, texture moderately fineThe wood is very heavy, with a density of1050-1210 kg/ms at 12% moisture content. Thewood air dries very slowly with little degrade,although there is a slight tendency to surfacechecking. The rates of shrinkage from green to

PHYLLARTHRON 529

I.

Phyllorthron ingdogoscoriense - wild

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oven dry are quite high, 5.0-6.9% radial and7.2-13.6% tangential. At 12% moisture con-tent, the modulus of rupture is 214-282N/min2, modulus of elasticity 14,800-23,540N/min2, compression parallel to grain 90-122N/mm2, shear 11 N/mm2, cleavage 23 N/mmand Chalais-Meudon side hardness 9.1-15.3The wood is rather difficult to saw and workThe use of stellite-tipped sawteeth is recoin-mended. It can be finished to a smooth surface

Pre-boring is needed in nailing. The woodglues, stains and polishes satisfactorily. It isvery durable. It is resistant to impregnationwith preservatives.The roots, heartwood and stem bark containlapachol, a known elicitor of contact dermatitisThe roots and heartwood also contain sesamin,which also may cause contact allergy. Severalindoid and phenethyl glycosides were isolatedfrom the leaves, as well as flavonoids.

Botany Evergreen, small to medium-sizedtree up to 20(-40) in tall; hole branchless for upto 15 in, usually straight, sometimes fluted atbase, up to 80 cm in diameter; bark surfacefissured, blackish, inner bark fibrous; twigsglandular hairy. Leaves opposite, occasionallyin whorls of 3, consisting of 2 flattened andwinged articles formed by petiole and rachis,without leaflets; stipules absent; basal articleobovate-oblong, 4.5-15 cm x 1.5-4 cm, terminalarticle elliptical to oblong-lanceolate, 4-19 cmx I-5.5 cm, leathery, glabrous, pinnatelyveined with numerous lateral veins. Inflo-rescence an axillary or terminal cyme or ra-ceme, up to 20-110wered. Flowers bisexual, zy-gomorphic, 5-merous, large; pedice1 0.5-I cmlong; calyx campanulate, 0.5-I cm long, ribbed;corolla funnel-shaped but narrowly cylindricalat base, 3-4(-5) cm long, 2-lipped with 2 upperand 3 lower obtuse lobes, white to pink or pur-PIish red, with yellow markings in throat,hairy; stamens 4 in 2 unequal pairs, insertedon the corolla, included; disk annular; ovarysuperior, c. 3 min long, slightly 4-angled, 2-celled, style slender, 12-15 mm long. Fruit afleshy spindle-shaped berry 6-8 cm x c. 2 cm,smooth, covered with sticky resin, indehiscent,many-seeded. Seeds globose to obovoid, I-1.5cm long.Phyllorthron inodugoscoriense flowers andfruits from November to May. The flowers arepollinated by large insects such as bees and bysunbirds. The fruits are eaten by lemurs, whichprobably disperse the seedsPhy!IQrthron comprises about 15 species and isrestricted to Comoros (I species) and Madagas-

car. It can be easily recognized by its leavesconsisting of flattened articles (phyllodes) andby its large, sticky, indehiscent fruits.Phyllurthron orticulotum (Desf Poir. )K. Schum. is a smallto medium-sized tree up to25 in tallthat is widespread in eastern Mada-gascar. It differs from Phyllorthron madugos-conense in its leaves having 3-5 articles. Itswood is similar, also known as 'zahana', andused for the same purposes. Other Phy!-IQrthron spp. are either too smalltrees or occurtoo local to be of importance for their timberHowever, some of them are used in traditionalmedicine

Ecology Phy!Jarthron inodogoscoriense usu-any occurs in more humid types offorest, oftenin littoral forest on sand, but up to 1600(-2200)in altitude

Genetic resources and breeding Thereare no indications that Phyllorthron ingdogos-curiense or Phyllorthron orticulotum are jin-mediateIy threatened. Both species are coin-paratively widespread and locally common ineastern Madagascar, whereas they are alsolocally planted.

Prospects Intorination on propagation, plant-ing and growth rates ofPhyllorthron mudugos-curiense and other Phyllorthron spp. is neededbefore their prospects as timber plantationtrees can be judged

Major references BOTza & Keating, 1972;Gu6neau, Bedel & Thie1, 1970-1975; Ferner deto Bathie, 1938; Rakotovao at al. , en prepara-tion; Takahashi, 1978.

Other references Bolteau, Bolteau & A1-10rge-Bolteau, 1999; Debray, Jacquemin & Ra-zafindramba0, 1971; Gu6neau, 1971a; Harman-tenaina at a1. , 2001; Hudson, Lee & Rasoa-naiv0, 2000; Parant, Chichignoud & Rakotovao,1985; Sallenave, 1955; Sallenave, 1964; Schatz,2001; Tillequin, Henri & Paris, 1977

Authors R. H. M. J. Lemmens

ex

PHYl, LOCOSMUSAFRICANUS (Hook. f. )}00tzsch

Protologue Abh. K6nigl. Akad. WISS. Berlin1856: 232 (1857)

Family IxonanthaceaeSynonyms Ochthocosmus @1nconus Hook. f

(1848).Origin and geographic distribution Phyl-

IOCosmus o17iconus occurs from Guinea Bissauto the Central African Republic and DR Congo

Uses The wood is used for poles and posts inhut and house building, and for carpentry. It is

.

Phyllocosmus ofriconus - wild

suitable for hydraulic construction work, floor-ing, mine props, ship building and railwaysleepers, but not recommended for joinery,interior trim, furniture, musical instruments,toys, novelties, carvings and turnery. The PIia-ble twigs are used to make traps for game. Thewood is considered a good firewood and is alsopopular for charcoal production, especially indemand by traditional blacksmithsThe bark is used in traditional medicine. Barkdecoctions are taken to treat diarrhoea, dizzi-ness, stiffness and dysmenorrhoea, and as ano-dyne

Production and international trade Thewood of Phyl!ocosmus of neonus is only usedlocally and not traded on the internationaltimber market

Properties The heartwood is brown to red-dish brown and distinctly demarcated from thefairly wide, yellowish brown sapwood. Thegrain is straight or slightly interlocked, texturefine. Quarter-sawn surfaces show darkerstreaks or a ribbon figure. The wood containssomegum

The wood is heavy, with a density of 915-965kg/in3 at 12% moisture content, and hard butrather brittle. It should be air dried with careto avoid serious degrade. The rates of shrink-age are very high, from green to oven dry about9.2% radial and 14.3% tangential. It is recoin-mended to quarter-saw logs before drying.Once dry, the wood is unstable in serviceAt 15% moisture content, the modulus of rup-ture is 180-210 N/min2, compression paralleltograin 63-79 N/min2, shear 4 N/mm2, cleavage22.5 N/min and Chalais-Meudon side hardness6.8. The wood saws slowly and does not work

easily, but it can be planed and finished to asmooth surface. It holds nails well, although ithas a slight tendency to split. It is moderatelydurable, being moderately resistant to fungibut quite resistant to termite and marine borerattacks. The sapwood is susceptible to Lyctus

Botany Evergreen large shrub or small tomedium-sized tree up to 20(-35) in tan; bolebranchless for up to 17 in but usually muchshorter, usually straight and cylindrical, up to80(-100) cm in diameter, often with many ad-ventitious shoots at base; bark surface slightlylongitudinalIy fissured, with numerous smalllenticels, greyish to nearly black, inner barkfibrous, reddish to dark brown; crown large,with slightly sinuous and drooping branches;twigs glabrous. Leaves alternate, simple; stip-ules minute, caducous; petiole up to 0.5 cmlong; blade ovate to elliptical or obovate, 5-18cm x 2-6.5 cm, curieate at base, acuminate atapex, with glandular teeth at margins towardsapex of leaf, papery, glabrous, plrinately veinedwith 5-7 pairs of lateral veins. Inflorescence anaxillary raceme up to 10 cm long, short-hairy.Flowers bisexual, regular, 5-merous; pedicel I-3 mm long; sepals slightly fused at base, ovate-oblong, c. 1.5 mm long; petals free, ovate-oblong, 3-4 mm long, whitish to yellowish;stamens alternating with petals, slightly fusedat base, up to 5 min long; ovary superior, ovoid,c. 0.5 mm long, style 3-4 min long, stigma 5-lobed. Fruit an ovoid to nearly globose capsule4-6 min x 2.5-5 min, dehiscent with 5 valves,I-3-seeded, with persistent sepals and petalsat base. Seeds ellipsoid, c. 3 mm long, enclosedby a reddish anI. Seedling with epigeal germi-nation; hypocoty1 2-4 cm long, epicotyl veryshort; cotyledons leafy, elliptical, 6-8 min long;first leaves alternate

In C6te d'Ivoire trees usually flower in June-August, but they can also be found flowering inDecember-March. The flowers are fragrantand visited by flies, which are probably themain pollinators. Fruits take about 3 monthsto ripen after flowering. The seeds, with theirreddish anIs, are eaten by birds, which serveas seed dispersersPhyllocosmus comprises about 5 species and isrestricted to Africa. It has long been includedin Ochthocosmus, which is now considered tobe restricted to tropical America.Phy1!o008mus Iemoireonus roe Wild. & T. Durand)T. Durand & HDurand (synonym: Ochthocos-mus Iemoireonus De Wild. & T. Durand) is ashrub or smalltree up to 10(-20) in tall, occur-ring in southern DR Congo, Tanzania, Malawi,

L.

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PHYl. LOGOSMUS 531

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532 TIMBERS2

Zambia, Angola and northern Mozambique. Itswood is used for poles in house building and asfirewood. Twigs are used as tooth brush, andthe smoke from the wood is inhaled to treatcolds, cough and sore eyes. Root decoctions aretaken as vermifuge

Ecology Phyllocosmus of neonus occurs invarious lowland forest types up to about 500 inaltitude, from rather dry to moist forest, perl-odically inundated forest and gallery forest,sometimes even in wooded savanna and ontermite mounds in savanna. In the forest, it isoften an understorey tree

Management Fruits are usually pickedfrom the tree before they open, and the smallseeds are collected from the dehisced fruitsafter drying in the shade. The germination rateis 30-40%; seeds start germinating 3-6 weeksafter sowing. Seedlings prefer shade. In timberproduction forest, Phyllocosmus of neonus isoften considered a weedy species, which wasformerly sometimes killed by girdling or poi-soning, e. g. by sodium arsenite. The tree bolesare often small and short, yielding limitedamounts of wood, but a hole of 17 in long with adiameter of 70 cm felled in DR Congo yielded5.5 ing of wood

Genetic resources and breeding Phy!-IOCosmus of neonus is widespread and locallycommon in various forest types and is not ex-PIOited on a large scale. Consequently, it is notliable to genetic erosion.

Prospects The holes are usually too smalltobe of interest for commercial timber exploita-tion. Moreover, the wood is rather difficult todry, saw and work. Phyllocosmus o17iconusand other Phyl!ocosmus species will remain ofsome Interest for local applications, especiallyfor poles in construction.

Major references Badr6, 1972d; Bolza &Keating, 1972; Burki11, 1994; Fouarge, G6rard& Sacr6, 1953; Wilczek & Boutique, 1958.

Other references Aubr6ville, 1959b; Badr6,1973c; Chilufya & Tengnas, 1996; de Koning,1983; Hawthorne, 1995; Hawthorne & Jong-kind, 2006; Neuwinger, 2000; Normand &Paquis, 1976; Savill & Fox, 1967; Tailfer, 1989

Authors R. H. M. J. Lemmens

I.

PIOcodiscuspseudostipuloris - wild

discus pseudost;PMloris is distributed in SierraLeone, Liberia, 06te d'TVoire and Ghana.

Uses In Liberia the wood of PIOcodiscuspseudostjpuloris is used for poles in housebuilding and for tool handles. It has similarcharacteristics concerning quality and aspectas the wood of European walnut (Juglons regtoL. ), and is suitable for similar purposes such asfirst-class cabinet work and veneer. It is alsoused as firewood. The bark, soaked in water, isused as an embrocation to give relief to achingfeet and legs.

Properties The heartwood is pale to darkbrown. The grain is usually straight, texturefine. The wood is fairly lustrous. It is heavywith a density of about 800-920 kg/ing at 12%moisture content, hard and strong. The rates ofshrinkage during drying are medium. Oncedry, the wood is moderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 131-153 N/min2, modulus of elasticity17,540 N/min2, compression parallel to grain49-57 N/mm2, Janka side hardness 17,760 Nand Chalais-Meudon side hardness 5.3-6.1.

The wood is somewhat brittle and moderatelydifficult to saw and work due to its hardness,but can be finished smoothly. It is durable.The leaves contain saponins and tannins.

Description Dioecious smalltree up to 15 intall; hole slender, often crooked or twisted, upto 30 cm in diameter; bark surface finely platedor covered with bumps and warts, greenishblack, inner bark dark reddish brown; twigsglabrous. Leaves alternate, panpinnately coin-pound with 2 pairs of leaflets, sessile; rachis I-9 cm long; petiolules up to 8 min long; leafletsovate to elliptical, obovate or lanceolate, lower

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PLACODISCUS PSEUDOSTIPULARIS Radlk.

Protologue Sitz. -Ber. Bayer. Akad. 20: 242(1890)

Family SapindaceaeOrigin and geographic distribution PIOco-

leaflets 2-6 cm x I-4 cm, usually clasping stemand resembling stipules, upper leaflets 6-24cm x 1.5-8 cm, curieate at base, acuminate atapex, margins thickened, leathery, glabrous,pinnately veined with 7-12 pairs of lateralveins and with Teticulate fine nervation. Inno-

rescence an axillary false raceme up to 35 cmlong. Flowers unisexual, regular, pale green,nearly sessile; calyx with tube c. 3.5 min longand wide, with 5 short lobes, hairy; petals ab-sent; stamens 8, free; disk hairy; ovary superI-or, 3-lobed and 3-celled; male flowers with ru-dimentary ovary, female flowers with reducedstamens. Fruit a I-2-10bed berry up to 3 cmlong, short-hairy when young but becomingglabrous, yellow to orange when ripe, withfleshy whitish pulp, indehiscent, I-2-seededSeeds ellipsoid, lateralIy flattened, pale brown.Seedling with hypogealgermination.

Other botanical information PIOcodiscus

comprises about 15 species and is restricted totropical Africa. Even though most species areonly small understorey trees, the wood is ap-preciated for its strength and resilience. Thismay be the case for at least the following spe-

Iygona1; 25: intervessel pits small(4-7 prn); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; 41: mean tangential diameter ofvessellumina 50-100 prn; 47: 5-20 vessels persquare minimetre; 58: gums and other depositsin heartwood vessels. Tracheids and fibres: 61

fibres with simple to minutely bordered pits;66: non-septate fibres present; 69: fibres thin-to thick-walled; (70: fibres very thick-walled)Axial parenchyma: (78: axial parenchymascanty paratrachea1); 79: axial parenchymavasicentric; (80: axial parenchyma allform);(85: axial parenchyma bands more than threecells wide); 89: axial parenchyma in marginalor in seemingly marginal bands; 91: two cellsper parenchyma strand; 92: four (3-4) cells perparenchyma 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); 1/6: z 12 rays per mm. Mineralinclusions: 136: prismatic crystals present; 138:prismatic crystals in procumbent ray cells; 142prismatic crystals in chambered axial paren-chyma cells.(S. N'Danikou, P. Baas & H. Beeckman)

Growth and development PIOcodiscuspseu-dosttpuloris grows slowly. In C6te d'TVoire itflowers in July and fruits mature in February-March

Ecology PIOcodiscus pseudostipuloris occursin the understorey of evergreen forest

Propagation and planting There areabout 100 seeds per kg. The germination rateis quite high and germination starts 10-30days after sowing.

Management In forest managed for coin-mercial timber production PIOcodiscus pseudo-stipuloris is usually considered and treated asa weed

Genetic resources PIOcodiscus pseudostjpu-Ions is on the IUCN Red List as 'endangered'.The effects of reduction in forest area in West

Africa have been severe for this speciesProspects For commercial timber producers

PIOcodiscus pseudostipuloris and other PIOco-discus spp. are not very interesting because ofthe small size and often poor shape of the holeand their slow growth. They will remain ofsome local importance because of the excellentwood properties for special purposes such ashigh-quality cabinet work, or where durabilityis required for instance for poles in housebuilding

Major references Bouquet & Debray, 1974;

cies.

PIOcodiscus boricoensis Aubr6v. & Fellegr. is asmall tree up to 15 in tall, occurring locally inevergreen forest in C6te d'TVoire and Ghana,where it is considered vulnerable. The bark has

been used to treat asthma.

PIOcodiscus boyo Aubr6v. & Fellegr. , a smalltomedium-sized tree up to 20 in tall, occurs inclosed forest in C6te d'TVoire and Ghana, whereit can be quite common. The limited distribu-tion area and the severe loss of its habitat ren-

der it vulnerable.

PIOcodiscus glandulosus Radlk. is a small for-est tree of Nigeria, Cameroon and Gabon, hav-ing very strong heartwood.Floeodiscus pontculotus Hauman, a small tomedium-sized tree up to 25 in tall and with ahole diameter up to 40 cm, Is endemic to north-eastern DR Congo and considered vulnerable.The wood is reddish, very hard and used tomanufacture arrows. The fruitis eaten

PIOcodiscus riporius Keay is confined to ripari-an, flooded forest of Guinea Bissau, Guinea,Sierra Leone and Liberia, and is a smalltreeup to 10 in tall

Anatomy Wood-anatomical description JAWAhardwood 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-

PmcoDiscus 533

534 TIMBERS2

Chevalier, 1909; Cooper & Record, 1931; Kryn& Fobes, 1959; Sallenave, 1955; Takahashi,1978

Other references Burki11, 2000; de Koning,1983; de in Mensbruge, 1966; Foullloy & Hall6,1973a; Hall, 1980; Hauman, 1960; Hawthorne,1995; Hawthorne, 1998; Hawthorne & Jong-kind, 2006; Hegnauer, 1990; Keay, 1958j;Ndjele, 1998; Normand, 1955

Authors C. H. BOSch

POGAOLEOSAPierre

Protologue Bull. Mens. SOC. Linn. Paris 2:1254 (1896)

Family killsophylleaceaeVernacular names African brazil nut, inoi

nut (En). Erable d'Mrique (Fr).Origin and geographic distribution Pogo

o1eoso occurs in southern Nigeria, Cameroon,Equatorial Guinea and Gabon

Uses The wood, known in Cameroon as 'ovo-ga' and in Gabon as 'afo', is used for joinery,Interior trim, furniture, boxes, crates, veneer,plywood, particle board and fibre board. It istraditionally used for canoes. It is suitable forlight construction, vehicle bodies, musical in-struments, toys, novelties, vats and turnery.The sweet fruit pulp is edible. The seeds can beeaten fresh or after roasting. They are oftenadded to sauces. They resemble the brazil nut(Bertholletio excelso Humb. & Bonpl. ) fromtropical America in flavour and taste. The seedoil has some resemblance to olive oil and

groundnut oil, and is used for cooking. Theresidual mealremaining after expression of theoilis suitable as feed for cattle. Bark decoctions

are taken as emetic, whereas powdered bark isapplied to wounds and skin diseases. The stonewall of the fruit is burnt and pulverized, andthen applied to soothe toothache. Seed o111staken as purgative in the treatment of gonor-rhoea and as a massage oil. The bark, which isrich in tannin, has been used for dyeing clothblackish

Production and international trade The

wood of Pogo o1eoso has been exported in smallamounts from Gamero0n and Gabon. In 1961

Cameroon exported 1300 ina of logs. Gabonexported on average 230 in3 of sawn wood inthe period 1959-1964 and Equatorial Guinea3200 in31year of logs in the period 1959-1968.The export of logs from Gabon was 100 ms in2003, 265 in3 in 2004 and 780 in3 in 2005. InGabon a ban on commercial exploitation hasbeen introduced in 2009 for a period of 25years. Seeds are fairly commonly available onlocal markets in Cameroon and Gabon. Theyare imported in Equatorial Guinea from Came-roon.

Properties The heartwood is pinkish whiteto pinkish brown, becoming greyish upon expo-sure, and distinctly demarcated from the whiteto greyish, 2-5 cm wide sapwood. The grain isstraight to slightly wavy, texture coarse. Quar-ter-sawn surfaces show a silver-grain figure.The wood is lustrous.

The wood is lightweight, with a density of 400-500 kg/ina at 12% moisture content. It air driesand kiln dries fairly rapidly, with some risk ofdistortion, checking and warping in hacksawnboards. However, the drying rates of boards arequite variable. The rates of shrinkage are inod-erate, from green to oven dry 2.5-3.2% radialand 6.9-9.6% tangential. Once dry, the wood ismoderately stable in serviceAt 12% moisture content, the modulus of rup-ture is 78-90 N/mm2, modulus of elasticity6370-7160 N/min2, compression parallel tograin 35-38 N/min2, shear 6-6.5 N/mm2 cleav-age 7.5-13 N/min and Chalais-Meudon sidehardness 0.9-1.8

The wood is easy to saw and work with ordi-nary equipment. Planing is somewhat difficultdue to the coarse texture. The use of a filler is

needed to obtain a good finish and polish, andalso for painting and varnishing. The nailingand screwing characteristics are satisfactory,as well as the gluing properties. The woodpeels and slices well, producing good-qualityveneer. It is moderately durable, being moder-ately resistant to fungi, fairly resistant to dry-wood borers and susceptible to termites. The

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wood Is easy to treat with preservatives.The weight of 100 dry seeds is about 80 g. Theseeds contain per 100 g total mineral matter3.8 g, protein 19 g, fat 69 g, cellulose 0.6 g andother extractive matters 6.5 g. Hexane extrac-tion of the seeds resulted in an oil which is

fluid at ambient temperature and is mainlycomposed of triglycerides. Methanolic fruitextracts showed anti-inflammatory activity,inhibiting carrageenan-induced paw oedema inrats, and antibacterial activity against Stophy-IOCoccus oureus and Bocillus subtilis.

Adulterations and substitutes The fruit of

Pogo o1eoso strongly resembles that of PondoDieoso Pierre, which is used for similar purpos-us but differs in the more smooth stone.

Description Evergreen, medium-sized tolarge tree up to 40(-45) in tall; bole branchlessfor up to 20 in, usually straight and cylindrical,up to 100(-150) cm in diameter, often withbroad and rounded buttresses; bark surfacesmooth to rough or irregularly scaly, grey togreyish brown, inner bark granular, pink withwhitish bands; crown rounded, with large, as-cending branches; twigs glabrous. Leaves al-

ternate, simple; stipules up to 1.5 cm long, ear-Iy caducous; petiole I-2 cm long, stout; bladebroadly elliptical, 7-15 cm x 5-7 cm, roundedat base and apex, margins entire but slightlymrolled, leathery, glabrous, with glandulardots below, glossy, pinnately veined with nu-merous lateral veins. Inflorescence a paniclecomposed of spikes, short-hairy, many-flowered. Flowers usually functionally unisex-ual, regular, 4-merous, small, whitish, sessile;sepals fused at base, triangular, c. 1.5 mmlong; petals free, c. 3 mm long, 5-7-lobed withgland at apex of the slender lobes; stamens 8,free, nearly sessile; ovary Inferior, 4-celled,styles 4, short. Fruit a globose to ellipsoid,fleshy drupe 4-7 cm in diameter, greenish withnumerous brownish lenticels; stone nearly glo-bose, 3.5-5 cm in diameter, distinctly rippled,very hard, 2-4-seeded. Seeds ovoid, 2-2.5 cm xI-1.5 cm, brown, oily.

Other botanical information Pogo coin-prises a slrigle species, and is classified togeth-er with Antsophylleo (pantropical), Combreto-corpus (tropical Asia) and Polygononthus (trop-ical America) in the family Antsophy!legceoe.Formerly these genera have been included inRhizophoroceoe

Anatomy Wood-anatomical description JAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; (23: shape of alternate pits po-Iygona1); 25: intervessel pits small (4-7 pin);26: intervessel pits medium (7-10 F1m); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; 43: mean tangential diameter ofvessellumina a 200 I'm; 46: S 5 vessels persquare minimetre; (58: gums and other depos-its in heartwood vessels). Tracheids and fibres:61: fibres with simple to minutely borderedpits; (62: fibres with distinctly bordered pits);66: non-septate fibres present; (68: fibres verythin-walled); 69: fibres thin- to thick-walledAxial parenchyma: 79: axial parenchyma va-siCentric; 80: axial parenchyma allform; 82:axial parenchyma winged-allform; 83: axialparenchyma confluent; (92: four (3-4) cells perparenchyma strand); 93: eight (5-8) cells perparenchyma strand; (94: over eight cells perparenchyma strand). Rays: 98: larger rayscommonly 4- to 10-seriate; 99: larger rayscommonly > 10-senate; 102: ray height > Imin; 103: rays of two distinct sizes; 106: bodyray cells procumbent with one row of upright

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536 TIMBERS2

and/or square marginal cells; 107: body raycells procumbent with mostly 2-4 rows of up-right and/or square marginal cells; 109: rayswith procumbent, square and upright cellsmixed throughout the ray; (110: sheath cellspresent); 1/4: S 4 rays per mm; 1/5: 4-12 raysper min

(N. P. Monel, P. E. Gasson & E. A. Wheeler)Growth and development In a test in Ga-

bon, 57% of seedlings planted in the open sur-vived after one year and 87% of seedlingsplanted in the undergrowth of opened-up for-est; after 6 years the survival rates were 18%and 76%, and after 11 years 8% and 61%, re-spectively. The young trees in the opened-upforest reached an average height of 22 in and amean hole diameter of 24 cm 11 years afterplanting. Pogo o1eoso is classified as a shadebearer.

In Gamero0n fruits ripen in June-August andJanuary-March. Trees produce fruits aboutonce every two years' The fruit stones are dis-persed by elephants which eat the fruits. How-ever, in a test in Cameroon, stones collectedfrom elephant dung as well as fresh stonesfailed to germinate within the test period ofone year. Gorillas also eat the fruits

Ecology Pogo o1eoso is a canopy tree of low-land humid evergreen forest, where it is oftenfound in valleys, also in swamp forest. It maybe found in secondary forest associated withokoum6 (Aucoumeo hmmeono Pierre), where itoften shows better growth than the latter spe-cies. It is sometimes a relic in farmland

Propagation and planting The germina-tion offruit stones is irregular and may take along period. Wildlings can be used for planting,butthey are often rare in the forest.

Management In general, Pogo o1eoso isquite uncommon in the forest; only locally it ismore common or even abundant. In forests in

south-western Cameroon, the average densityof Pogo o1eoso trees with a hole diameter ofmore than 60 cm is 0.06 per ha, with an aver-age wood volume of 0.62 mama. In Gabon theaverage wood volume has been recorded as0.56 mama. The trees are sometimes left stand-

ing after forest clearance because they are val-ued by the local population for their oil-richseeds.

In Gabon Pogo o1eoso has been planted in re-forestation projects. It was considered promis-ing, but needed to be planted under light tomoderate shade conditions.

Diseases and pests In Gabon longicornbeetles have been recorded as a common pest

in standing treesYield A hole branchless for 18 in and with a

diameter of 90 cm may yield 10 in3 of woodThe mean weight of the fruitis 90 g, that of thestone 40 g, and that of the seed 5 g

Handling after harvest Logs should beremoved rapidly from the logging sites afterharvestlrig because they are moderately sus-ceptible to attacks by fungi and insects.

Genetic resources Pogo o1eoso has a re-stricted area of distribution and is usually notabundant in the forest. It might be easily liableto genetic erosion, although it has been exclud-ed from commercial exploitation in Gabon

Prospects Although Pogo o1eoso is a multi-purpose tree which is not only important for itstimber, but also for its edible, oil-rich seeds andas medicinal plant, little is known about manyaspects, particularly about SIIviculture, man-agement and propagation. At present, Pogoo1eoso does not seem to have good prospects formore intensified timber exploitation. However,tests in Gabon showed rapid growth of seed-lings planted in the undergrowth of opened-upforest, and this may offer possibilities for en-richment planting in natural forest. The maindifficulty is the irregular and slow germinationof seeds

More research is needed on the properties andmarket opportunities of the seed oil. Some ap-PIications in traditional medicine have beenconfirmed by Dharmacologicalstudies, and thismay offer possibilities for drug development.

Major references Bolza & Keating, 1972;Burki11, 1997; CIRAD Forestry Department,2008; CTFT, 1950b; Eyog Matig at al. (Editors),2006; 0gbole at a1. , 2007; Raponda-Walker &Sillans, 1961; Vivien & Faure, 1985; Vivien &Faure, 1996; Wilks & TSSemb6, 2000

Other references Chudnoff, 1980; de Saint-Aubin, 1963; Gassita at al. (Editors), 1982;Keay, 1989; Koumba Zaou at a1. , 1998; Louppeat a1. , 1999; Nchanji & Plumptre, 2003;Neuwinger, 2000; Normand & Paquis, 1976;Painbou Tchivounda at a1. , 1992; Sallenave,1955; Sallenave, 1964; Sunderland & Obama,1999; Tailfer, 1989; Takahashi, 1978; Tobe &Raven, 1988; White & Ahemethy, 1997.

Sources of illustration Raponda-Walker &Sillans, 1961; White & Ahemethy, 1997; Wilks& Issemb6, 2000

Authors AT. Tchinda

POLYSCIASFULVA (Hiern) Harms

Protologue Engl. & Prantl, Nat. Pflanzen-fom. 111, a 45 (1894)

Family AraliaceaeVernacular names Parasoltree (En)Origin and geographic distribution Poly-

setos Iuluo is widespread from Guinea east toEthiopia and Kenya, and south to ingola,Zimbabwe and Mozambique, but it occursmainly in mountain regions

Uses The wood is used for interior joinery,doors, utensils, musical instruments, contain-ers, boxes, crates, beehives, carvings, matches,veneer and plywood. In Cameroon it is valuedfor carving to make handicrafts and masks,and in Uganda for making drums. It is alsoused as firewood, although of low qualityIn traditional medicine in DR Congo, bark in-fusions or decoctions are taken for the treat-

merit of fever and malaria, as an enema totreat colic, and as a purgative. A bark macera-tion is applied as drops to the nostrils to treatmental illness. Pulverized bark is snuffed as

anodyne, and to treat cough, haemoptysis andtuberculosis. In Gamero0n the bark is used in

mixtures with other plants to treat epilepsyLeaf decoctions are taken to treat intestinal

complaints including those caused by para-sites, whereas pounded leaves are applied ex-ternally to treat fractures and internallyagainst peptic ulcers. Leaves make good mulch,and the tree is suitable for intercropping withbanana, coffee and cocoa. It is planted in lifefences. The flowers are a good source of nectarand pollen for honeybees.

Properties The heartwood is whitish to paleyellow-brown, and indistinctly demarcated

from the sapwood. The wood is soft and brittleThe grain is usually straight, texture moder-ately fine to rather coarse. The wood has nodistinct smell. The wood is lightweight, with adensity of 330-450 kg/ms at 12% moisture con-tent. It is fairly easy to air dry, but splittingand ring shaking may occurAt 12% moisture content, the modulus of rup-ture is 61 N/mm2, modulus of elasticity 8625N/min2, compression parallel to grain 29N/min2, cleavage 12.5 N/min and Chalais-Meudon side hardness 1.3. The wood is easy tosaw and work, but often planes to a ratherwoolly surface. It is liable to splitting uponnailing and screwing, but glues well. The woodis not durable; it is susceptible to attacks byfungisuch as blue stain and brown and whiterot, but it is easy to treat with preservativesSome triterpene glycosides have been isolatedfrom the stern bark. One of these, u-hedenn,showed antifungal activity against Candidoo16icons and Cryptococc!, s neoformons.

Botany Deciduous medium-sized tree up to30 in tall; hole branchless for up to 15(-25) in,straight and cylindrical, up to 100 cm in diam-eter; bark surface grey, smooth, with lenticels,inner bark whitish; crown umbrella-shaped,with main branches often whorled, bendingupward; branches with prominent leaf scarsLeaves arranged spiralIy, clustered at ends ofbranches, usually imparipinnately compoundwith (3-)5-12(-15) pairs of leaflets; stipulesabsent; petiole up to 25 cm long, slightlygrooved; petiolules up to 8(-14) min long; leaf-lets ovate to oblong or lanceolate, 6-20 cm x 2-10 cm, rounded to slightly cordate at base,acute to acuminate at apex, margins entire toslightly wavy, papery to leathery, yellowishbrown hairy below, pinnately veined. Inno-rescence an axillary panicle up to 70 cm long,yellowish hairy. Flowers bisexual, regular, 5-merous, creamy to greenish yellow, honey-scented; pedice1 I-3 min long, Jointed; calyxwith very short teeth; petals free, ovate, 1.5-2min long, early caducous; stamens alternatingwith petals; ovary inferior, 2-celled, styles 2,0.5-I mm long, persistent in fruit. Fruit anovoid drupe-like berry up to 6 mm long, flat-tened, ribbed, purplish black, glabrous orslightly hairy, 2-seeded. Seeds ovoid, slightlycompressed, 3-4.5 mm long, ribbed, glabrous.Polyscios fu!Do is fast growing. In plantationsin Garnero0n seedlings reached a height of 2(-3) in after 4 years and 15 in with a bole diame-ter of 20(-40) cm after 20 years. The flowersare pollinated by insects such as bees and flies.

POLYSCIAS 537

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Polyscius Iuluo- wild

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Polyscios comprises more than 100 species andis distributed in tropical Africa, Asia, Australiaand islands in the Pacific Ocean. Mainland

tropical Africa has about 10 species and Mada-gascar about 50.The wood of Polyscios hikuye, 1sts Summerh. , a

to 25 in tall with amedium-sized tree up

straight hole up to 120 cm in diameter andendemic to central Kenya, is similar to that ofPolyscios Iuluo and used for similar purposes,especially for food containers, boxes and ply-wood. In Madagascar the wood of some Poly-setos spp. is used by the local population, andis known together with the wood of severalother genera of Arolioceoe as 'voantsilana'. Oneof these is Polyscios ornifolio (Baker) Harms, asmallto medium-sized tree up to 20 in tall, thewood of which is heavier (density about 630kg/ing at 12% moisture content), stronger andharder than that of Polyscios Iuluo, and locallyused for light construction and flooring, as wellas for matches

Ecology Polyscios Iuluo is found in differenttypes of forest up to 2450(-2750) in altitude,often in secondary forest, also in mountaingrassland and in vegetations dominated bybamboo. In West Africa it seems to be confined

to mountain areas. The mean annual rainfall

in its area of distribution is 1500-2000 mm.

Polyscius Iuluo is sometimes considered a PIO-neer species, but in Cameroon natural regen-eration seems to be rare. It is sensitive to fire,but may recover by producing sprouts andsuckers

Management Seeds and cuttings are usedfor planting. One kg contains about 300,000seeds. The seeds germinate after 5-7 weeks,with a germination rate of up to 75%. Fruitscan best be collected from the trees as soon as

they have become purplish black, but they aresometimes also collected from the ground. Theyshould be dried in the shade for I-2 days. Thenthe fruits should be soaked in water for 4-6

hours and the seeds squeezed out; they float inthe water. After drying the seeds in the shade,they can be stored for up to 2 years, preferablyat 3'C and 7-10% moisture content. Seedlingsare gradually exposed to the sun and can betransplanted from the nursery into the fieldafter 4-6 months. Wildlings are commonly col-Iected for plantingThe logs are susceptible to attack by blue stainfungi directly after felling, and should be treat-ed with anti-stain solution or transported fromthe logging area and processed immediately.

Genetic resources and breeding Poly-

setos Iuluo has a very wide distribution area,but it is uncommon in several regions, e. g. inWest Africa. In Kenya it has been reportedthat Polyscios Iuluo is becoming rare in itsnatural habitat. This is also locally the case inCameroon and Uganda, where the species hasbeen overexploited for the production of handi-crafts. Monitoring of populations in such re-gions where it is under pressure because oftimber harvesting is recommended, but Poly-setos Iuluo is not yet subject to serious geneticerosion, as is the case in Kenya with the en-demic Polyscios Athuyensis that is already in-cluded as vulnerable in the IUCN Red List.

Prospects Although the wood of PolysciosIu!Do is of rather poor quality for building pur-poses, it seems to have good prospects forcommercial production of carvings, veneer andplywood. The fair growth rates and the devel-opment of a long and straight hole are advan-tageous. Polyscios Iuluo has good characteris-tics as a shade tree and auxiliary plant, whichmake it interesting for agroforestry systems. InUganda it has already been recommended tostart on-farm production of Polyscios Iuluo toguarantee future supply of the wood for pur-poses for which it is much in demand, such asdrum production. However, research is stillneeded on propagation techniques and ongrowth and development.

Major references Latham, 2007; Maundu& Tengnas (Editors), 2005; Takahashi, 1978;Tennant, 1968; World Agroforestry Centre,undated

Other references Bamps, 1974; Bedir at al. ,2001; Beentje, 1994; Bekele-Tesemma, 2007;Chikamai at al. , undated; Katende, BITnie &Tengnas, 1995; Neuwinger, 2000; Noumi &Fozi, 2003; Tshibangu at a1. , 2002; Vivien &Faure, 1985

Authors R. H. M. J. Lemmens

POPULUS ILICIFOLIA (Engl. ) Rouleau

Protologue Rhodora 47: 362 (1945).Family SalicaceaeVernacular names Tana River poplar(En).Origin and geographic distribution Popu-

Jus incifolio is endemic to eastern Kenya andnorth-eastern Tanzania

Uses The holes of Populus incifolio trees areused to make dug-out canoes. They are the firstchoice of Pokomo people for canoes, and thesewere often sold down the Tana River to fisher-men on the coast. These canoes were very

.

Populus incifolio - wild

common in the Malindi-Lainu area of Kenyaand were often taken aboard sea-going dhowsfor use as dinghies. The wood is also used forpoles, posts, utensils and beehives. Populusincifolio is suitable for the production of ply-wood. The wood is used as firewood, but is oflow quality for this purpose. The tree is usefulfor the stabilization of river banks and as or-

namental. The foliage is eaten by livestock.Properties The wood is brown, with a

coarse texture. It is lightweight, with a densityof about 500 kg/ms at 12% moisture content,and rather soft. The wood air dries well withsome distortion and end checking. At 12%moisture content, the modulus of rupture isabout 70 N/mm2 and compression parallel tograin 36 N/min2. The wood is easy to saw andwork. The durability of the wood is low. It hasbeen reported that dug-out canoes last for up to2 years

Botany Deciduous, dioecious, medium-sizedtree up to 25(-30) in tall; hole cylindrical; barksurface smooth and whitish to pale brown, butbecoming longitudinalIy fissured and red-brown or grey-brown in older trees; crown coni-cal to rounded, open; twigs longitudinalIy fis-sured, glabrous, reddish brown, with scatteredlenticels. Leaves alternate, simple; stipulesstrap-shaped or obovate, up to 5 mm long, thinand reddish, deciduous; petiole (0.5-)2.5-4.5cm long; blade angular ovate or obovate, onjuvenile growth linear to narrowly elliptical, 4-7.5 cm x (0.5-)4-5 cm, base curieate to round-ed, apex acute to lobed, margins irregularlylobed or coarsely toothed, papery, glabrous orsparsely hairy, pinnately veined with 4-7(-8)pairs of lateral veins. Inflorescence a unisexual

catkin; male inflorescence axillary, 1.5-2 cmlong, 6-9-flowered; female inflorescence termi-nal on short branches, I-4 cm long, 5-15-flowered. Flowers unisexual, in axils of spatu-late bracts 3-6 min long, lobed or toothed atapex, without perianth, disk thin, reddish,deeply fringed; male flowers with pedicel c. Imin long, stamens many, c. 1.5 min long; fe-male flowers with pedice1 3-5(-8) mm long,ovary ovoid, 3.5-5 min long, hairy, style c. Imm long, stigmas 4, 6-7 mm long, 2-3-10bed.Fruit an ovoid capsule 0.5-1.5 cm x 0.5-I cm,furrowed, rough with pale lenticels, dehiscentwith 2-4 valves, many-seeded. Seeds ovoid, I-2min long, with tuft of long white hairsPopulus comprises about 40 species, most ofthem confined to temperate and subtropicalareas of the northern hemisphere. Populusincifolio is the only species of the genus that isa native to tropical Africa. Several Populusspecies and hybrids have been introduced intropical Africa for timber and pulpwood pro-duction, or as ornamental. They are mostlygrown in the highlands. Populus o160 L. andPopulus deltoides Marshall are most widelyplanted in tropical Africa. Both are medium-sized trees up to 30 in tall. The hybrid Populusxeonodensis Moench (Populus deltoides x Po.pulus nigro L. ) has been planted in Mauritiusto serve as a windbreak

Populus incifolio is a shade-intoIerant pioneerspecies. It is short-lived, with a lifespan ofabout 50 years. Leaf fallis usually during therainy season. It flowers irregularly, dependingon moisture levels. Pollination is by wind. Ripefruits are most common in August-December,and the seeds with their long hairs are widelydispersed by wind

Ecology Populus ilicifolio is found in river-me forest and woodland, and on sand banksand mud flats, from sea-level up to 1000 inaltitude. It is most frequently encountered onlow-lying sandy point bars that are frequentlyflooded, but it requires deep soils. The meanannual rainfall within the area of distribution

is 200-800 mm and the mean day tempera-tures vary between 17'C and 35'C

Management Populus theifolio producesnumerous seedlings on alluvial soils, but manyof these are washed away by floods and darn-aged by grazing animals. It can be propagatedby seed, wildlings, layering, and stem and rootcuttings. For propagation by seed, fruits shouldbe collected before they open. After opening ofthe fruits, the seeds are placed in seed traysfilled with fine sand flooded with water. They

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POPULUS 539

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germinate within a few days. They can also bestored for up to one year at O'CStern cuttings of 8-10 cm long and I-2 cm indiameter are recommended; they should bekept moist and at about 25'C, rooting after 3-6weeks. Root suckers can be planted, as is thecase for other Populus spp. Trees coppice quitepoorly. When they are planted as ornamental,it Is recommended to keep some distance frombuildings and pavements because of the largesuperficial roots and because trees are short-living with easily breaking branches. Inter-cropping with crops such as maize, rice, milletand banana is possible as the quite open treecrown gives little shadeSeedlings are commonly attacked by blight andstem-rot caused by FMSorium, caterpillars ofPholontho butterflies and by scale insects. Inlarger-sized holes, heart rot is common.

Genetic resources and breeding Populusthatfolio is classified in the IUCN Red List asvulnerable because of its restricted distribution

area and habitat degradation. However, aslong as suitable habitats along rivers areavailable, It Is not threatened because of itspioneer nature, regenerating easily on alluvialsoils. Some collection of germplasm has beendone in Kenya.

Prospects Populus incifolio is a promisingplantation tree coinparable to other Populusspp. in temperate regions. Its rapid growthenables high production of wood, which is,however, not suitable for purposes where dura-hinty is required. The wood is suitable for SImi-Iar purposes as that of other Populus spp. , es-pecially for boxes, crates, utensils, matches,veneer, plywood and pulp for paper productionThe establishment of experimental plantationsseems worthwhile, but the rather narrow eco-logical niche of Populus inctfolto should betaken into consideration.

Major references Chikamaiet al. , undated;Maundu & Tengnas (Editors), 2005; 0balla,1996; 0balla at a1. , 1996; Wilmot-Dear, 1985.

Other references Beentje, 1994; Dale &Greenway, 1961; Jobansson, 1991; Maingi &Marsh, 2006; Marais, 1985; MeIkle, 1989;Nioroge at a1. , 2010; van Wyk & Gencke, 2000;Weiss, 1973; Wilmot-Dear, 1991a.

Authors C. H. BOSch, L. FA. Oyen & R. H. M. J.Lemmens

POUCHETIAAFRICANAA. Rich. ex DC

Protologue Prodr. 4: 393 (1830)Family RubiaceaeOriginandgeographicdistributionPouche-

It0 @1ncono is distributed from Senegal andGambia to Gabon and DR Congo.

Uses In Sierra Leone the stems are used as

supports for fishing nets. In traditional medi-cine in Senegal a decoction of leafy twigs istaken in draught and used in frictions for thetreatment of kidney pain, stitch, lumbago,stiffness and other pains. A leaf decoction istaken in draught for the treatment of dysen-tery. In Benin leaf decoctions are externalIyapplied on fractures

Botany Evergreen shrub or smalltree up to6 in tall; hole cylindrical, slender; branchesslender, horizontal. Leaves opposite, simpleand entire; stipules triangular, 3-6 min long;petiole 3-8(-10) mm long, grooved; blade ob-long-elliptical, 2-15 cm x I-6.5 cm, base obtuseor curieate, apex broadly acuminate, glabrousor with hairs in the axils of the main lateral

veins, pinnately veined with 4-8 pairs of Iat-eralveins. Inflorescence an axillary or terminalraceme or panicle up to 11 cm long, lax, pendu-IOUs, glabrous; peduncle 3-5 cm long. Flowersbisexual, regular, 4-5-merous, glabrous, fra-grant; calyx campanulate, lobes triangular, c.05 mm long; corolla funnel-shaped, whitish orgreenish, glabrous, tube 3-4 min long, gradual-Iy widening, lobes elliptical or oblong, 1.5-3.5mm long, more or less erect; stamens Insertedon corolla tube, alternating with lobes, 3-4 mmlong, arithers linear, nearly sessile; ovary info-nor, 2-celled, style bifid. Fruit an ellipsoid her-ry I-1.5 cm long, acute, red to blackish purple,

.

Pouchetio of neono - wild

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crowned by persistent calyx, up to 15-seeded.Seeds compressed.Pouchetio of neono is quite variable and 2 van-eties have been distinguished. In BeninPouchetio ofrico, IQ flowers and fruits in Janu-ary-FebruaryPouchetio comprises 4 species, distributed inWest and Central Africa. Pouchetio porui/lordBenth. is a shrub or small tree up to 5 in tall,distributed from Guinea and Sierra Leone toSao Tome at Principe and Bioko (EquatorialGuinea). The wood is used for rafters in hutconstruction.

Ecology Poucheti0 o17icono occurs in galleryforest in savanna regions and in coastal vege-tation, but also in swamp forest and humidforest

Genetic resources and breeding In viewof its wide distribution, Poucheti0 @1nconoseems notthreatened with genetic erosion

Prospects Poucheti0 o17icono is only locallyused as a source of stakes and traditional med-ICines. In view of its small size its importanceas a source of wood is unlikely to increase, butresearch on pharmacological activity is war-ranted because the leaves are fairly widelyused for medicinal purposes.

Major references Ako6gninou, van der Burg& van der Maesen (Editors), 2006; Burkill,1997; Han6, 1970; Hawthorne & Jongkind,2006; Hepper & Keay, 1963

Other references Aubr6ville, 1950; Figueire-do, 2005; Irvine, 1961; Keyharo & Adam, 1974.

Authors M. Brink

.

POUPARTiA 541

POUPARTiAsiLVATicAH. Ferner

PDUportio silootico- wild

Protologue Mein. Mus. natl. Hist. nat. , Parisn. s. 18: 247 (1944)

Family AnacardiaceaeOrigin and geographic distribution Pou-

portiu snugtico is endemic to western Mada-gascar, where it is widespread from the ex-treme north to the south.

Uses The wood is sometimes used for con-struction, although it is not durable, and forknife handles. It is suitable for the productionof veneer, plywood and particle board, and forpanelling, light boxes and crates. The fruitpulp Is occasionally eaten.

Properties The wood is white to pale greyThe grain is straight, texture medium. Thewood is lightweight, with a density of 320-500kg/ina at 12% moisture content, and soft. It airdries rapidly and the rates of shrinkage are

moderate, from green to oven dry 2.4-3.8%radial and 4.5-7.2% tangential. Once dry, thewood is stable in service. At 12% moisture con-

tent, the modulus of rupture is 49-110 N/min2,modulus of elasticity 5000-14,000 N/min2,compression parallel to grain 20-42 N/mm2,cleavage 18-24 N/min and Chalais-Meudonside hardness 05-1.7. The wood works easilyand finishes well. It is not durable, although intests it showed fair resistance to fungal at-tacks. It is quite easy to impregnate with pre-servatlves

Botany Deciduous, dioecious, small to medi-urn-sized tree up to 20(-30) in tall; hole usuallystraight, up to 60(-100) cm in diameter; barksurface greyish, cracking and scaly, inner barkreddish brown, with reddish exudate; youngtwigs short-hairy. Leaves arranged spiralIy,clustered near apex of twigs, imparipinnatelycompound with 5-11 leaflets; stipules absent;petiole and rachis flattened above; petiolules5-12 min long, but 2-4 cm long in terminalleaflet; leaflets opposite, ovate to lanceolate, 5-9.5 cm x 2.5-5 cm, asymmetrical at base, acuteto acuminate at apex, glabrous, pinnatelyveined with 8-9 pairs of lateral veins. Inno-rescence an axillary panicle or raceme up to 15cm long, short-hairy. Flowers unisexual, regu-Iar, 5-merous, whitish; pedicel up to 4 minlong; sepals free, up to I min long, rounded;petals free, ovate, 2-3 mm long; stamens 10,free; disk thick, annular, slightly 10-lobed;ovary superior, spherical, 5-celled, with 5 ses-sile stigmas; male flowers with strongly rudi-meritary ovary, female flowers with rudimen-tary stamens. Fruit an ovoid drupe c. 2.5 cm x2 cm, yellowish to orange when ripe, with

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fleshy pulp; stone hard and bony, with 3 0per-CUIes below the apex and with 5 irregular cavi-ties, I-seeded. Seed kidney-shaped, c. I cm indiameter. Seedling with epigealgermination.Growth is rather slow. in annual growth rateof seedlings of 25 cm has been recorded. Pou-portio siluottco usually flowers in September-December, but mainly at the beginning of therainy season, and fruits mature a few monthslater, stillin the rainy season.Pouportio comprises about 7 species and isrestricted to Madagascar and the MascareneIslands. It is closely related to Seierocoryo fromcontinental Africa, which is sometimes evenincluded in Pouportio.Pouportio chopelieri(Guillaumin) H. Perrier isendemic to eastern Madagascar, where it iswidespread from Antsiranana in the north toTaolaiiaro in the south, in humid, evergreenforest, from sea-level up to 1500(-1700) in alti-tude. It is an evergreen smallto medium-sizedtree up to 20(-30) in tall, with usually straighthole, up to 100 cm in diameter. The wood isused for the same purposes as that of Poupor-tio snugtico. The fruit pulp is sometimes eaten.Protium ingdogoscoriense Engl. (Burseroceoe)closely resembles Pouportio chopeliert and thetwo species are often confused. Slashed bark ofProtium madogoscoriense has a strong turpen-tine smell, which lacks in Pouportio chopelieri.POMPortio orientolis Capuron ex A. Randriana-solo & J. S. Miller is an evergreen, small to me-dium-sized tree up to 20 in tall from easternMadagascar south to Toamasina. It differsfrom POMPurtio chopeliert in its flowers withlonger pedicels and its fruit stone with 5 0per-CUIes (I-2 in POMPortio chopelieri). Its wood isprobably similar and can be used for the samepurposes.

Ecology POMPortio siluotico occurs in sea-sonally dry forest, from sea-level up to 800(-1200) in altitude. It is often found on calcare-ous soils

Management Seeds can be collected aroundFebruary below the trees. The germinationrate is about 50%. The seed still showed fair

germination rates after 2 years of storage.Fruits should be dried for 4-5 days before stor-age. Seedlings are ready for planting into thefield after 12 months, but the survival rate isoften low. They should be protected from wildpigs and be keptfree from weeds.

Genetic resources and breeding Poupor-tio si!uotico is widespread in eastern Madagas-car and locally common. It is not exploited se-Iectively. Therefore, it does not seem to be un-

der threat of genetic erosion at present.Prospects Pouportio siloottco and other

Pouportio spp. of Madagascar may have goodprospects for veneer and plywood production,but although this has been suggested alreadydecades ago, little research has been done untilpresent. Tests on growth performance andpropagation methods are warranted

Major references Blaser at a1. , 1993; CFPF,2008; Gu6neau, Bedel & Thie1, 1970-1975; Pa-rant, Chichignoud & Rakotova0, 1985; Perrierde in Bathie, 1946a.

Other references Bolteau, Boiteau & A1-10rge-Bolteau, 1999; Rakotovao at al. , en pre-paration; Randrianas010, 1992b; Randrianasolo& Miller, 1999; Schatz, 2001

Authors R. H. M. J. Lemmens

PRIORIABALSAMIFERA ryermoesen) Breteler

Protologue Wageningen Agric. Univ. Pap.99-3: 21 (1999).

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Synonyms Gosswei!erodeizdron 60180miferumryermoesen) Harms (1925).

Vernacular names Agba (En). ToIa (Fr).ToIa branca (Po).

Origin and geographic distribution Priorio60180mifero is distributed from southern Nige-ria eastward to DR Congo and south to Cabin-da (Angola)

Uses The wood, known in trade as 'toIa' or'agba', is used for construction, flooring, join-ery, interior trim, ship building, furniture, box-es, crates; shingles, veneer and plywood. It issuitable for vehicle bodies, toys, novelties, vats,

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Priorio balsamifero- wild

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carvings, turnery, hardboard, particle boardand pulpwood. The holes are traditionally usedto make canoes. The resin exuding from thesapwood has been used as an illuminant, alt-hough it reportedIy causes a smoky flame, andfor caulking canoes.

Production and international trade In

1961-1970 Nigeria exported on average 22,000in3 of logs and 11,000 ing of sawn wood ofPriori0 60180mifero per year. The export of logsfrom Gabon increased from 20,500 ms in 1991to 56,000 ina in 2001. In 1999 Priori0 601somi-Iero ranked 7th on the list of most importantGabonese export timbers. Exports decreased to13,800 in3 in 2005. In 2004 Congo exported13,000 ms of logs at an average price of Us$14/1m3, and 1000 ms of veneer at an averageprice of Us$ 3091m3. Cameroon exported 5600ina of logs in 1999, 12,300 in' in 2000, 3000 msin 2001 and only 850 in3 in 2003. In 2006 smallamounts of plywood were exported from Came-Toon at an average price of Us$ 8671m3

Properties The heartwood is pale yellowishbrown, darkening to pale reddish brown uponexposure, and not distinctly demarcated fromthe up to 10 cm wlde, slightly paler sapwoodThere is an intermediate zone of 2-3 cm wide

between sapwood and heartwood. The grain isstraight or slightly interlocked, texture moder-ately fine and even. Wood surfaces are Ius-trous. Freshly cut wood has a resinous orslightly peppery odour. The intermediate zonebetween sapwood and heartwood exudes agreenish resin in freshly cut logs, and resinpockets may also be present near the heart ofthe hole

The wood is lightweight to medlurn-weight,with a density of 445-580 kg/in' at 12% ino1s-ture content, and moderately hard. It air driesrapidly, with only slight tendency to distortionand checking. Kiln drying should be done withcare to avoid collapse;it is recommended to usemoderate temperatures to avoid excessive resinexudation. The rates of shrinkage are moder-ate, from green to oven dry 2.4-3.0% radial and5.5-6.2% tangential. Once dry, the wood isstable in service. At 12% moisture content, themodulus of rupture is 70-94 N/min2, modulusof elasticity 6075-10,900 N/min2, compressionparallelto grain 33-43 N/mm2, shear 6.5-11.5N/min2 cleavage 9-14 N/min, Janka sidehardness 2440-3300 N, Janka end hardness3510-4400 N and Chalais-Meudon side hard-ness 1.2-2.3.

The wood is easy to saw and work with bothhand and machine tools, with little blunting

effect on saw teeth and cutting edges. However,tools may be clogged by resin and boards maystick together. After drying of the wood, thedifficulties due to the presence of resin havedisappeared. The nailing and screwing proper-ties are satisfactory, and the wood glues, paintsand varnishes well although the presence ofresin may cause difficulties and some stainingmay occur. It polishes to an excellent finish.The steam bending properties are moderatelygood. The logs are suitable for rotary peeling,giving a good-quality veneer that can be gluedsatisfactorily to produce plywood. The wood ismoderately durable, being moderately resistantto fungi, but rather susceptible to termite, dry-wood borer and marine borer attacks. Theheartwood is moderately resistant to Impreg-nation with preservatives, the sapwood is per-meable. Saw dust can cause irritation of eyesand mucous membranes in wood workers.

Tests on the pulping properties for paper pro-duction showed moderate results; the paperwas of medium-quality. The wood contains 39-40% ceUulose, 27-30% jignin, 16.5-19.5% pen-tosan, 0.3-0.4% ash and little silica. The solu-binty is 10.1-11.3% in alcohol-benzene, 1.0-1.7% in hot water, and 17.9-20.2% in a I%NaOH solution.

Adulterations and substitutes The woodof Priorio balsamifero resembles that of Mri-can mahogany (Errtondrophrogmo and inlayospp. ) and has been traded as a substitute

Description Evergreen, large to very largetree up to 55(-60) in tall; hole branchless for upto 25(-30) in, straight and cylindrical, up to150(-300) cm in diameter, without buttresses;bark surface scaly with elongate scales, some-times slightly fissured, greyish brown to red-dish brown, inner bark fibrous, reddish; crownusually comparatively small, wlth sinuousbranches; twigs glabrous. Leaves alternate,imparipinnately or panpinnately compoundwith (4-)6-8(-9) leaflets; stipules small, veryearly caducous; petiole I-2(-3) cm long, rachis4.5-7(-8) cm long, grooved; petiolules twisted,I-3(-4) mm long; leaflets alternate, obliquelyovate-elliptical to obovate-elliptical, (2.5-)5-7(-9) cm x (1.5-)2-3(-4) cm, rounded to obtuse atbase, obtuse to rounded, sometimes slightlyacuminate at apex, thin-leathery, glabrous,pinnately veined with 9-13 pairs of lateralveins. Inflorescence an axillary compound ra-ceme up to 12 cm long, with branches up to 7cm long, short-hairy. Flowers bisexual, regular,whitish; pedicel up to 0.5 min long; sepals 4(-5), elliptical, I-2 mm long, spreading, hairy at

PRIORIA 543

544 TIMBERS2

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Priorio joueri(Normand ex Aubr6v. ) Breteler(synonym: Gossweilerodendronjoueri Normandex Aubr6v. ) is a large tree up to 40 in tall, withhole up to 150 cm in diameter. It is found inrainforest in Cameroon, Equatorial Guinea andnorthern Gabon, and differs from Priori0 601-sumifero in more numerous lateral veins in itsleaflets and in its fruit tapering to a narrow,curved top. In trade, logs of this species areundoubtedly mixed with those of Priori0 601-sqmtfero and its wood can probably be used forsimilar purposes. Priorio joueriis classified asvulnerable in the IUCN Red List.

Anatomy Wood-anatoimcal description (IAWAhardwood codes)Growth rings: L growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; 13: sim-PIe perforation plates; 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;24: intervessel pits minute (s 4 pin); 25: inter-vessel pits small (4-7 prn); 29: vestured pits;30: vessel-ray pits with distinct borders; SImi-Iar to intervessel pits in size and shapethroughout the ray cell; 42: mean tangentialdiameter of vessellumina 100-200 pin; 47: 5-20 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;69: fibres thin- to thick-walled. Axial paren-chyma: (76: axial parenchyma diffuse); 79: axi-al parenchyma vasicentric; (80: axial paren-chyma anform); 83: axial parenchyma conflu-ent; 85: axial parenchyma bands more thanthree cells wide; 89: axial parenchyma in mar-ginal or in seemingly marginal bands; 90: fusi-form parenchyma cells; 91: two cells per paren-chyma strand; (92: four (3-4) cells per paren-chyma strand). Rays: 97: ray width I-3 cells;98: larger rays commonly 4- to 10-senate; 104:all ray cells procumbent; 106: body ray cellsprocumbent with one row of upright and/orsquare marginal cells; 1/5: 4/2 rays per min.Secretory elements and cainbial variants: 129:axial canals diffuse. Mineral inclusions: (136prismatic crystals present); (142: prismaticcrystals in chambered axial parenchyma cells).(E, Uetimane, P. E. Gasson & E. A. Wheeler)

Growth and development Priori0 60180-mileru is shade tolerant. Natural regenerationcan be abundant. In semi-deciduous forest in

south-western DR Congo, up to 7800 seedlingsand saplings have been counted per ha. Seedgermination can be gregarious under a closedcanopy, but it seems that for good growth ofseedlings openings in the canopy are needed.

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margins; petals absent; stamens usually 10,free, 3-4 min long, filaments hairy; ovary supe-rior, with a c. I min long stipe, hairy, I-celled,style slender, curved. Fruit an ovoid, flattened,indehiscent, I-seeded pod with large wing atbase, (7-)12-15 cm x (2-)3-4.5 cm, with nu-merous resin pockets, wing reticulately veinedSeed rounded to elhpsoid, 2-3 cm long, shal-lowly grooved. Seedling with hypogeal germi-nation; eplcoty1 8-22 cm long, with some smallscales; flrst leaves alternate, with 4-6 leaflets

Other botanical information Priorio coin-prises 14 species, 7 in Africa (most of them inCentral Africa), 6 in tropical Asia and on is-lands of the Pacific Ocean, and I in CentralAmerica. Until 1999 the name Priorio was re-served for the American species, but then thegenera Kingiodendron from Asia and the Pacif-IC region, and Gossweilerodendron and Orystig-ing from tropical Africa have been added basedon inorphological evidence. Wood-anatomical,pollen and molecular studies support the closerelationship, but more studies are needed toassess generic delimitation.

In Nigeria and GarnerDon, flowering trees aremainly found from January to March andfruits ripen a few months later. In DR Congoflowering and fruiting trees have been recordedthroughout the year. Fruits are dispersed bywind, sometimes quite far from the parenttree.

Ecology Priori0 60180mifero is most coin-mon in lowland semi-deciduous forest up to 600in altitude, but can also be found in evergreenforest. It prefers deep ferruginous soils and canalso be found on sandy soils.

Management Priori0 601somifero usuallyoccurs quite scattered in the forest, althoughoften in small groups of a few trees. In Came-roon the average number of trees with a holediameter of more than 60 cm is 0.05-0.1 perha, with a mean wood volume of 0.3-1.25inalba, but locally up to 6 in3ftia. Locally ineastern Gabon and Congo, 4-6 exploitabletrees per ha have been recorded.

Harvesting Some caution is needed duringharvesting operations because holes may havebrittle heart and ring or wind shakes. The min-jinum hole diameter allowed for harvesting is80 cm in Gabon and DR Congo, and 100 cm inGamero0n

Yield A hole of 13 in long and 120 cm in di-ameter yielded 8.7 ms of wood

Handling after harvest Freshly harvestedlogs are sometimes attacked by pinhole borers,but chemical protection of logs is usuaUy notneeded when the logs do not stay in the forestfor longer periods. They float in water and thuscan be transported by river

Priorio balsamiferoGenetic resources

suffered from habitat loss and heavy exploita-tion and has therefore been included as endan-

gered in the TUGN Red List of threatened spe-cies. FAO recommends that the genetic mate-rial of this species should be protected for afuture planting programme

Prospects Priori0 60180mtfero produces animportant commercial timber for the Interna-tional market, but unfortunately there hasbeen no attention for proper management prac-tices to ensure sustainable production. It de-serves research attention on genetics and SIIvi-culture for future planting programmes.

Major references ATIBT, 1986; Aubr6ville,1970; BOLza & Keating, 1972; Breteler, 1999;Burki11, 1995; CTFT, 1979; de Saint-Aubin,1963; Fouarge & G6rard, 1964; Tailfer, 1989;Wilczek at a1. , 1952

Other references African Regional Work-shop, 1998; hatoine, Berben & Sauvage, 1959;Christy at a1. , 2003; Chudnoff, 1980; CTRAD

Forestry Department, 2008; Cordiez, 2000;Fouarge, Quoilin & Roosen, 1970; Fouarge,Sacr6 & Mottet, 1950; G6rard at a1. , 1998;Keay, 1989; Lewis at a1. , 2005; Normand &Paquis, 1976; Sallenave, 1964; Takahashi,1978; UNEP-WCMC, 2006; Vivien & Faure,1985.

Sources of illustration Breteler, 1999;Wilczek at a1. , 1952

Authors JR. Cobbinah & E. A. Obeng

PRIORIA 545

PRIORIAOXYPHYLLA (Harms) Breteler

Protologue Wageningen Agric. Univ. Pap99-3: 37 (1999)

Family Caesalpiniaceae (Leguminosae - Cae-salpinioideae)

Synonyms PterygopodLum oryphyllum Harms(1913), Orystigmo aryphyllttm (Harms) J. Leo-nard (1950).

Vernacular names Tchitola (Fr). ToIa chin-futa, toIa in afuta (Po).

Origin and geographic distribution Prioriooxyphyllo is distributed from southern Nigeriaeastward to the Central African Republic, andsouth to DR Congo and Cabinda (hagola)

Uses The wood, known in trade as 'tchitola',is used for light construction, joinery, furni-ture, boxes, crates, shingles, veneer and ply-wood. It is suitable for light flooring, interiortrim, vehicle bodies, railway sleepers, handles,ladders, toys, novelties, agricultural imple-merits, turnery and pulpwood. The holes aretraditionally used to make canoes. The thickand tough bark is used to make containers. Theresin from the bark is heated to combat para-sites such as lice and Jiggers.

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Production and international trade The

timber of Priorio oryphyllo seems to be tradedon the international market in small amounts

only, although it may be traded in mixed con-signments, e. g. together with toIa (Priori0 601-sqmifero ryermoesen) Breteler). Gabon exportssmall amounts of Priorio oryphyllo wood,mainly as veneer. In 2006 small amounts ofveneer were exported from Congo at an aver-age price of Us$ 3591m3.

Properties The heartwood is pinkish brownto coppery brown, darkening to reddish brownupon exposure, with blackish streaks, and dis-tinctly demarcated from the up to 10 cm wide,pale yellow-pink sapwood. All intermediatezone up to 12 cm wide is often present betweensapwood and heartwood. The grain is straightor slightly interlocked, texture moderatelycoarse. The wood contains a greenish resin,becoming brownish to blackish upon exposureThe wood is medium-weight, with a density of580-670 kg/in" at 12% moisture content, andmoderately hard. It air dries fairly rapidly,with only slight tendency to distortion andchecking. It should not be dried too rapidlybecause spots caused by resin may developthen. The rates of shrinkage are medium tohigh, from green to oven dry 3.7-4.6% radialand 8.0-13.1% tangential. Once dry, the woodis moderately stable to stable in service. At12% moisture content, the modulus of ruptureis 85-137 N/mm2, modulus of elasticity 9310-14,960 N/mm2, compression parallel to grain37-72 N/mm2, shear 14 N/mm2, cleavage 14-17N/mm, Janka side hardness 5560 N andChalais-Meudon side hardness 3.2

The wood is easy to saw and work with bothhand and machine tools, with little bluntingeffect on saw teeth and cutting edges. However,tools may be clogged by resin. The resin con-tent can be high and steam and hot watertreatments are sometimes needed to reduce the

amounts to manageable levels. The nailing andscrewing properties are satisfactory, and thewood glues, paints and varnishes well whenthe surfaces are free of resin. The logs are suit-able for rotary peeling, giving a good-qualityveneer that can be glued satisfactorily to pro-duce plywood. The wood is moderately durable,being fairly resistant to fungi, termites anddry-wood borers, but susceptible to marineborer attacks. The heartwood is moderatelyresistant to impregnation with preservatives,the sapwood is permeable. Saw dust can causeirritation of eyes and mucous membranes inwood workers.

Adulterations and substitutes In Europeand the United States the veneer of Priorio

oryphyllo is considered a substitute for that ofwalnut (Juglons regio L. ) and jatoba or guapi-nol (Hymenoeo COMrboril L. ), mainly used forfurniture and decorative purposes

Description Evergreen, large tree up to 50in tall; hole branchless for up to 20(-40) in,straight and cylindrical, up to 130(-300) cm indiameter, without buttresses; bark surfacescaly with elongate scales, sometimes slightlyfissured, grey to greenish grey, Inner bark fi-brous, pinkish to reddish; crown usually coin-paratively small, with sinuous branches; twigsglabrous. Leaves alternate, imparipinnately orpanpinnately compound with (4-)6-8(-100leaflets; stipules small, c. 3 min long, very earlycaducous; petiole (0.5-)I-2(-2.5) cm long, ra-chis (4-)6-11 cm long; petiolules 3-6(-7) mmlong; leaflets alternate, obliquely elliptical toovate-elliptical, (3-)6-11 cm x (1)2-4.5 cm,curieate to rounded at base, short-acuminate atapex, leathery, glabrous, pinnately veined with6-11 pairs of lateral veins. Inflorescence anaxillary compound raceme up to 20 cm long,

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with branches up to 15 cm long, short-hairy.Flowers bisexual, regular, yellowish white;pedice1 I-1.5 min long; sepals 5, 0bovate-elliptical, 1.5-2 mm long, spreading or erect,usually hairy at margins only; petals absent;stamens 10, free, 2-5 mm long, filamentshairy; ovary superior, c. I min long, denselyhairy, I-celled, style slender, slightly curved.Fruit an ovoid, flattened, indehiscent, I-seededpod with large wing at base, 6-11.5 cm x 2-4cm, sometimes with numerous resin pockets,wing longitudinalIy and reticulately veinedSeed ellipsoid, flattened, up to 2.5 cm longSeedling with hypogeal germination; epicotyl9-17 cm long, with some small scales; firstleaves alternate, with 2 leaflets

Other botanical information Priorio coin-

prises 14 species, 7 in Africa (most of them inCentral Africa), 6 in tropical Asia and on is-lands of the Pacific Ocean, and I in CentralAmerica. Until 1999 the name Priorio was re-

served for the American species, but then thegenera Kingiodendron from Asia and the Pacif-it region, and Gosswetlerodendron and Orystig-ing from tropical Africa have been added basedon inorphological evidence. Wood-anatomical,pollen and molecular studies support the closerelationship, but more studies are needed toassess generic delimitationPriorio buchh012ii(Harms) Breteler (synonym:Oxystigmo buchholzii Harms, Oxystigmo dewe-orei De Wild. ) is a medium-sized tree up to 30in tall, with hole up to 75(-180) cm in diameterIt is found in periodically inundated forestalong rivers from Cameroon east to DR Congoand south to northern Angola. It is character-ized by glabrous inflorescences and unwingedfruits circular to obcordate-obovate in outline.Its wood is used for similar purposes as that ofPriorio oryphyllo, e. g for joinery and tradition-ally for canoesPriorio monnii (Baill. ) Breteler (synonym:Oxystigmo monrrit (Baill. ) Harms) is a medi-urn-sized tree up to 30 in tall, with hole up to70 cm in diameter. It occurs in swamp forestalong the coast from eastern Nigeria to north-ern Gabon. It resembles Priorio buchholzii, butdiffers in short-hairy inflorescences and morerough and dullfruits. Its wood is used for con-struction and utensils, and traditionally for

It has similar properties as that ofPriorio oryphyllo, but the density is slightlylower, about 510 kg/in3 at 12% moisture con-tent, and it is slightly less hard and strong, andnot durable

Prioriu msoo (Harms) Breteler (synonym: Ory-

stigmo msoo Harms) is a large tree up to 40(-50) in tall, with bole up to 150(-300) cm in di-ameter. It is restricted to evergreen forest inKenya and Tanzania. It differs from Prioriomonnii in its sepals which are more hairy in-side, and in its shortly winged fruits with 10n-gitudinalveins. Its wood is used in house build-ing and for canoes, utensils, tool handles, carv-ings and plywood, and as firewood. It has SImi-Iar properties as that of Priorio monnii, with adensity of about 510 kg/ing at 12% moisturecontent, and it is rather soft and not durableThe tree serves as forage for honey bees.Priorio msoo has been classified as vulnerable

in the IUCN Red List

Anatomy Wood-anatonitcal 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);(26: intervessel pits medium (7-10 prn)); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;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; 66: non-septate fibres present;69: fibres thin- to thick-walled. Axial paren-chyma: 79: axial parenchyma vasicentric; 80:axial parenchyma allform; 81: axial parenchy-ina lozenge-allform; 82: axial parenchymawinged-allform; 83: axial parenchyma conHu-ent; 86: axial parenchyma in narrow bands orlines up to three cells wide; 89: axial paren-chyma in marginal or in seemingly marginalbands; 92: four (3-4) cells per parenchymastrand; (93: eight (5-8) cells per parenchymastrand). Rays: 97: ray width I-3 cells; 104: allray cells procumbent; 106: body ray cells pro-cumbent with one row of upright and/or squaremarginal cells; 1/5: 4-12 rays per min; 1/6: Z12 rays per min. Secretory elements and cam-bial variants: 129: axial canals diffuse. Mineralinclusions: 136: prismatic crystals present; 138:prismatic crystals in procumbent ray cells; 142prismatic crystals in chambered axial paren-chyma cells.(S. N'Danikou, P. E. Gasson & E. A. Wheeler)

Growth and development In DR Congoflowering and fruiting trees have been recordedthroughoutthe year

PRIORIA 547

canoes

548 TIMBERS2

Ecology Priorio oryphyllo occurs in lowlandrainforest on well-drained soils

Management Priorio oryphyllo occurs scat-tered in the forest. In Cameroon and Gabon, itIs apparently comparatively rare, but it Is morecommonin CongoandDR Congo

Harvesting The minimum hole diameterallowed for harvesting is 60 cm in Cameroon,70 cm in Gabon and 80 cm in the Central Mri-can Republic and DR Congo.

Yield In DR Congo a hole of 39 in long and110 cm in diameter yielded 19.8 ina of wood

Handling after harvest Freshly harvestedlogs are sometimes attacked by pinhole borers,but Immediate chemical protection of logs isusually not needed. They usually float in waterand thus can be transported by river, but somelogs sink because the density of green wood isclose to 1000 kg/ina

Genetic resources Although Priorio ory-phyllo has a fairly large area of distribution, itoccurs scattered and is in many regions withinits area of distribution uncommon. Like Priorio60180mtfero, it suffered from habitat loss andheavy exploitation, but it is not classified onthe TUGN Red List of endangered species likePriorio balsamijero. Monitoring of the popula-tions of Priorio oryphy!!o is recommended be-cause it may become easily liable to geneticerosion.

Prospects Priorio oryphyl!o produces tim-her that is of great interest for the internation-al market, but the volume of timber availableseems to be limited. Information on growthrates and natural regeneration is needed todraw up directives for proper managementpractices to ensure sustainable production

Major references ATIBT, 1986; Aubr6ville,1970; Bolza & Keating, 1972; Breteler, 1999;Burki11, 1995; CTFT, 1952b; de Saint-Aubin,1963; Fouarge & G6rard, 1964; Tailfer, 1989;Wilczek at a1. , 1952.

Other references Aubr6viUe, 1968; Beentje,1994; Brenan, 1967; Christy at a1. , 2003;Chudnoff, 1980; CIRAD Forestry Department,2008; Fouarge, G6rard & Sacr6, 1953; G6rardat a1. , 1998; Keay, 1989; Lewis at a1. , 2005;Lovett & Clarke, 1998; Lovett at a1. , 2007;Normand & Paquis, 1976; Raponda-Walker &Sillans, 1961; Sallenave, 1955; Takahashi,1978; Vivien & Faure, 1985; Wilks & Issemb6,2000

Sources of illustration Aubr6ville, 1970;Breteler, 1999; Wilks & Issemb6, 2000

Authors R. H. M. J. Lemmens

PROTOMEGABARIASTAPFLANA(Benle) Hutch

Protologue Pram, Icon. PI. 30: t. 2929 (1913).Family Euphorbiaceae(APG: PhyUanthaceae)Chromosome number2n=26

Origin and geographic distribution Proto-megobang stopfiono is distributed from SierraLeoneto Gabon

Uses The wood of Protomegoborio stopfionois widely used for planks, and is also used forconstruction and furniture. It is considered avaluable firewood

Properties The heartwood is yellowish topinkish brown with dark streaks, and notclearly demarcated from the pale brown sap-wood. The grain is mostly straight, texturemedium. The wood is fairly lustrous. It is me-dium-weight, with a density of about 700 kg/msat 12% moisture content. At the same moisturecontent, compression parallelto grain is about53 N/min2, Janka side hardness 6750 N andJanka end hardness 7560 N

The wood is not difficult to work and it finisheswell to a smooth surface. It splits easily and isnot resistant to decay.

Botany Evergreen, dioecious, small to medi-urn-sized tree up to 30 in tall; bole short, low-branching, up to 25 cm in diameter, buttresseswell developed; bark surface brownish withgreen patches; twigs soft-hairy, soon becomingglabrous. Leaves alternate, simple and entire;stipules triangular, 2-4 min long, caducous;petiole 3-11 cm long; blade narrowly oblong-elliptical, 15-37 cm x 7-15 cm, base curieate torounded, apex rounded to short-acuminate,sparingIy soft hairy below and with few glands,pinnately veined with 10-20 pairs of lateralveins. Inflorescence a glabrous raceme, male

I.

Protomegoborio stopfiono - wild

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one c. 10 cm long, female one c. 4 cm long.Flowers unisexual, regular, 5(-6)-merous; ped-ice1 I-3 min long; sepals triangular, up to 2min long; petals absent; male flowers withstamens 2-3 min long, disk 5-10bed, ovary ru-dimentary; female flowers without disk, ovarysuperior, usually 3-celled, styles (2-)3, 2-branched. Fruit a nearly globose capsule 3-4cm in diameter, often slightly 3-10bed, gla-brous, usually 3-seeded. Seeds obovoid, c. I cmlong, smooth, shiny. Seedling with epigealgermination; hypocoty12-3 cm long, epicoty1 3-5 min long; cotyledons leafy, roundish, c. 4 cmx 3.5 cm; first leaves alternateIn Sierra Leone Protomegoborio stopfionoflowers from March untilthe start of the rainyseason. Fruits are mature about 6 weeks after

floweringProtomegoborio is restricted to West and Cen-tral Africa and comprises 3 species. Records ofProtomegoborio ingcrophyl!o Hutch. and itsuses for West Africa have been based on wrongidentifications as this species is restricted tosouthern Cameroon and Gabon.

Ecology Protomegoborio stopfiono is foundin the understorey of primary and secondarylowland forest. It sometimes forms purestands. It is found both in swamps as well ason well-drained localities. In swamps the treestend to develop adventitious roots

Management Caution is needed at fellingbecause holes often have a spongy heart

Genetic resources and breeding Proto-megaborio stopfiono is widely distributed, of-ten abundant and not much sought after be-cause of its small and often poorly shaped hole.Therefore it is not under threat of genetic ero-

PROTORHUS DITIMENA H. Ferner

Protologue Mein. Mus. natl. Hist. nat. , Pa-ris n. s. 18: 260 (1944).

Family AnacardiaceaeOrigin and geographic distribution Pro-

torhus ditimeno Is endemic to northern and

eastern MadagascarUses The wood of Protorh"s ditimeno and

other Protorhus spp. , known as 'ditimena', isused in Madagascar for construction, joinery,flooring, panelling, moulding and railwaysleepers. The leaves are used in traditionalmedicine as sedative

Properties The heartwood is chestnutbrown and streaked;it is distinctly demarcatedfrom the paler, up to 6 cm wide sapwood. Thegrain is generally straight, texture fine. Thewood is moderately heavy, with a density of750-810 kg/in3 at 12% moisture content. It airdries easily with little checking and warpingThe rates of shrinkage during drying are me-dium, from green to oven dry 3.4-4.6% radialand 9.1-10.3% tangential. At 12% moisturecontent, the modulus of rupture is 156-177N/min2, modulus of elasticity 11,800-14,400N/mm2, compression parallel to grain 58-70N/mm2 shear 7.5-8.5 N/min2, cleavage 21-25N/min and Chalais-Meudon side hardness 5.8

The wood works easily, glues satisfactorily andthe nailing properties are moderate. The woodis moderately durable, with fair resistance totermites, but it is rather susceptible to fungalattacks. Impregnation with preservatives un-der pressure showed fairly good results. Forusage as railway sleepers, It is advised to treatthe wood with preservatives under pressure

Botany Evergreen, small to medium-sized

PROTORHUS 549

SLOn

Prospects The wood of Protomegoborio stop-florid will probably remain of some local jin-portance only

Major references Burki11, 1994; Cooper &Record, 1931; Hawthorne, 1995; Kryn & Fobes,1959; Takahashi, 1978.

Other references Breteler, 2012b; Chave,2005; Dalzie1, 1937; de in Mensbruge, 1966;Hawthorne & Jongkind, 2006; Keay, 1958d;Keay, 1989; Leonard, 1995; Normand &Paquis, 1976; Savill& Fox, 1967

Authors C. H. BOSch

.

Protorhus ditimeno - wild

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tree up to 25 in tall; hole up to 35 cm in diame-ter; bark surface slightly rough, finely flaky,blackish, inner bark reddish, exuding a whitishresin; young twigs yellowish brown short-hairyLeaves opposite to alternate, simple and entire;stipules absent; petiole 5-12 min long; bladeobovate to elliptical, 2-7.5 cm x 2-3 cm, curie-ate to obtuse at base, rounded to notched atapex, leathery, brownish green above and red-dish below, glabrous, pinnately veined with15-25 lateral veins. Inflorescence an axillaryshort panicle, densely reddish brown short-hairy, densely flowered. Flowers unisexual,regular, 4--5-merous, whitish or yellowish, ses-sile; calyx lobes rounded, c. I min long; petalsfree, ovate, 2-3 min long, densely short-hairy;stamens c. 2 min long; disk slightly lobed,hairy; ovary superior, hairy, 3-celled, styleshort, 3-branched; male flowers with rudimen-tary ovary, female flowers slightly larger thanmale ones and with rudimentary stamensFruit an ovoid to oblong drupe up to 3 cm long,slightly grooved, reddish brown short-hairy,with resinous pulp, I-seededProtorhus ditimeno grows slowly. Trees flowerin September-December and fruits mature 2-3months later. The fruits are eaten by birds,which probably serve as seed dispersersProtorhus comprises nearly 20 species, all en-demic to Madagascar except Protorhus longifo-Ito (Bernh. ) Engl. , which is from South Africa.It has been proposed to transfer the speciesfrom Madagascar to a separate genus Abro-hornio. Micronychio, an endemic genus fromMadagascar, is closely related to Protorhus, butdiffers in sigmoid, mango-like fruits.The wood of several other Protorhus spp. isused for similar purposes as that of Protorhusditimeno. One of these is Protorhus grandtdieriEngl. , a small to medium-sized tree up to 25 intallfrom western Madagascar. The gum-resin,bark and leaves are used in traditional medi-

cine, but no details are available. The wood ofProtorhus senceo Engl. , Protorhus IOUuelitH. Ferner and Protorhus thouuenotii Lecomte,all occurring in eastern Madagascar, is similarin properties and uses to that of Protorhus di-timeno. A leaf infusion of Protorhus senceo is

administered to treat indigestion. The leavesare also valued in traditional medicine to treat

heart complaints and as disinfectant.Ecology Protorhus dittmeno occurs in ever-

green forest up to 1250(-1600) in altitude. It iscommonly found along watercourses.

Genetic resources and breeding Proto-rhus dirtmeno is quite widespread, but it oc-

curs scattered in the forest and is limited to

threatened types of forest. Therefore, caution isneeded in harvesting this species to preventgenetic erosion. This is even more the case insome other Protorhus spp. with limited areas ofdistribution.

Prospects Protorhus ditimeno and otherProtorhus spp. have been recommended forenrichment planting in natural forest, and asnon-pioneer light-demanders that also toleratesome shade they are suited for that purposeHowever, they grow slowly and long cuttingcycles are needed for timber production, andtherefore commercial exploitation on a sus-tainable basis seems to have little prospects

Major references Blaser at a1. , 1993; Gu6-neau, Bedel & Thie1, 1970-1975; Ferner de inBathie, 1946a; Rakotovao at al. , en prepara-tion.

Other references Boiteau, Bolteau & A1-10rge-Bolteau, 1999; Gunb-Fakim & Brendler,2004; Sallenave, 1955; Sallenave, 1971; Schatz,2001

Authors R. H. M. J. Lemmens

PSYDRAXPARVIFLORA (atzel. ) Bridson

Protologue Kew Bull. 40: 700 (1985)Family RubiaceaeSynonyms Conthium rubrocostotum Robyns

(1929), Conthium uulgore (K. Schum. ) Bullock(1932)

Origin and geographic distribution Psydroxporui/!o70 is widespread from Senegal east-ward to Ethiopia and Kenya, and southward toAngola, Zimbabwe andMozambique

Uses In Kenya the wood is sometimes used

I.

Psydroxporui/1070 - wild

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for Joinery, interior trim, furniture and imple-merits, and in Tanzania in boat construction. Itis suitable for construction, flooring, mineprops, railway sleepers, ladders, sportinggoods, agricultural implements, tool handles,draining boards, pattern making, turnery, ve-neer and plywood. It is also used as firewoodand for charcoal productionRipe fruits are eaten fresh. Preparations ofleaves and flowers are applied against lice.Root decoctions are taken as arithelmintic and

tonic. Fruit extracts are administered againstcough and influenza. Unknown parts of theplants have been used to prepare medicine totreat rheumatism, oedema and sores. Psydroxparui/10ro is occasionally planted as ornamen-talshade tree. The flowers are visited by honeybees

Production and international trade The

wood is only used locally and not traded on theinternational timber market. The fruits are not

marketed

Properties The heartwood is pale brown todark brown, often with a purplish tinge, andindistinctly demarcated from the paler sap-wood. The grain is straight, texture fine andeven. The wood is moderately heavy, with adensity of 720-800 kg/ms at 12% moisture con-tent, and hard. It dries well and rapidly, withonly occasional surface and end splitting. Therates of shrinkage are moderate. Once dry, thewood is fairly stable in service.The wood saws and works moderately wellwith both hand and machine tools. It is quitebrittle. It holds nails and screws satisfactorilyand polishes, stains and glues well. It has goodturning properties and can be made into good-quality veneer. The wood is moderately dura-ble. The heartwood is very resistant to impreg-nation with preservatives

Botany Evergreen or seml-deciduous shrubor small to medium-sized tree up to 25(-30) intall; bole straight, up to 60 cm in diameter,sometimes fluted; bark surface slightly fissuredor cracked, pale grey to brown, inner barkcream-yellow with orange flecks or darker lay-ers, with a smell of almond; twigs usually an-gular, glabrous. Leaves opposite, simple andentire; stipules triangular, 2-7 mm long; peti-o1e up to I cm long; blade elliptical to ovate,5.5-15.5 cm x 2-8 cm, rounded to curieate atbase, acuminate at apex, leathery, glabrous,pinnately veined with 4-8 pairs of lateralveins. Inflorescence an axillary umbel-likecyme 2-6 cm wide, short-hairy, many-flowered;peduncle up to 2 cm long. Flowers bisexual,

regular, 4-merous; pedicel up to I cm long; ca-Iyx up to 2.5 min long, with lobes shorter orlonger than the tube, glabrous to short-hairy;corolla whitish, tube 2-3 min long, hairy in-side, lobes 1.5-2.5 mm long; stamens insertedat corolla throat, reflexed; ovary inferior, 2-celled, style slender, up to 8 min long. Fruit a2-10bed drupe 0.5-ICm x I-1.5 cm, black whenripe; stones 2, 0.5-I cm in diameter, each I-seeded. Seeds finely reticulatePsydro, c poruiftoro is variable and 4 differentsubspecies have been distinguished. The fruitsare eaten by birds, which disperse the seedsDuiker antelopes have been reported to eattheleaves

Psydrox comprises about 100 species and oc-curs in the tropics of Asia and Africa, with ap-proximateIy 35 species in Africa. It is related toConthium andKeetio

The wood of several other Psydrox spp. is usedin tropical Africa for similar purposes as that ofPsydroxporui/lord, but most of these are moreimportant for their use in traditional medicine.The wood of Psydrox foulhneroe Bridson isused in Kenya for building poles. Psydroxfoulkneroe is a shrub or smalltree up to 7 intall, occurring in woodland and thickets insouth-eastern Kenya and eastern Tanzania.The yellowish brown to reddish brown, heavyand hard wood ofPsydrox o6000to (Notzsch exECkl. & Zeyh. ) Bridson (synonym: Conthiumo6000tum Notzsch ex ECkl. & Zeyh. )is used inSouth Africa in house building, for main postsbut also for walllaths and roof laths. Psydroxo60uoto prevents erosion in coastal dunes. It isa shrub or smalltree up to 15 in tall, occurringin coastal forest, mixed evergreen rainforestand woodland in Zimbabwe, Mozambique andSouth Africa

Ecology In West Africa Psydrox poruiftorois found in dry forest and gallery forest in sa-vanna regions, but in East Africa it occurs inhumid lowland forest and rainforest in the

mountains up to 2000(-2800) in altitude. Insouthern Africa it is found in evergreen rain-forest and woodland up to 2000 in altitude,sometimes on termite mounds. In Ghana trees

of all size classes are more abundant in forest

that is regularly burned than in undisturbedforest

Management In Tanzania ripe fruits arecollected in January-May. Psydro" poruiftorois rarely planted, but can be propagated byseed

Genetic resources and breeding Psydroxparuiftoro is very widespread and locally coin-

PSYDRAX 551

552 TIMBERS2

mon, and not subject to genetic erosion. How-ever, two of the subspecies have restricted are-as of distribution in montane forest, and mightbe more easily liable to habitat degradation

Prospects Psydroxporui/10ro is valuable asa multipurpose tree, providing wood for localapplications but also edible fruits and ingredi-ents for traditional medicine. More research is

needed on its wood properties and pharmaco-logical activities, and to explore its possibihtiesfor use in agroforestry systems

Major references Bolza & Keating, 1972;Bridson, 1985; Bridson, 1998; Chikamai at al. ,undated; Ruffo, Birnie & Tengnas, 2002

Other references Beentje, 1994; Burkill,1997; Coates Palgrave, 2002; Gaugris at al. ,2007; Hawthorne, 1995; Hawthorne & Jong-kind, 2006; Lovett at a1. , 2007; Neuwinger,2000; Verdcourt & Bridson, 1991; Wimbush,1957.

Authors R. H. M. J. Lemmens

Tanzania and Zambia

Uses A yellow to reddish brown fat, called'kombo butter' or 'Angola tallow', is extractedfrom the seed and is important in West andCentral Africa for illumination and in soapmaking. It is not edible. The seeds somewhatresemble those of nutmeg (Myristico frogronsHoutt. ) and are burnt as candles. In CentralAfrica they are used as spice. Traditionally thewood is highly valued as fuel and is used tomake split planks, known as 'calabot' or 'cara-board' in the coastal zone of Garnero0n. Be-

cause it is easy to work, it is used to makeshingles both for roofing and covering the sidesof native houses, and planks for doors andwindow frames. The long straight hole makes itsuitable for making canoes. Since the SecondWorld War the wood has become an importanttimber for plywood corestock, veneer, mould-Trigs, interior trim, Interior joinery, furniturecomponents and paper pulp. In agroforestryPycnonthus origolensis is planted or retainedfor shade in coffee and cocoa plantations in thehumid lowlands of Cameroon, in Uganda oftenalso in banana plantations. Farmers in Came-roon consider it a good indicator of soil fertility.In Uganda it has been planted as an amenitytree

Throughout its area of distribution, variouspreparations of the bark, and to a lesser extentother parts of the tree, are used medicinalIy totreat skin infections, especially of the mouthPreparations made from the bark are used as apotent purgative, to cleanse the milk of Iactat-Ing mothers and to treat coughs and chestcomplaints. In Ghana a decoction of the bark istaken to treat anaemia, in C6te d'Ivoire as apoison antidote and against ascites and Iepro-sy. In Congo the bark is used to treat a numberof gynaecological problems, from infertility togonorrhoea. In C6te d'Ivoire a root maceratemixed with parts of other plants is taken bydraught to treat schistosomiasis. In Sao Tomethe bark is used to treat malaria.

Production and international trade No

information is available on the trade in kombo

butter. Trade in the timber 'nomba' began afterthe Second World War due to an increased

demand for plywood and improvements inwood conservation techniques and also as asubstitute for okoum6 (Aucoumeo hintneonoPierre). Trade in ilomba increased spectacular-Iy between 1946 and 1959 from 100 to 5600boles. Gabon and Cameroon became the first

major exporters in 1952/1953, followed by C6ted'TVoire in 1954 and Congo in 1955. For several

PYCNANTHUSANGOLENSIS ayelw. ) Warb

Protologue Notizbl. Konigl. Bot. Gart. Ber-Iin I: 100 (1895)

Family MyristicaceaeChromosome number2n=38

Synonyms Pycnortthus hornbo (Baill. ) Warb.(1897).

Vernacular names African nutmeg, boxboard(En). nomba, faux muscadier, arbre a suif (Fr)Menebantamo (Po). Mkungu inwitu (Sw)

Origin and geographic distribution Pyc-norithus origolensis is found in the forest zoneof tropical Africa, from Senegal and Guinea toAngola, and through DR Congo to Uganda,

Pycnonthus origolensis - wild

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years, nomba was among the most valued tim-hers in Central Africa. Between 1950 and 1960,the quantity of wood exported from Gabon was3000 in3 from Gamero0n 278,000 ms. Came-roon has enforced a ban on exports of nombalogs since 1999. Exports of nomba have fallendrastically. In 2003 the combined exports ofveneer, sawnwood and plywood from Gamero0namounted to 72 ina, from Gabon to 816 ina. Ex-ports from the Congo basin dropped to 0.06% oftotal timber exports or about 3000 ing in 2003In 2001 11,000 ina of nomba veneer were ex-ported from C6te d'TVoire at an average price ofUs$ 2401m3, and 5000 in3 from Ghana at anaverage price of Us$ 3511m3. The export ofplywood from C6te d'Ivoire in 2001 amountedto 3000 in3 at an average price of Us$ 3291m3,and from Ghana in 2002 to 1000 ms at an av-

erage price of Us$ 4561m3Properties The seeds of Pycnonthus origo-

lensis are aromatic, but information on volatileconstituents is not available. The seeds yield45-70% of a yellow to reddish brown solid fatknown as 'kombo butter', which tastes bitterand is suitable for making soap and candles,while the residue is used for manure as it Isunsuitable as cattle feed. The melting point ofthe fat is 51'C. The fatty acid composition ofkombo butter is lauric acid 5.5%, myristic acid61.5%, palmitic acid 3.6%, myristoleic acid23.6%, oleic acid 5.7%. Crude kombo buttercontains about 20% kombic acid (a diny-droxymethylphenyl derivative of hexadecatet-raenoic acid) and sargaquinoic acid (a quinonederivative) and several of their derivativesThese terpenoid quinonic acids have promisinganti-oxidant properties for pharmacology, cos-metics and the stabilization of plastics. Theyhave also shown hypoglycaemic activity in dia-betes patientsThe bark contains dihydroguaiaretic acid,which has shown non-selective toxicity towardsseveral human tumour celllines. Extracts ofthe bark also showed the presence offIavonoids(2'-hydroxy-formononetin), tannins and sapo-rim glycosides, which might be responsible forits biological activities. Terpenoid quinonesthat have shown hypoglycaemic activity in bothinsulin-dependent and insulin-independentdiabetes have been extracted from the hole andleaves.

The heartwood is whitish to pinkish brown,sometimes with yellowish markings and iridis-tinctly demarcted from the sapwood. The grainis generaUy straight, the texture medium tocoarse. The wood has no lustre and when fresh-

Iy sawn it has an unpleasant odour which dis-appears on drying.At 12% moisture the density is 440-570 kg/inaThe wood is rather difficult to dry, it is prone tocollapse, end splitting and distortion. Goodventilation is required for air drying. Kiln dry-ing can give good results if done carefully.Shrinkage rates from green to oven dry are4.6% radial and 8.4% tangential. Drying ofbeams more than 55 mm thick is very difficultand steaming for 2 days is recommended. At12% moisture content, the modulus of ruptureis 62-72 N/mm2 modulus of elasticity 8300-12000 N/mm2, compression parallel to grain38-39.5 N/mm2, shear 5.4-8.9 N/min2, cleavage13-24 N/mm, Janka side hardness 2700-3400N

The wood is easy to saw and plane with normaltools; blunting effects are moderate. It is diffi-cult to polish. Nailing and screwing are easyand holding properties are good. The wood maystain in contact with tools. It peels and sliceswell to produce good-quality veneer and ply-wood, although steaming is recommended be-cause of the occasional presence of numeroussmallhard spots. It glues well with antypes ofglue. It paints wellbutis rather absorbentThe wood is not durable and liable to attack bytermites, powder-post beetles, pinhole borersand marine borers, but it is permeable to pre-servatives

Description Evergreen, monoecious or dioe-CIOus, medium-sized to large tree up to 25-35(-40) in tall; hole usually straight and cylindrical,branchless for up to 15(-25) in high, up to120(-150) cm in diameter, usually withoutbuttresses; outer bark greyish brown, with

crown small, withorange-brown exudate;branches at right angles to the bole; twigsslender, pendulous, densely rusty hairyLeaves distichously alternate, simple and en-tire, without stipules; petiole I-2 cm long;blade oblong to oblong-lanceolate, 7.5-30(-40)cm x 4.5-Ile16) cm, base cordate, apex acu-inmate, dark green above, glaucous below,young leaves velvety reddish brown hairy, butglabrescent, pinnately veined with 20-40 pairsof lateral veins. Inflorescence an axillary pant-cle, often on leafless branches, 10-30 cm long,rusty hairy, with flowers in numerous head-shaped clusters. Flowers unisexual, regular,very small, sessile, with 3-lobed perianth cov-ered with dark brown hairs; male flowers with2-4 stamens, filaments merged into a column;female flowers with superior, sessile, I-celledovary, stigmas 2, sessile. Fruit an ellipsoid to

PYCNANTHUS 553

554 TIMBERS2

3

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SIonally used.Anatomy Wood-anatomical description OAWA

hardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; (14: scalariformperforation plates); (15: scalariform perforationplates with s 10 bars); 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 prn); 27:intervessel pits large (Z 10 F1m); (31: vessel-raypits with much reduced borders to apparentlysimple: pits rounded or angular); 32: vessel-raypits with much reduced borders to apparentlysimple: pits horizontal(scalariform, gash-like)to vertical (palisade); 42: mean tangential di-ameter of vessellumina 100-200 pin; 46: S 5vessels per square minimetre; 47: 5-20 vesselsper square minimetre; 56: tyloses common.Tracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibrespresent; (68: fibres very thin-walled); 69: fibresthin- to thick-walled. Axial parenchyma: 75:axial parenchyma absent or extremely rare; 78:axial parenchyma scanty paratrachea1; 93:eight (5-8) cells per parenchyma strand; 94:over eight cells per parenchyma strand. Rays:97: ray width I-3 cells; 102: ray height > Imm; 107: body ray cells procumbent with most-Iy 2-4 rows of upright and/or square marginalcells; 108: body ray cells procumbent with over4 rows of upright and/or square marginal cells;109: rays with procumbent, square and uprightcells mixed throughout the ray; 1/5: 4-12 raysper min. Secretory elements and cainbial van-ants: (132: Iaticifers or tanniferous tubes)Mineral inclusions: 152: crystals of othershapes (mostly small).on. Thiam, P. Derienne & E. A. Wheeler)

Growth and development Seeds of Pyc-norithus origolensis are recalcitrant. The dura-tion of germination is 16-36 days. The cotyle-dons are pulpy and the first two leaves whichappear after two months are SLmple, oppositeor alternate, later leaves alternate. A deep sec-ondary root system develops during the firstseven months of growth. In natural standsnumerous seedlings appear around the mothertree. In the first year the stem height reaches20-30 cm and it can reach 50 cm in the secondyear. In Sierra Leone a mean annual incre-merit in diameter of 1.6-2.4 cm has been ob-served. Because of the long straight trunk, thevolume/trunk ratio is higher than in most otherAfrican forest tree species. Pycnonthus origo-lensis is evergreen, and at any latitude in its

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Pycnunthus origolensis - I, 60se o1601e, 2, led/ytwig, . 3, inflorescence, 4, fruit, ' 5, seed.Redrot{!it grid adopted by Ishoh Syumsudin

oblong or globose drupe, 3-4.5 cm x 2-4 cm, inbunches, yellowish orange when ripe, fruit wallrather hard and tough, 2-10 min thick, split-ting longitudinalIy with 2 valves, I-seededSeed ellipsoid, aromatic, 1.5-3 cm x I-1.5 cm,dark brown, with pink to red anI, Iaciniatealmost to the base. Seedling with epigeal ger-mination, but cotyledons remaining in the tes-ta.

Other botanical information Pycnonthuscomprises 3-4 species, all in Africa. Pycnon-thus origolensis is variable, especially in thehairiness of the leaves, the size and shape ofthe fruits, and reportedIy also in the quality ofthe timber. Two subspecies have been distin-guished: subsp. origolensis and subsp. schwein-furthit (Warb. ) Verdc. , the latter occurring DRCongo and East Africa, but possibly also moreto the west, and differing from subsp. origolen-sis in having larger, often more globose fruitswith thicker fruit wall. The wood of Cepho!0-sphoero usQinborensis (Warb. ) Warb. and sev-eral American Virolo species closely resemblesthat ofPycnonthus origolensis. Cepholosphoerousermborensis is restricted to eastern parts ofKenya and Tanzania, where its timber is occa-

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range leaf fall and flushing occur SImultane-ously. The flowering period is long and dependson the location. In Cameroon it flowers in Oc-

toher-May with male and female flowers atseparate parts of the same tree, generally alsoat differenttimes, while it fruits in September-April. Dehiscence takes place on the tree or thewhole intructescence falls before dehiscence

Ecology Pycnonthus origo!errsis occurs inupland and wet evergreen forest and semi-deciduous forest with more than 1600 minrainfall. It is especially abundant in old fallowsand secondary forest as its rate of natural re-CTuitment after disturbance of the forest is

high. In southern Africa it occurs in rivermeand swamp forest, but in West Africa it doesnot occur in swamps. In Uganda it also occursin gallery forest. It is mostly found in smallgroups or solitary and it regenerates in smallto medium-sized gaps in the forest. Its abun-dance increases with rainfall, the optimumbeing about 2000 min/year; above 2600mm/year numbers decline strongly. It occa-SIonally occurs where rainfallis only 1300 mmor less with 4-5 dry months. Seedlings are verysensitive to drought. Pycnonthus origolensis isa light-demanding tree typical of the dominantforest strata, although it can tolerate slightshade when young. It occurs up to 1200(-1400)in altitude. Pycnonthus origo!errsts tolerateslight and heavy soils, but is scarce on sandysoils, while other reports indicate that it isoften found on poor soils.

Propagation and planting Pycnonthus on-gole, 1sts is propagated by seed. There are about500 seeds per kg. Young broken or cut treesresprout easily, but in a trial vegetative propa-gation by stem cuttings failed to succeed. Seedsshould not be dried, but sown as soon as possi-ble because of their short viahinty. Germina-tion is easy and with proper care the germina-tion rate offresh seed can reach 100%. Soakingin cold water for 24 hours hastens germination.In the case of unsorted seeds, the germinationrate is about 50%. Seeds can be planted direct-Iy in the field or in an open field nursery pref-erably in polythene bags. It is important toprotect the seeds from rodents. A mixture ofsand and arable soil(50150) is a suitable ger-ruination substrate. The seedling rapidly growsa large taproot, whose development should bechecked timely in the nursery. Cutting thetaproot when it is large greatly reduces theplant's growth rate. It is advisable to trans-plant seedlings after I-2 years when 30-50 cmtall, at the beginning of the rainy season. A

slight mulching is recommended. In the humidlowlands of Gamero0n farmers used to retain

or transplant seedlings from the wild whenclearing new fields. To improve growth, coin-post or chemical fertilizer may be applied. Indirect sowing in the field, the recommendationis to plant 3-5 seeds per hole and thin to a sin-gle plant after germination. Field spacing hasbeen 4 in x 5 in, but recent recommendationsare 9 in x 10 in (110 treesA1a)

Management Protection and retention ofnatural Pycnonthus origo!errsis trees has longbeen done by farmers in the humid lowlandforest of West and Central Africa. In planta-tions the initial thinning should be done whentrees are about 7 years old to reduce the densi-ty to 300-350 trees/ha; when trees approachthe age of 12 years a second thinning shouldreduce the density to 150-200 trees/ha.

Diseases and pests Although its leaves areoften marred by small holes, no Important dis-eases or pests have been detected in Pycnon-thus origolensis in either the natural state orplantations and from a phytosanitary point ofview, SIIviculture of the species is very easyNevertheless, some sporadic insect (Monocho-mus SCObtosus, Mol!odori dowriesi, Bryochoetointerrupto) and fungi(Ophiostomo sp. , Micro-thyriel!o sp. ) attacks have been reported inC6te d'Ivoire, Gamero0n and Gabon

Harvesting In good plantations in the ever-green forest zone the exploitable diameter of 50cm is reached when trees are 30 years old, anda diameter of 60 cm at 45 years

Yield Little information on seed yield isavailable; an average tree may produce 60-100seeds annually. In wellgrowing plantations theannual increment at 15 years of age is 15ms/ha/year, at 30 years it can be 10 mama/year

Handling after harvest Logs should betreated with preservatives and be convertedsoon after felling to avoid discoloration by fungiand damage by insects. Logs can be floated andbe transported by river

Genetic resources Because of its wide dis-tribution and occurrence in secondary forest,there is little risk of genetic erosion. No geneticconservation programme is known to exist.

Breeding Pycnonthus origolensis is one ofthe most important agroforestry tree species ofthe humid lowland forest of West and CentralAfrica identified by the World Agroforestry Gen-tre (ICRAF) for a domestication programme.

Prospects Pycnonthus origolensis is an jin-portant medicinal plant in the humid forestregion. It is traditionally protected by farmers

PYCNANTHUS 555

556 TIMBERS2

during forest clearing. Large amounts of tim-her have been exported, but recently volumeshave dropped markedIy. The export of the woodas veneer and plywood has been most jin-portant in recent years. New opportunities forexploiting the oil and medicinal propertiesshould be investigated. However, as a fairlyfast-growing species that is not very liable todiseases and pests, Pycnonthus origolensisseems to have good prospects for timber plan-tations and for sustainably managed naturalproduction forest

Major references Bone, 2000; CTFT, 1975;Duguma, Tonye & Depommier, 1990; Katende,Birnie & Tengnas, 1995; Mapongmetsem at al. ,1995; Mapongmetsem at a1. , 1999a; Mapong-metsem, Nkongmeneck & Duguma, 2002;Normand & Paquis, 1976; Richter & Danwitz,2000; Verdcourt, 1997

Other references Adjanohoun at a1. , 1991;Adjanohoun at a1. , 1996; ATIBT, 2004; Berhaut,1979; CTFT, 1961a; Dalzie1, 1937; Dounias,1995; Forest Product Laboratory, 1999; In-sideWood, undated; Irvine, 1961; Laird, 1999;Letouzey, 1955; Luo at a1. , 1999; Mapongmet-sem at a1. , 1999b; Mique1, 1985; Perez at al. ,2005; Pope, 1997; Raponda-Walker & Sillans,1961; SImon at a1. , 2005; Taylor, 1960; Vabi &Mala'a, 1995; World Agroforestry Centre, un-dated

Sources of illustration Vetdcourt, 1997;Voorhoeve, 1965; Wilks & TSSemb6, 2000

Authors P. -M. Mapongmetsem

,.

RAWSONIALUCIDA Harv

Protologue Harv. & Sond. , F1. cap. I: 67(1860).

Family F1acourtiaceae (APG: AChariaceae)Synonyms Rott!sonio reticulata GIIg (1901).Vernacular names Forest peach (En).Origin and geographic distribution Rowso-

itto lucido occurs from Sudan east to Somaliaand Kenya and south to Angola, Zimbabwe,Mozambique, eastern South Africa and Swazi-land.

Uses The wood is locally used for walkingsticks, pestles, spoons and tool handles, andalso as firewood and for charcoal productionThe fruitis recorded as edible.

Properties The pale pink to reddish wood isheavy, hard and tough. The cyanogenic glyco-sides gynocardin and cyclopentenylglycinehave been isolated from the leaves

Botany Evergreen shrub or small to medi-

Rowsonio lucido - wild

urn-sized tree up to 20 in tall; hole up to 50 cmin diameter, sometimes fluted at base or withbuttresses; bark surface smooth, peeling inrounded scales, grey to reddish brown, innerbark red; crown with spreading branches;twigsshort-hairy to glabrous. Leaves alternate, sim-PIe; stipules narrowly oblong, c. 5(-10) minlong, caducous; petiole 0.5-I(-1.5) cm long;blade oblanceolate to oblong-obovate, 7-20 cmx 2-8.5 cm, curieate to obtuse at base, acumin.ate at apex, margins usually distinctly toothed,thin-leathery, glabrous, glossy, pinnately veinedwith 6-9 pairs of lateral veins. Inflorescence anaxillary raceme up to 2.5(-4) cm long, short-hairy to glabrous, densely flowered. Flowers bi-sexual or male, regular, scented; pedicelup to 2mm long; sepals 4-6, free, unequal, rounded toelliptical, (2-)3-4(-5) min long, short-hairy onmargins; petals 4-8, similar to sepals but 5-8min long and with basal hairy appendage in-side; stamens numerous, up to ICm long; ovarysuperior, glabrous, I-celled, style up to 2 mmlong, with 3-4 short branches. Fruit a globoseberry-like capsule 2-2.5(-4) cm in diameter,short-pointed, glabrous, yellow becoming red-dish, tardily dehiscent with 45 valves, up to10-seeded. Seeds ovoid to nearly globose, 6-10min long, rough.Rowsonio lucido sometimes flowers when only3 in tall. In southern Africa trees flower in Sep-ternber to November, and fruits ripen in No-veinber to February. They have been reportedto fruit in Kenya in June, ~uly. Trees may pro-duce masses of fruits that may carpet the for-est floor in the fruiting season. They are eatenby monkeys, which probably disperse theseeds.

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Rowsonio comprises 2 species and is related toDOSylepis, Erythrospermum and Scottellio. Row-sonio burtt-douyi(Edlin) Finiite is a small tomedium-sized tree up to 20 in tall, with hole upto 30 cm in diameter, endemic to Mt Mulanjein Malawi

Ecology Rowsonio lucido is most character-istic for inId-altitude and montane semi-

evergreen forest up to 2200 in altitude, but italso occurs in the lowlands. It is found in dry tomoist evergreen forest and in riverme forest,usually in the understorey

Management Rowsonio lucido can be prop-agated by seeds and cuttings. It is recommend-ed to soak seeds in water for 24 hours before

sowing. They should be covered with I cm ofsoil and kept at about 25'C, constantly moistbut not wet. The seeds germinate in 3-6 weeks.

Genetic resources and breeding Row-sonio lucido is widespread and locally common,and certainly not endangered by genetic ero-SLOn.

Prospects The wood of Rowsonio lucido willremain of limited importance for local applica-tions. The small size of the bole makes that it

has no prospects for the international timbermarket. As an evergreen tree with shinyleaves, lt may have prospects as ornamental.

Major references Coates Palgrave, 1983;Dowsett-Lemaire & White, 1990; Lovett, Ruffo& Gereau, 2003; Palmer & Pitman, 1972-1974;SIeumer, 1975

Other references hadersen at a1. , 2001;Barnps, 1968; Beentje, 1994; Chase at a1. , 2002;Enge1, 2000; Hyde & Wursten, 2011; In-sideWood, undated; Wild, 1960; Wimbush,1957

Authors R. H. M. J. Lemmens

.

RHABDOPHYLLUM 557

Rhobdophyl!urn colophyl!urn - wild

leaves taken together with fruits of Afro-momum melegueto K. Schum. are consideredaphrodisiac

Production and international trade The

wood of Rhobdophyllum CUIophy!!urn is onlyused locally.

Properties The pinkish brown wood is hard,flexible and durable.

Botany Erect shrub or smalltree up to Toe15) in tall; hole up to 15 cm in diameter; barksurface smooth, grey; twigs robust, glabrousLeaves alternate, simple and entire; stipulesfused at base, falling early; petiole 3-6 mmlong, slightly corky; blade narrowly elliptical toelliptical-obonate, (6-)9-20(-25) cm x 2.5-8.5cm, base curieate, apex acute or acuminate,leathery, pinnately veined with very numerouslateral veins. Inflorescence an axillary falseraceme (4-)6-11 cm long, glabrous, denselyflowered, with fascicles of (2-)3-5 flowersFlowers pendant, bisexual, regular, 5-meTous;pedice1 7-10 min long; sepals free, oblong, 7-10(-12) mm x 2.5-6 min, persistent in fruit;petals free, elliptical-obovate, 6-8 min long,yellow or cream, falling early; stamens 10, sur-rounding the style, arithers nearly sessile; car-pels 5, free, style I, long and slender. Fruitpendant, consisting of up to 5 globose, blackdrupelets, each drupelet I-seeded, with redsepals at base.In Ghana flowering occurs in December, andfruits mature in January-February.Rhobdophyllum comprises 8 species and is res-tricted to tropical Africa, where it occurs fromGuinea to Zambia. It is related to Campylo-spermum and Idertio

Ecology Rhobdophyllum colophyl!urn occurs

RHABDOPHYLLUM CALOPHYLLUM (Hook. f. )Tiegh.

Protologue Bull. Mus. natl. Hist. nat. , Paris8 : 216 (1902).

Family OchnaceaeChromosome number it = 12

Synonyms Ouroteo colonhyllo (Hook. f. ) Englex Gilg (1893).

Origin and geographic distribution Rhob-dophy!Ium colophyllum occurs from Guinea toGhana, and in Garnero0n, Equatorial Guinea,Gabon, Congo and Cabinda (Angola)

Uses The wood of Rhobdophy!Ium CUIophyl-Ium is used for poles in house building and fortool handles, and also as firewood. In Congo

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558 TIMBERS2

in primary and secondary evergreen and semi-deciduous forest on well-drained soils, butsometimes also in swampy localities and occa-SIonally in shrub vegetation near the coast oreven in savanna, up to 750 in altitude. In Gha-na it is mainly found on mild hill slopes in ev-ergreen forest

Genetic resources and breeding Rhobdo-phyllum CUIophyllum is widespread and coin-mon, and there are no indications that it is indanger of genetic erosion

Prospects The wood of Rhobdophyllum co-lophyllum is likely to remain of limited, localuse only. The tree deserves attention as anornamental

Major references Aubr6ville, 1959a; BurkiU,1997; Farron, 1965; Keay, 1954e; SOSef, 2008

Other references Bouquet, 1969; Cooper &Record, 1931; Hawthorne, 1995; Hawthorne &Jongkind, 2006; Irvine, 1961; Neuwinger, 2000;Raponda-Walker & Sillans, 1961

Authors L. FA. Oyen

dendron heude!otii occurs from southern Sene-

gal eastwards to Kenya, and southwards toAngola and Mozambique

Uses The seeds of Ricinodendron heude!otii

are widely used in cooking in West and CentralAfrica. An edible o111s extracted from the seeds

and a paste made by crushing dried kernels issometimes used as a thickening agentfor soupsand stews. A paste from the dried and poundedkernels is also stored for making porridge intimes of food shortage. The protein-rich leavesare eaten as a cooked vegetable with dried fishand are used as forage for goats and sheep.The wood, called 'erimado' or 'essessang' intrade, is very light, soft and perishable, but isoccasionally used in carving and for makinghousehold utensils, furniture, boxes and crates.In Uganda the Semliki and Unyoro people useit for making doors for their huts, while insouthern Nigeria and DR Congo well-soundingdrums are carved from it. It is a potential sub-stitute for balsa wood (Ochromo pyramidole(Cav. ex Lain. ) Drb. ) for making floats and life-belts. The wood is also suitable for boat build-

ing, sporting goods, toys and novelties, hard-board, particle board, plywood, wood-wool andwood-pulp. The ash of the wood is used as veg-etable salt in cooking, indigo dyeing and soapmaking. The seeds are used in rattles and ascounters in games. In Bas Congo (DR Congo)the tree is planted to attract edible caterpillars(lin 670sio epimetheo), and several other ediblecaterpillars are collected from it. The leavesare used as wrapping material and for mulch-ing. In DR Congo Ricirtodendron heudelotiiisplanted as amenity tree, as live fence and forerosion control.

Many parts of the tree are used in medicine.Bark of the root and stem is used in decoctions

or lotions to treat constipation, cough, dysen-tery, rheumatism, rickets in children, oedema,elephantiasis, fungal infection, blennorrhoea,painful menstruation, and to prevent miscar-nage, relieve pain in pregnant women, cureinfertility in women, give strength to prema-ture babies, and to mature abscesses, furunclesand buboes. The sap is instilled into the eyeagainst filaria and ophthalmia and leaf decoc-tions are used as rebrifuge. Leaves are alsoused to treat dysentery, female sterility, oede-ina and stomach pain. Roots are used as aph-rodisiac in C6te d'TVoire. Fruits and latex are

used in West Africa to cure gonorrhoea anddiarrhoea

Production and international trade Ker.nels of Ricinodendron heudelotii are traded in-

RICINODENDRONHEUDELOTll(Baill. ) Pierreex Heckel

Protologue Ann. Inst. Bot. -Geol. Colon.Marseille 5(2): 40 (1898).

Family EuphorbiaceaeChromosome number2n=22

Synonyms Ridnodendrongiftconum Milll. ATg.(1864).

Vernacular names Groundnut tree, cork-wood tree, African oil-nut tree (En). BSsang,essessang (Fr). Menguela, inunguella (Po).Muawa (Sw)

Origin and geographic distribution Ridno-

..

Ricinodendro, " heudelotii- wild

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ternationally and are found in many marketsin West and Central Africa; they are exportedto Europe from Cameroon as 'ridjanssang'. Thehumid forest zone of Cameroon appears to bethe main production area. In 1995, 36,000 kg ofseeds were marketed in this zone, for a totalvalue of about Us$ 79,000.

Properties The dried seeds of Ricirtoden-dron heudelotii contain on average per 100 gwater 6 g, energy 2200 kJ (530 kca}), protein 21g, fat 43 g, carbohydrate 23 g, Ca 611 ing, P926 ing, Fe 0.4 ing, thiamin 10 Fig, riboflavinand niacin traces (Leung, Busson & Jardin,1968). Some sources give a fat content of up to60%. The fat is pale yellow and liquid butsomewhat viscous at ambienttemperatures. Itsfatty acid composition is: palmitic acid 6-10%,stearic acid 6-7%, oleic acid 7-9%, linoleic acid28-36%, u-eleostearic acid 30-51%. The fatalso contains small amounts of ^-eleostearicacid, catalpic acid, gadoleic acid and jignocericacid. When exposed to air in a thin layer itdries to a frosted film; when the oilis heatedfirst to 280'C it dries to a hard clear filmThe heartwood is whitish to pale yellow, and isnot differentiated from the sapwood. The wooddarkens on exposure to light. The grain isstraight, texture coarse and even. The wood islight-weight with a density of 130-300 kg/ms.soft and brittle. It dries rapidly and with littleor no degrade. The shrinkage rates are low:from green to oven dry 1.9-2.4% radial and4.7-5.4% tangential. At 12% moisture content,the modulus of rupture is 29-46 N/min2, inodu-Ius of elasticity 3700-4800 N/mm2, coinpres-SIon parallel to grain 14-21 N/mm2, shear 2.2-3.2 N/mm, , cleavage 5.0-7.3 N/mm (tangential)and Chalais-Meudon side hardness 0.2-0.6

The wood saws and works easily, and nailswithout splitting, but turning and planing aredifficult. The wood is liable to decay and attackby termites, powder-post beetles and marineborers. The wood is permeable to preservatives

Description Deciduous, dioecious, medium-sized tree up to 30(-45) in tall; hole straightand cylindrical, up to 120(-150) cm in diame-ter, base with short, thick buttresses often ex-tending into heavy superficial roots; outer barksmooth at first, becoming rough and fissured,grey; inner bark pink to red, densely inottledand granular; crown candelabra-shaped, coin-monly with many broken branches; twigs withfew lenticels, densely brown stellate hairy butsoon glabrescent, with thick pith. Leaves alter-nate, palmately compound with (3-)5-7(-8)leaflets; stipules fan-shaped, I-5 cm x 1.5-4

RICiNODENDRON 559

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Ricinodendron heudelotii - I, base of 601e, 2,port of 670nch with young fruits, . 3, mole flow-er, 4, fruit, 5, seedRedrown and odopted bylshok Syomsudin

11"' 11' "!

II I

cm, with gland-tipped teeth, persistent; petioleup to 5-30(-40) cm long; leaflets obovate toelliptical-lanceolate, median leaflet 10-30 cm x5-15 cm, lateral ones sinaUer, base curieate,apex long-acuminate, margin almost entire toshallowly glandular-toothed, thinly papery,glabrous above, glabrous to densely stellatehairy below. Inflorescence a terminal panicle,densely stellate hairy but glabrescent; bractsawl-shaped to linear, 3-7 min long; male inflo-rescence up to 40 cm long; female one up to 20cm long. Flowers unisexual, regular, (4-)5-merous, pedicellate; sepals fused at base, c. 4mm long, densely stellate hairy; petals lateral-Iy coherent, oblong, c. 6 mm long, greenishwhite to pale yellow; disk lobes yellowish; maleflowers with 6-14 stamens c. 6 mm long; re-male flowers with superior, globose ovary, 2-3-celled, stellate hairy, styles 2-3, bifid. Fruit a2-3-10bed drupe 2.5-3.5 cm x 4-5 cm, greenwhen young, black when ripe, each lobe con-taming one I'Seeded stone. Seeds globose, c1.5 cm in diameter, reddish brown to black.Seedling with epigeal germination; hypocotyl

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560 TIMBERS2

up to 20 cm long, epicotyl short; cotyledonswith petiole 1.5-2.5 cm long, blade leafy, 6-7cm x 5-6 cm, glandular at margins, palmatelyveined; first leaf 3-lobed

Other botanical information Ridnode, I-

dron comprises a single species. It is closelyrelated to Schirtziophyton. In Rictnodendronheude!otit 2 subspecies are recognized: subspheudelotii occurring from Senegal to Benin,and subsp. o17iconum (Mtill. Arg. ) J. Leonardfrom Nigeria eastwards and southwards. Theformer has mostly 3-lobed fruits, in the latter2-10bed fruits are more common.

Anatomy Wood-anatointcal description OAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 27: intervessel pits large (Z 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; 43: mean tangen-tialdiameter of vessellumina z 200 prn; 46: S 5vessels per square minimetre; 56: tyloses coin-mon. Tracheids and fibres: 61: fibres with sim-

PIe to minutely bordered pits; 66: non-septatefibres present; 68: fibres very thin-walled. A, :I-al parenchyma: 76: axial parenchyma diffuse;77: axial parenchyma diffuse-in-aggregates; 93eight (5-8) cells per parenchyma strand; 94over eight cells per parenchyma strand. Rays:96: rays exclusively uniseriate; 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; (109: rayswith procumbent, square and upright cellsmixed throughout the ray); 1/5: 4-12 rays permin. Mineral inclusions: 136: prismatic crystalspresent; 141: prismatic crystals In non-

chambered axial parenchyma cells; 154: morethan one crystal of about the same size per cellor chamber

(N. P. Monel, P. 06tienne & E. A. Wheeler)Growth and development The roots ofRi-

cmodendron heude!otii reach deep and causelittle competition for nutrients and water inthe upper soillayers with adjacent crops. Thetree starts bearing fruits at 8-10 years of age.In Sierra Leone flowering takes place in April-May, and fruits are produced in September-October; trees are leafless for a few weekswhen the fruits fall. In central Cameroon fruits

are collected in July-September. Bats, horn-bills and rodents are believed to contribute to

the dispersal of the seed. Fruits also breakopen and scatter their seed when they fall onthe ground

Ecology Ricinodendron heudelotii occurs inclearings in rainforest; it is characteristic ofhumid secondary forest and common in aban-doned farmland at 200-500 in altitude. The

inlnimum annual rainfall required is about1000 mm, but annual rainfall may be as highas 10,000 min/year as in Dibunscha, CameroonIt is a fast-growing and light-demanding tree,requiring mean annual temperatures of 18-32"C. Ricinodendron heudelotii prefers medi-urn-textured and freely draining acidic soils.

Propagation and planting Seeds start ger-minating 3-6 weeks after sowing. Scarificationbefore sowing accelerates germination. Vegeta-tive propagation is possible by rooting of leafystern cuttings, layering and side grafting.

Management There is stilllittle experiencewith management of planted Ricirrodendronheudelotii. Trials are in progress at ICRAF,Garnero0n. In DR Congo stakes are sometimesplanted to create a live fence as they easilystrike root. Although the species loses itsleaves during the dry season, some farmers inCameroon use it to shade cash crops such as

Coppicing is possible, but reports onregrowth are contradictory

Diseases and pests Some caterpillars havebeen reported to defoliate Ricinodendron heu-delotii in DR Congo, such as Lobobunoeophoe-duso, Jinbrosio spp. and one identified IOCaUyas 'inimpemba'. However, these caterpillarsalso constitute a considerable protein supplyfor local people. In Gamero0n, a psyllid (Dieli-dophiebio rugnt), and aphids have been report-ed to cause serious damage to young plants.

Harvesting Fallen fruits are collected fromthe ground.

Handling after harvest After collection,the fruits are left to rot in big piles. Once thefruit pulp is rotten, the stones are extracted bywashing and boiling the fruits vigorously. Thenthe stones are removed from the hot water, putin cold water and left overnight. They areboiled vigorously once more until the shellscrack. Shells are then removed using a knifeAfter extraction, seeds are dried.Logs felled for timber should be extracted fromthe forest and converted rapidly because theyare prone to staining.

Genetic resources Ridnodendron heude-lotiiis very widespread in tropical Africa and

cocoa

genetic variation is large. Within a sample of47 accessions, considerable variation was foundin fruit size, seed size and oil content of theseed (49-63%). Because of its wide distributionand prevalence in secondary forest and onfarmland there is no risk of genetic erosion. Nogermplasm collections are known to exist.

Breeding Domestication of Ricinodendronheudelotii has started recently under the TreeDomestication Program of the World Agrofor-estry Centre (ICRAF) in Cameroon. Selectionwork is still in its infancy. Plant characterspreferred by farmers have been identified.They include high yield, long fruiting season,stable yield, thin shell, self-cracking stories andgood taste. It appears that fruit size is onlyweakly correlated with seed size. The self-cracking shell character is not related to shellthickness

Prospects Continuing intensification ofagriculture in humid tropical Africa will in-creasingIy rely on domesticated, fast-growing,multipurpose tree species that fit well in agro-forestry systems. Ifselections can be made thatmeet farmers' requirements and if appropriatepackages of management practices can be de-veloped, Ricinodendron heude!otiiis likely tobecome a more important component of suchsystems and contribute to the regional demandfor edible and industrial oil.

Major references kingbogu, 1996; Ayuk ata1. , 1999a; Fondoun, Tiki Manga & Kengue,1999; Franzel, Jaenicke & Janssen, 1996; Lat-ham, 2004; Ndoye, Ruiz-Perez & Eyebe, 1998;Ngo Mpeck at a1. , 2003; Shiembo, Newton &Leakey, 1997; Tchoundjeu & Atarigana, 2006;TikiManga at a1. , 2000.

Other references Beentje, 1994; Berhaut,1975a; Burki11, 1994; Firestone, 1999; Inside.Wood, undated; Katende, Birnie & Tengnas,1995; Leonard, 1962; Leung, Busson & Jardin,1968; Radcliffe-Smith, 1987a; Richter & Dan-witz, 2000; Tabuna, 1999; Tane, 1997; Tchie-gang at a1. , 1997

Sources of illustration Govaerts, Frodin &Radcliffe-Smith, 2000; Radcliffe-Smith, 1987a;Wilks & TSSemb6, 2000.

Authors Z. Tchoundjeu & A. R. Atarigana

.

RINOREA 561

Rinore0 670chypetolo- wild

red 670chypetolo is widespread in tropical Mri-ca from Guinea east to Kenya, and south toZambia and Angola.

Uses The wood of Rinoreo byochypetolo isused in Kenya and Uganda for hut construc-tion, walking sticks and knobkerries. In DRCongo it is used in construction and to makehousehold utensils. Boiled leaves are eaten as a

vegetable and the dried, powdered leaves aresniffed to relieve headache.

Properties The wood is yellowish white,fine textured and medium-weight.

Botany Shrub or smalltree up to 6.5 in tall;bark greyish green; twigs hairy or glabrousLeaves arranged spiralIy, simple and entire;stipules awl-shaped, c. 8 mm long, early cadu-cous; petiole 0.5-5.5 cm long; blade obovate tooblong-elliptical or elliptical-oblanceolate, 6-23cm x 3-10 cm, rounded to curieate at base,acute or shortly acuminate at apex, nearly gla-brous, pinnately veined with 6-10 pairs of Iat-eral veins. Inflorescence a terminal, many-flowered thyrse up to 17 cm long, short-hairyFlowers bisexual, regular, 5-merous, nodding,cream-coloured to greenish yellow; pedice1 I-2min long; sepals free, broadly ovate to oblong,2-4 mm long, thick, usually short-hairy; petalsfree, oblong to obovate, 4-6 mm long, thick, tipstrongly recurved; stamens c. 3.5 mm long,filaments fused into a ring, arithers with alarge central appendage and 2 smaller lateralappendages; ovary superior, obovoid, glabrous,style 3-4 mm long, thickened upwards. Fruit a3-lobed capsule I-1.5 cm long, 3-6-seededSeeds angular, c. 6 min x 5 min.Rinoreo is a large pantropical genus of shrubsand smalltrees with about 340 species recog-

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RINORBABRACHYPETALA (Turcz. ) Kuntze

Protologue Revis. gen. PI. I: 42 (1891).Family ViolaceaeSynonyms RinoreopoggeiEng1. (1902)Origin and geographic distribution Rino-

562 TIMBERS2

nized at present, most of them occurring inAfrica. Madagascar has about 20 species, near-Iy all endemic. For continental Africa a propertaxonomic study is long overdue. There areabout 50 well-defined species in the NeotropicsRinoreo orboreo (Thouars) Baill. (synonym:Alsodeio orboreo Thouars), called 'inkandaa-inwitu' in Swahili, is found in coastal Kenya,Tanzania and Mozambique, as well as in Mad-agascar. The wood is used for hut building andto make toolhandles. In Kenya a root decoctionIs drunk to cure stomach pain.Rinoreo aylmeri Chipp is restricted to Guinea,Sierra Leone, Liberia and C6te d'TVoire. Thewood is used for walking sticksRinoreo incifoliu (011v. ) Kuntze, called 'inku-rute' in Swahili, is widespread in tropical Mri-ca from Guinea east to Kenya, and south toAngola and Mozambique. The hard and heavywood Is used in Uganda to make wooden ham-mers, tool handles and walking sticks. In WestAfrica decoctions of the whole plant are drunkto cure epilepsy. In C6te d'Ivoire sap of youngleafy twigs is added to palm wine for its aphro-disiac properties. In Kenya the pulped root issteeped in water and the liquid is drunk totreat cough.Rinoreo At66iensis Chipp, a shrub or smalltreeup to 7 in tall, is distributed from C6te d'Ivoireto Cameroon. The wood is used to make uten-SIIs and the twigs are used as toothbrushRinoreo oblongifolio (C. H. Wright) Marquandex Chipp is one of the taller species of the ge-nus, reaching 12 in tall and 30 cm in hole di-ameter. In DR Congo the wood is used in con-struction and for household utensils, and inSierra Leone for spoons and combs. In the Gen-tralAfrican Republic a root extract is drunk asa purgativeThe wood of Rinoreo seieensis De Wild. andRinoreo subsessilis M. Brandt, both found inthe Central African rainforest, is used for ar-row shafts and house construction, respective-Iy. The bark sap of the latter species is used todye hair.

Ecology Rinoreo brochypeto!o is found inthe undergrowth of evergreen forest and gal-Iery forest, but also in savanna. In East Africait is found up to 1900 in altitude.

Genetic resources and breeding Rinoreobrochypetolo is widespread and not intensiveIyexploited. Therefore, it does not seem to be indanger of genetic erosion

Prospects In view of the small size of thebole, wood of Rinoreo brochypetolo and otherRLnoreu spp. Is not of commercial interest, and

use will remain restricted.

Major references Aubr6ville, 1959; Burkill,2000; Grey-Wilson, 1986; Neuwinger, 2000;Taton, 1969

Other references Bakker, van Gemerden &AChoundong, 2006; Beentje, 1994; BOS, 1989;Bouquet & Debray, 1974; Dale & Greenway,1961; Kokwar0, 1993; Normand & Paquis,1976; Terashima & Ichikawa, 2003; Wahlert &Ballard Jr, 2009.

Authors C. H. Boseh

SACOGLOTTiS GABONENSis (Baill. ) Urb.

Protologue Mart. , F1. bras. 12(2): 449 (1877)Family HumiriaceaeVernacular names Bitterbark tree, cherry

inahogony (En). Bidou (Fr)Origin and geographic distribution SOCo-

glottis gobonensis occurs from Senegal andGambia east to the Central African Republicand south to Angola.

Uses The wood of SOCoglottis 8060nensis iswidely, though locally, used for hydraulic con-struction, house and bridge construction, car-pentry, heavy flooring, poles, ship building,vehicle bodies, railway sleepers and mineprops. It is suitable for interior trim, furniture,cabinet work, toys, novelties, turnery and pat-tern making. In Liberia and Nigeria, it is usedto make canoes and in Gabon for the ribs of

boats. The wood is a good firewood and produc-us a valuable charcoal

Infusions of the stem bark of SOCoglottis 8060-itensis are commonly taken to treat fever, diar-rhoea, gonorrhoea and abdominal pain, andsometimes they are used to treat hypertension

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SOCoglottis gobonensis - wild

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and diabetes. In coastal Cameroon the KOIa

pygmies and Mvae people use a decoction ofthe crushed bark mixed with leaves of Di-

o800reo minuti/1070 Engl. as a rectal enema totreat acute abdominal pain. In Congo a decoc-tion of the stem bark is used to cure difficultcases of dermatitis. In Sierra Leone a barkdecoction is used to treat stomach-ache and italso used as a spice in food to induce heat innursing and pregnant mothers. In coastal C6ted'TVoire the diluted stem sap is used inhipbaths to promote muscle tone in womenafter childbirth. In Gabon an extract of thestem bark is drunk as an emetic. In Senegaland Congo a stem bark decoction Is mixed withother plants and added to bath water to treatovarian troubles, vaginal infections and chil-dren with fever.

Stem bark is used as a palm wine additive, asit is claimed to prolong the shelf life of thewine, add potency, reduce foaming and imparta bitter taste. It is reported to have aphrodisiacproperties. The bark is used as a fish poison.The fruit is edible, sweet and tastes like hana-na bread. It is mainly eaten by children andcan be used to make an alcoholic drink. Incoastal Cameroon the seeds are grilled andeaten by pygmies. The sweetness of the resinstored in the cavities of the stone is much

sought by beesProduction and international trade Timber

of SOCoglottis gobonensis is known in the in-ternational trade as 'ozouga', but volumestraded are small. Stem bark is sold for medici-

nal purposes in local markets. Quantities trad-ed are unknown.

Properties The heartwood is brown to pur-PIish red and rather indistinctly demarcatedfrom the narrow, yellowish sapwood. The grainis straight or wavy, texture fine and even. Thewood is heavy, with a density of 870-950 kg/msat 12% moisture content. It should be air dried

slowly and with care to avoid serious checkingand warping. The rates of shrinkage are quitehigh, from green to oven dry 5.0-5.8% radialand 9.0-10.0% tangential. Once dry, the woodis moderately stable to unstable in serviceAt 12% moisture content, the modulus of rup-ture is 109-233 N/min2, modulus of elasticity13,330-21,770 N/mm2, compression parallel to

66-104 N/min2, shear 10-13 N/min2,grain

cleavage 22-23 N/min, Janka side hardness9065-11,005 N, Janka end hardness 12,495 Nand Chalais-Meudon side hardness 7.3-10.9.Considering its high density and hardness, thewood is not difficult to saw and work, but it has

a severe blunting effect on saw teeth and cut-ting edges; stellite-tipped saw teeth and tung-sten-carbide cutting edges are needed. Thewood is difficult to nailand screw, butthe hold-Ing power Is good; pre-boring is recommended.The finishing and gluing properties are satis-factory, but the wood is not suited for veneerand plywood production. It is durable, beingresistant to fungal and insect attacks. Theheartwood is resistant to impregnation withpreservatives.Bergenin, an ISOCoumarin, was identified asthe main active compound of the stem barkextract of SOCoglottis gobonensis. The stembark extract is reported to have hepatoprotec-tive properties. Both the extract and hergeninreduced the rate of formation of intermediatesof the lipid peroxidation pathway (lipid hy-droperoxide aldehydes, carbonyls) as well ascomplementing the primary anti-oxidant en-zymes catalase and superoxide dismutase dur-ing 2.4-DNPH-induced membrane lipid peroxi-dation in rat liver and red blood cells. In invo inrats, hergenin protects against 2.4-DNPH-induced hepatotoxicity and toxicity to red bloodcells. However, in another experiment, a stembark extract given orally to rats showed hepa-totoxicity even at low doses. Bergenin also pro-tects stored vegetable oils against peroxidativedeterioration over a period of time. A stembark extract added to drinking water of ratsincreased prothrombin and thrombin levels ofblood plasma.Further analyses of stem bark extract haveshown tannins in appreciable amounts with atrace of saponins. They have also revealed thepresence of 2 CIS/trans isomers of 11gnans (ca-lopiptine and galgravine) which may play arole in cancer preventionThe alcoholcontent of palm wine from Raphiohookeri G. Mann & H. Weridl. with stem barkadded became almost twice as high as that ofuntreated palm wine. It was found that theethanol tolerance and OSmotolerance of the

yeast SOCchoromyces cereuisioe were enhancedby the bark extract, whereas 110cculation andinvertase activity were reducedThe endocarp contains 54% oil of unknowncoinposltlon

Adulterations and substitutes Bergeninis commercially extracted from Ardisio andBergenio spp. (Siberian tea and marlberrybush) and is used as a weight-loss product, e. g.in body building, and in Chinese medicine toincrease body heat.

Description Evergreen, medium-sized to

SACOGLOTTls 563

564 TIMBERS2

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green to yellow; stone with 10 grooves andmany resinous cavities, I-3-seeded. Seeds ob-long, c. 15 min x 3 min. Seedling with epigealgermination; hypocoty1 5-9 cm long, epicotyl c.2 cm long; cotyledons leafy, ovate, c. 1.5 cm x0.5 cm; first 2 leaves opposite.

Other botanical information SOCoglottiscomprises 9 species, which all occur in SouthAmerica except SOCoglottis gobonensis, whichis considered to be closely related to SOCoglottisQinozonico Mart. from South America and theCaribbean. The fruits of SOCog!ottis dingzonicuare eaten in Venezuela to cure diarrhoea. Thetimber is valued for heavy construction in Ven-ezuela and Brazil.

SOCog!ottis gobonensis may have evolved fromseeds, which arrived on the West African coastfrom South America, being one of the mostIllustrative cases of dispersal by water fromSouth America to Africa. The resin-filledchambers of the stone make it float in waterand seeds remain viable for up to 4 years.

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);14: scalariform perforation plates; 16: scalari-form perforation plates with 10-20 bars; 22:intervessel pits alternate; (23: shape of alter-nate pits polygonal); 25: intervessel pits small(4-7 prn); 26: intervessel pits medium (7-10Urn); 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; 47: 5-20 vessels per square minimetre; 56: tylosescommon. Tracheids and fibres: (60: vascu-Iar/vasicentric tracheids present); 62: fibreswith distinctly bordered pits; 63: fibre pitscommon in both radial and tangential walls;66: non-septate fibres present; 70: fibres verythick-walled. Axial parenchyma: 77: axial pa-Tenchyma diffuse-in-aggregates; 78: axial pa-renchyma scanty paratrachea1; (84: axial pa-renchyma unilateral paratrachea1); (92: four(3-4) cells per parenchyma strand); 93: eight(5-8) cells per parenchyma strand; (94: overeight cells per parenchyma strand). Rays: 97:ray width I-3 cells; (100: rays with multiseri-ate portion(s) as wide as uniseriate portions);(107: body ray cells procumbent with mostly 2-4 rows of upright and/or square marginal cells);108: body ray cells procumbent with over 4rows of upright and/or square marginal cells;(109: rays with procumbent, square and up-

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SOCoglottis gobonensis - I, flowering bronch, . 2,frui"Redrown and udopted by AChmod Sotiri Nur-hornon

fairly large tree up to 40 in tall; hole branchlessfor up to 20 in but often branching low, oftencrooked, knotty and deeply fluted, up to 180(-450) cm in diameter, with buttresses up to 2.5in high; bark surface in young trees fairlysmooth with horizontal lenticels, in old treesbrown to dark brown, very scaly, inner barkfibrous, brittle, pinkish brown to red-brown,exuding a clear, sticky sap; crown very large,wide-spreading, heavily branched, rounded andfairly open. Leaves alternate, simple; stipulesc. I mm long, early falllrig; petiole 0.5-I cmlong; blade ovate to elliptical or oblong, 6-15cm x 2.5-6 cm, base curieate, apex acuminate,margin obscurely toothed, thin-leathery, gla-brous, pinnately veined with 6-12 pairs of Iat-eral veins. Inflorescence a short axillary cymeup to 5 cm long. Flowers bisexual, regular, 5-merous; pedice1 0.5-1.5 min long, jointed atbase; sepals c. 1.5 mm long, hairy outside; pet-als free, linear, c. 7 min long, hairy outside,white; stamens 10, fused at base, 5 longer sta-mens c. 5 min long, 5 shorter ones c. 3.5 minlong; ovary superior, ovoid, 5-celled, glabrous,style grooved, c. 4 min long. Fruit an ellipsoidto globose drupe 3-4 cm x 2.5-3.5 cm, smooth,

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right cells mixed throughout the ray); 1/5: 4-12 rays per min. Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells(L. N. Banak, H. Beeckman & P. E. Gasson)

Growth and development In Liberia flow-ering of SOCog!ottis gobonensis occurs fromDecember-March. The fruits are adapted fordispersal by elephants. They develop slowlyover the dry season, ripening in September-December in Liberia and remaining dull green.They fallto the ground when ripe and developa strong yeasty smell. Chimpanzees and man-drills also eat the fruits. Like LOphtro glutoBanks ex P. Gaertn. and Aucoumeo hloineono

Pierre, SOCoglottis gobonensis is able to estab-lish in the edges of moist savanna and it Iscommon in younger forest types. It can doinI-nate this habitat, since its branches form hugearches, suppressing vegetation under them,hence cooling annual savanna fires, whichmight otherwise scorch their trunk. Early colo-nizers have low, round canopies and branchlow down, but later generations grow taller andbranch higher to escape from the shadowYoung trees close to the savanna edge oftenhave multiple trunks because they have cop-piced after being broken by elephants whichfeed on theirleaves and bark

Ecology SOCoglottis gobonertsis occurs inevergreen forest, on river banks, along lagoons,on forest and savanna edges and in secondaryforest. It is one of the dominant trees in thecoastal rainforests of West and Central Africaand is particularly abundant in Garnero0n (as-SOCiated with LOphtr0 o10to) and in Gabon (as-SOCiated with Aucoumeo hintneono). In C6ted'TVoire it is characteristic of the most humid

evergreen forests. It is usually found in smallpockets, but occasionally in quite large andalmost pure stands.

Propagation and planting Natural propa-gation of SOCoglottis gobonensis is only by seedNatural regeneration is not very abundant asmany seeds are damaged by Insects. Theweight of 100 stones is 560-1000 g. Seeds take2-4 months to germinate. The germinationrate Is quite low.

Management SOCog!ottis gobonensis is lessdesirable for commercial forestry as the largecrown suppresses regeneration of more valua-ble timber species. After cutting the tree, thestump coppices vigorously.

Harvesting The minimum bole diameter forharvesting is 70 cm in Liberia and Gabon. Thestern bark and wood of SOCoglottis gobonensis

is harvested from the wild.

Handling after harvest Logs do not floatin water and cannot be transported by riverThe bark of SOCoglottis gobonensis is generallysold in markets in sheets or rolls for use as an

additive to palm wine or as a medicine, lessfrequently as a powder for emetic use.

Genetic resources SOCoglottis gobonensisis widespread and locally common, and there-fore notthreatened by genetic erosion

Prospects SOCoglottis gobo, ,errsis plays anImportant role in the succession in and regen-eration of natural forests. As a widespread andoften common species It is important in thelocaleconomy as a timber and for firewood andcharcoal production. Because of the often poorshape of the hole, commercial exploitation ofthe timber for international trade is limited.

Selection for bole shape and research intoproper management practices could lead tobetter shaped holes with higher commercialvalue in the timber trade

The stem bark extracts of SOCoglottis gobonen-sis and its main isolated compound hergeninhave interesting hepatoprotective and anti-oxidant properties, but more research is need-ed to evaluate its potential as a lead drug. Itsuse to increase alcohol content in palm winedeserves more research attention, as does in-formation concerning health risks

Major references Badr6, 1972c; BurkiU, 1994;ITvine, 1961; Maduka, Okoye & Mahmood,2004; SIepel, Poorter & Hawthorne, 2004;Takahashi, 1978; Voorhoeve, 1965; White &Abernethy, 1997.

Other references Cuatrecasas, 1961; Ekouya& Itoua, 2005; Ezeronye, Elijah & Oninelukwe,2005; Gassita at al. (Editors), 1982; Haw-thorne, 1995; Hoshin0, 1985; InsideWood, un-dated; Liben, 1970a; Maduka, 2005; Maduka,Okoye & Eje, 2002; Maduka, Uhwache & Oko-ye, 2003; Madusolumuo & Okoye, 1993; 0jime-Iukwe, 2001; Okoye & Ohaeri, 1995; Raponda-Walker & Sillans, 1961; Renner, 2004; Udosen& OiOng, 1998

Sources of illustration Voorhoeve, 1965Authors E. Dounias

SARCOCEPHALUS 565

SARCOCEPHALUS FOBEGUINllPob6g.

Protologue Essaifi. Gum6efranc. : 313 (1906)Family RubiaceaeChromosome number2n=44

Synonyms Nouc!eopobeguinii(Pob6g. ) Merr(1915)

566 TIMBERS2

Vernacular names Water kusia (En). Bi-linga des marais, hots d'or (Fr)

Origin and geographic distribution Sureo-copholuspobeguiniiis distributed from Senegalto Zambia. It is occasionally planted.

Uses The wood is used for construction, join-ery and cabinet work. Although the wood hasbeen recorded to be suitable for making inor-tars, like that of the related NOMc!eo di-derrichii(De Wild. & T. Durand) Merr. , othersources indicate it is not used for making inor-tars in Ghana because it imparts a bitter tasteto food. The pulp of the infructescence is edible,but not much eaten.

Soreocepholus pobeguinii is widely used inAfrican traditional medicine. A root decoctionis taken as an arithelminthic in Guinea. In

Nigeria root preparations are used againstgonorrhoea. The powdered bark is taken inwater against Intestinal pain and diarrhoea inSenegal, and a bark decoction as an oxytocicIn Guinea a bark maceration is recorded to be

taken as an abortifacient, while in Garnero0nthe bark is said to be used against threatenedabortion. In C6te d'Ivoire bark preparations areused for the treatment offever andjaundice. InNigeria a bark decoction is drunk for thetreatment of stomach-ache. In Gabon a macer-

ation or decoction of the bark is drunk for the

treatment of gonorrhoea. In DR Congo barkdecoctions are taken for the treatment of ina-

Iana, a decoction of the barks of Soreocepholuspobegutnii and Symphonio globulifero L. f. istaken against sexual asthenia, and a barkpreparation Is applied as a suppository againstepilepsy. A leaf decoction is used against jaun-dice in Nigeria. In Gabon a leafinfusion is tak-en againstfever

Production and international trade The

wood is only used locally and not commercial-ized

Properties The heartwood is yellow or paleorange, the sapwood pale yenow to orange. Thegrain is irregular, texture fine. The wood ismedium-weight, with a density of 500-780kg/ina, and moderately hard to hard. It worksand glues well, and holds nails and screwswell

Aqueous, ethanolic and dichloromethane ex-tracts of the bark have shown in-vitro antima-

Ianal activity against Plusmodium foieiporumand in-vivo antimalarial activity against PIOs-inodium berghei and Pigsmodium yoellii inmice. No significant in-vivo toxic effects in micewere observed for aqueous and ethanolic barkextracts, but dichloromethane extracts haveshown high in-vitro cytotoxic activity. A by-dromethanolic extract of the bark showed SIg-nificant chromosome-damaging activity, buthad no cytotoxic activity. A clinical test with asmall group of human adult volunteers diag-nosed with uricomplicated PIOsmodium foiei-parum malaria has shown that oral admin-1stration during meals of capsules of an 80%ethanolic extract of the stem bark was effective

against malaria and well-tolerated, giving onlymild and self-resolving adverse effects. inka-lotds, especially the major compound strictos-amide, are assumed to be responsible for theantimalarial activity. However, neither stric-tosamide nor other alkaloids did show in-vitro

antimalarial activity, suggesting activation ofstrictosamide, probably by the gastrointestinaltract, Is necessary for its activity. The root barkcontains alkaloids and quinovic acid glycosidesAn aqueous extract of the leaves has shown in-vivo anti-oxidant activity in rats

Botany Deciduous, small to medium-sizedtree up to 30 in tall, sometimes a shrub; holebranchless for up to 15 in, straight and cylin-drical, up to 100 cm in diameter, without but-tresses; bark surface longitudinalIy fissured,flaking in papery scales c. 2 cm in diameter,whitish, grey or pale brown, inner bark fibrous,pale yellow-brown, turning darker on exposure;crown with whorled horizontal branches; twigsshort-hairy when young. Leaves opposite, SIm-PIe and entire; stipules triangular, 3-7 mmlong; petiole slender, 1.5-5 cm long; blade ob-long-elliptical to broadly elliptical, 8-25 cm x5.5-16 cm, curieate at base, rounded to acute oracuminate at apex, thin, glabrous, pinnatelyveined with 7-9 pairs of lateral veins. Inflo-rescence a solitary, terminal, rounded head 2-5

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Soreocepho!uspobeguinii- wild

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cm in diameter; peduncle I-3 cm long. Flowersbisexual, regular, 4-5-meTous, fragrant; calyxtubes fused to each other, lobes oblong-spatulate, 2-4 min long, densely hairy; corolla8-10 min long, pinkish white, white or cream-coloured, tube funnel-shaped, c. 7 min long,hairy inside, lobes oblong to ovate, 1.5-2 mmlong; stamens inserted in the throat of the co-rolla tube, alternating with the lobes, withoutfilaments; ovary inferior, 2-celled, style c. 12min long, with spindle-shaped thickening nearapex. Fruits fused into a globose, fleshy in-fructescence 3.5-10 cm in diameter, yellow,densely pitted because of the persistent calyxlobes, with yellow pulp and an apple-like smell,many-seeded. Seeds ellipsoid or ovoid, I-3 minlong, reddish, Teticulate. Seedling with epigealgerminationSoreocepholus comprises only 2 species, re-stricted to tropical Africa, and seems related toBurttdouyo andNouc!eoIn West Africa Soreocepholus pobeguiniiflow-ers in (April-)MayJuly, and fruits in Septem-her-November(-December). The fruits are afavoured food for chimpanzees in Guinea andgorillas in Central Africa. The seeds are rec-orded to be dispersed by water.

Ecology Soreocepholus pobeguinii occursfrom sea-level up to 1400 in altitude on riverbanks and lake sides, and in swamp forest,where it may grow in nearly pure stands

Management Seed germination takes I-3weeks, and the germination rate is usuallyhigh. In an inventory in forest in southernCameroon, an average wood volume of 0.25ms/ha has been recorded for trees with a hole

diameter of more than 15 cm.

Genetic resources and breeding As 807-cocepho!uspobeguiniiis widely distributed andnot much used, it seems not threatened withgenetic erosion

Prospects The wood of Soreocepho!us pobe-gumiiis only used locally and not traded inter-nationally, and there are no indications thatthis situation winchange. Quantitative data onthe wood properties are scarce, and infor-mation on the durability of the wood is lackingaltogether. Research on the pharmacologicalproperties, especially in relation to the treat-merit of malaria, showed promising results forpossible drug development in the pharmaceuti-calmdustry

Major references Bridson & Verdcourt,2003; Burki11, 1997; Mesia at a1. , 2010; Taylor,1960; Vivien & Faure, 1985

Other references de in Mensbruge, 1966;

Hall6, 1966; Hawthorne & Jongkind, 2006; Ka-dirt, Adegor & Asagba, 2007; Liu at a1. , 2011;Mesia at a1. , 2005; MeSIa at a1. , 2011; Vivien &Faure, 1996; Voorhoeve, 1979; Zeches at al. ,1985

Authors M. Brink

SCHEFFLERAABYSSINICA (Hochst. exA. Rich. )Harms

SCHEFFLERA 567

Protologue Engl. & Prantl, Nat. Pftanzenfam.111, a 38 (1894).

Family AraliaceaeOrigin and geographic distribution Schef-

Iler0 o6yssinico occurs in mountain regionsfrom Nigeria and Cameroon east to Ethiopiaand Kenya, and south to Malawiand Zambia

Uses The wood is used for furniture, boxesand agricultural implements. It is suitable forinner layers of plywood. Occasionally It is alsoused as firewood. The flowers are a source of

nectar and pollen for honey bees. Schemeroo6yssinico is planted as life fence

Properties The heartwood is yellowishbrown and indistinctly demarcated from thesapwood. The texture is coarse. The wood islightweight, with a density of about 460 kg/in3at 12% moisture content. It is soft and easy towork. Some triterpene saponins have been iso-lated from the leaves

Botany Smallto medium-sized tree up to 30in tall, sometimes an epiphyte; hole oftentwisted; bark surface smooth to fissured, corky,grey-brown to grey-black, inner bark whitish;crown large, spreading. Leaves arranged spi-rally, digitately compound with 5-8 leaflets;stipules triangular, I-1.5 cm long; petiole up to

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Sche//Zero o6yssinicu- wild

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568 TIMBERS2

42 cm long, slightly grooved; petiolules up to12.5 cm long; leaflets elliptical to ovate, 6-25(-40) cm x 3-15(-20) cm, rounded to cordate atbase, acuminate at apex, margins finelytoothed, papery to leathery, glabrous, pinnatelyveined with 9-14 pairs of lateral veins. Inno-rescence a false raceme up to 40 cm long, oftenmany together at ends of branches, consistingof stalked umbels; bracts at base of innores-cences broadly ovate, up to 1.5 cm long. Flow-ers bisexual, regular, 5-8-merous, creamy togreenish yellow, fragrant; pedice1 2-11 mmlong; calyx wavy; petals fused, 1.5-2 min long,falling as a smallcap; stamens the same num-her as petals, free, 2-3 mm long; ovary inferior,5-8-celled, styles 5-8, 0.5-I mm long. Fruit anearly globose drupe-like berry c. 5 min in di-ameter, ribbed, reddish when ripe, glabrous orslightly hairy, several-seeded. Seeds ellipsoid,compressed, smooth.In Ethiopia trees flower from February toMarch, and fruits mature 2-3 months later.The flowers are pollinated by insects includingbees. The fruits are eaten by birds, which mayserve as seed dispersersSchemera probably comprises about 700 spe-cies and is widely distributed in the tropics andsubtropics. In mainland Africa about 12 speciesoccur, and in Madagascar about 15Scheftlero umbellifero (Sond. ) Baill. is a smallto medium-sized tree up to 20 in tall, usuallywith a straight hole up to 60 cm in diameter,occurring in Malawi, Zimbabwe, Mozambiqueand South Africa. The whitish, lightweight andsoft wood is used for boxes and matches. A

bark extract is taken to treat malaria, leaf ex-tracts are drunk against rheumatism, colic andinsanity, and root infusions against Inflamma-tions. The use of the bark in traditional medi-

cine to treat malaria has been rationalized in

experiments with bark extracts, which showedin-vitro activity against chloroquine-resistantstrains of PIOsmodium foieiporum. Schemeroumbellifero is planted as an ornamental tree ingardens

Ecology Schemer0 o6yssinicomountain rainforest at (1200-)1400-2800 inaltitude, often in secondary forest. In Ethiopiait is locally one of the dominant forest trees,e. g. with Hogeni0 o6yssinico (Bruce) J. F. Ginelat 2400-2800 in altitude, whereas in mountainforest in southern Sudan it is common.

Management Schemer0 o6yssinico is prop-agated by seed, cuttings and wildlings. Seedsdo not require any pre-treatment; they startgerminating about 2 weeks after sowing. Cut-

tings should be planted at the end of the rainyseason

Genetic resources and breeding Schef-nerd abyssinico is widespread in mountainousregions of Africa, and is not threatened. How-ever, isolated populations, like those in Nigeriaand Garnero0n, may become threatened withongoing forest clearing.

Prospects Scheffler0 o6yssinico will proba-bly remain of some importance for local appli-cations, but does not seem to have prospects forcommercial exploitation, with its often poorlyshaped hole and low-quality timber

Major references Barnps, 1974; Bamps,1989; Bekele-Tesemma, 2007; Jacques-Fellx,1970; Tennant, 1968

Other references Beentje, 1994; Cannon,1978b; Coates Palgrave, 1983; Keay, 1989;Mbambezeli, 2005; Neuwinger, 2000; Pillay,Maharaj & Smith, 2008; Sommerlatte & Som-merlatte, 1990; Tapondjou at a1. , 2006; Tetyanaat a1. , 2002.

Authors R. H. M. J. Lemmens

SCHOTIABRACHYPETAIASond

Protologue Linnaea 23: 39 (1850).Family Caesalpiniaceae (I. ,eguminosae - Cae-

salpinioideae)Chromosome number2n=24

Vernacular names Weeping schotia, weep-ing boerbean, fuchsia tree, African walnut(En)

Origin and geographic distribution Schotiobrachypetolo occurs in eastern Botswana, Zim-babwe, southern Mozambique, eastern SouthAfrica and Swaziland

Occurs In

I.

Schotio brochypetolo- wild

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.

Uses The wood is used for furniture, floor-ing, wagon beams and carving. Roasted seedsare eaten. The flowers, which produce abun-dantly nectar, are sometimes chewed. Prepara-tions of the bark, and sometimes of roots andleaves, are used in traditional medicine. Barkdecoctions and infusions are taken as emetic,and to treat heartburn, hangovers, nervousconditions, mental health problems and fonta-nel depression in babies, and are applied toswellings. The bark and roots are used to treatdiarrhoea, and bark is applied in mixtures as atonic. Pulverized leaves are used as a dressingto treat ulcers, and the smoke of burntleaves isinhaled against nose bleeding. The bark hasbeen used for tanning and dyeing reddishbrown. Schoti0 670chypetolo is planted as or-namentaltree in gardens and parks because ofits glossy foliage and bright red flowers.

Properties The heartwood is dark brown tonearly blackish, often with a greenish tinge,and is distinctly demarcated from the pinkishsapwood, which changes to yellowish brownupon exposure. The texture is fine. The wood isheavy and hard. It is only moderately durable,but quite resistant to termites. It has beenreported that wood dust may cause allergicreactions to the eyesThe medicinal activity of the bark may be dueto the presence of tannins. Fatty acids withantibacterial activity against Boci!Jus subtilisand Stophylococcus oureus, and to a lesserextent against ESChertchio coli and K!ebsie!IQ

have been isolated from thepiteumon, oe,

leaves. Bark extracts exhibited pronouncedmonoamine oxidase-B inhibition activity, sup-porting the use in traditional medicine to treatmental health problems. Polyhydroxystilbeneshave been isolated from the heartwood; themajor compound is 3.3', 4.5,5'-pentahydroxystil-herie.

Botany Evergreen or briefly deciduous,smalltree up to 15 in tall; hole up to 60 cm indiameter; bark surface smooth with hoopmarks and reddish brown in young trees, be-coining rough and greyish brown to reddishbrown in older trees; crown rounded, withspreading branches; twigs short-hairy whenyoung. Leaves alternate, panpinnately coin-pound with (3-)4-6(-7) pairs of leaflets; stip-ules narrowly triangular, c. 5 mm long, cadu-cous; petiole I-2 cm long, rachis 3.5-17.5 cmlong, grooved; leaflets opposite, elliptical torhombic-ovate, 2-6.5 cm x I-3 cm, curieate torounded and often asymmetrical at base,rounded to slightly notched at apex, margin

entire to slightly wavy, sparsely hairy or gla-brous, pinnately veined. Inflorescence a con-gested panicle up to 13 cm long, usually onolder branches. Flowers bisexual, zygomorphic,deep red; pedice1 3-5 mm long, expanding intothe 3-5 min long hypantium; calyx lobes 4,broadly ovate, c. I cm long, unequal, gland-dotted; petals often strap-shaped and up to 3mm long or absent, but sometimes I-4 narrow-Iy obovate petals up to 2 cm long present; sta-mens 10, fused at base, up to 2 cm long; ovarysuperior, narrowly elliptical, flattened, c. I cmlong, with stipe c. 3 mm long fused to the by-pantium, style c. I cm long. Fruit an oblong,flattened pod 6-17 cm x 3.5-5 cm, thickenedalong upper suture, glabrous, tardily dehiscent,several-seeded. Seeds broadly elliptical torhombic, flattened, 1.5-2 cm x I-1.5 cm,smooth, pale brown, with cup-shaped, yellowanI at base

Schotio brochypetolo usually grows slowly, butwhen planted on deep sandy soils it may growquite rapidly; a planted tree reached 12 in tall17 years after being planted as a cutting. Thetree is usually evergreen, but in colder areas ofSouth Africa it is deciduous for a short periodYoung foliage is pink to red, turning graduallyto coppery and light green, and is often accom-panied by the deep red flowers. Trees usuallyflower in September-October. Nectar is oftenso abundant that it drips from the flowers,which are visited by sunbirds, but also by in-sects such as bees, and by monkeys. Fruitsripen about 6 months after flowering. Theseeds are probably mainly dispersed by birdsattracted by the yellow anIs.Schotio comprises about 4 species and is con-fined to southern Africa. Its affinity is still un-certain. Schotio ofro (L. ) Thunb. is a shrub orsmall tree up to 7 in tall found in southernNamibia and South Africa. The reddish brown

wood is used for yokes and felloes, and as fire-wood;it is hard and durable. Roasted seeds areeaten. The foliage is eaten by livestock. Schotioo170 is an attractive ornamental. The bark hasbeen used for tanning.Schotio capitato Bone is a shrub or smalltreeup to 7 in tall found in Zimbabwe, southernMozambique, eastern South Africa and Swazi-land. The wood is sometimes used for furni-ture. Schotio capitato is occasionally plantedas ornamental

Schotio lotifo!to Jacq. is a smalltree up to IOC15) in tallendemic to South Africa, but recentlyit has also been recorded from southern

Mozambique. The tough, whitish wood is used

SCHOTiA 569

570 TIMBERS2

for fence posts. Roasted seeds are eaten. Barkdecoctions are taken to treat stomach-ache.

The bark of Schotio lotifolio is commonly usedin South Africa to treat tick-borne diseases of

livestock. The bark has been used as a dye pro-ducing a greenish colour, and for tanning.Schotio lotifolio is occasionally planted as anornamental

Ecology Schotio brachypeto!owoodland and thickets up to 1350 in altitude. Itis often found on termite mounds and on river

banks. The tree is drought resistant and slight-Iy frost tolerant

Management Propagation can be donebyusingseeds or truncheon cuttings. Young trees per-form best when they have been planted in deepsandy soils with plenty of well-decomposedcompost added

Genetic resources and breeding Thereare no indications that Schoti0 670chypetolo isin risk of genetic erosion. However, the bark islocally popular and much collected, and thismay have serious impact on local populationsof the species

Prospects The holes of Schoti0 670chypetoloand other Schotio spp. are usually too smalland too poorly shaped to be of commercial in-terest for timber. However, Schotio sppvalued as multi-purpose trees, especially asornamental and for medicinal purposes. Someof the known pharmacological activities de-serve more research attention, e. g. the mono-amine oxidase-B inhibition activity, whichcould be of interest for the development of newmedicines against Parkinson's and Alzheimer'sdisease. Schotio trees are also important as asource offo0d for wildlife.

Major references Brummitt at a1. , 2007a;Coates Palgrave, 1983; Mbambezeli & Notten,2001; Palmer & Pitman, 1972-1974; van Wyk,van Oudtshoorn & Gencke, 1997.

Other references Drewes & F1etcher, 1974;Grace at a1. , 2002a; Huxham at a1. , 1998;Masika, Sultana & ato1ayan, 2004; Mathabe ata1. , 2006; MeGaw, Jager & van Staden, 2002a;MCGaw, Jager & van Staden, 2002b; Neuwing-er, 2000; Stafford at a1. , 2007; van Wyk &Gencke, 2000.

Authors R. H. M. J. Lemmens

Chromosome number2n = 44, 46Origin and geographic distribution Schre-

bero gluto is widespread, from Eritrea andEthiopia southwards through eastern DR Con-go and East Africa to Angola, northern SouthAfrica and Swaziland.

Uses In Kenya the wood is mainly used forconstruction in house building, poles, furnitureand tool handles. It is suitable for flooring,joinery, Interior trim, mine props, vehicle bod-ies, sporting goods, toys, novelties, agriculturalImplements, turnery, pattern making, veneerand plywood. The wood is excellent as firewood,and is also used for charcoal productionSeveral plant parts are used in traditionalmedicine. Pounded roots, twigs and leaves arechewed to treat tooth complaints and as anti-tussive, and in water they are applied as awash to ulcers. Bark is also chewed to treat

toothache, and bark decoctions are applied asanodyne. Leaves are chewed to treat tonsillitis,pharyngitis and headache, and leaf decoctionsare administered as a vapour bath for treat-merit of headache and taken to treat colds,cough, fever, and as emetic, oxytocic and tonic.Schreberu o10to ls occasionally planted as or-namentaltree

Properties The heartwood is whitish to palebrown, often with slightly darker streaks, notdistinctly demarcated from the sapwood. Thegrain is straight to slightly wavy, texture fineand even. The wood is moderately heavy, witha density of 780-835 kg/ms at 12% moisturecontent, hard and strong. It should be driedslowly and carefully to avoid serious splitting,checking and distortion. Once dry, it is moder-ately stable in service.The wood saws and works fairly easily with

Occurs In

are

SCHREBERAAl, ATA (Hochst. ) WeIw.

Protologue Trans. Linn. SOC. London 27: 39(1869).

Family Oleaceae

I.

Schreber0 o10to - wild

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both hand and machine tools. It planes to asmooth surface, and polishes well. Pre-boring isneeded for nailing. The wood glues, paints andvarnishes satisfactorily, and it turns fairlywell. The heartwood is moderately durable.The sapwood is susceptible to Lyctus attack.The heartwood is resistant to preservatives.Tests in rats showed that bark extracts have

analgesic effects, confirming the use in tradi-tional medicine, but they also demonstratedsome toxicity, causing hepatic injuries.

Botany Deciduous, small to medium-sizedtree up to 20(-25) in tall; hole usually shortand crooked, up to 60 cm in diameter, oftenfluted; bark surface smooth or longitudinalIyfissured, sometimes flaking with thin scales,pale grey to yellowish brown, inner barkcream-coloured with orange markings, darken-Ing upon exposure; crown fairly open; twlgsinitially hairy but becoming glabrous, withlenticels. Leaves opposite, imparipinnatelycompound with (3-)5(-7) leaflets; stipules ab-sent; petiole I-5 cm long, slightly winged; ra-chis 2-5 cm long, winged; leaflets opposite,nearly sessile, elliptical to obovate, 2-14 cm xI-6 cm, curieate and often asymmetrical atbase, apex rounded to short-acuminate or occa-sionally notched, glabrous or sparsely short-hairy, pinnately veined with up to 10 pairs oflateral veins. Inflorescence a terminalcyme upto 10 cm long, glabrous to hairy, few-floweredto many-flowered. Flowers bisexual, regular,5-7-merous, heterostylous, sweet scented; ped-icelup to 3(-5) min long; calyx campanulate, 2-4(-7) mm long, short-hairy; corolla white,sometimes flushed with pink, with reddishbrown hairs at the base of lobes, tube funnel-shaped, I-1.5 cm long, lobes 3-7 min long,spreading; stamens 2, inserted in the upperpart of the corolla tube, with short filaments;ovary superior, c. 1.5 min in diameter, 2-celled,style slender, short or long. Fruit a pear-shaped or obovoid woody capsule 2-4.5 cm x I-1.5 cm, pale brown, dehiscing with 2 valves,with up to 8 seeds. Seeds with a large wing, upto 2.5 cm x ICm.

Young Schreber0 o10to trees grow fairly rapid-Iy when they are planted in fertile soil. InUganda young trees showed an annual growthof up to I in/year at 1500 altitude and on deepand moist soils, but growth was much less athigher altitudes and on drier sites. In southernAfrica trees flower in December to February,and fruits ripen about 4 months after flower-ing. Fruits usually open while still attached tothe tree. The seeds with their wings are dis-

persed by wind; they spin during fallingSchrebero comprises about 8 species, of which5 occur in tropical Africa including Madagas-car, 2 in tropical Asia and I in South AmericaIt seems most closely related to Coinoronthusfrom Comoros and Madagascar, which also hasa woody capsuleSchrebero trichoclodo WeIw. is a shrub or

smalltree up to 10 in tallwith hole up to 30 cmin diameter, occurring from DR Congo andTanzania southwards to Angola, Zimbabwe,Mozambique and Madagascar. Its wood is 10-cally used for implements such as spoons. Barkand leaves are chewed as anodyne, leaf macer-ations are applied to the eyes to treat ophthal-inia, and root infusions are used as an eye 10-tion.

Ecology Schrebero dinto occurs in evergreenforest, riverme forest, open woodland andsometimes savanna with scattered trees, up to2500 in altitude. In East Africa it is often asso-ciated with Juniperus, Podoco, pus and 01eospp. , and occurs particularly in forest marginsand clearings.

Management Schreber0 o10to can be propa-gated by seeds and wildlings. Fruits should beharvested from the tree just before they open.They can be dried in the sun untilthey openand the seed can be collected. It is recommend-ed to sow seeds with wings pointing upwards;they can be sown without pre-treatment. Theseeds can be stored in a dry and coollocality forlonger periods. Trees can be managed by cop-picing and pruning, and intercropped with cof-fee.

Genetic resources and breeding Schrebe-70 o10to is quite widespread in various habitatsand not selectively harvested on a large scale.Therefore, there is no reason to consider It 11a-ble to genetic erosion

Prospects The wood of Schrebero gluto willremain of some local importance, but it doesnot seem to have prospects for trade on theinternational market because the tree hole is

usually too short and crooked or fluted.Schreber0 o10to has been recommended as afirewood crop for highland farmers in EastAfrica. It deserves more attention as an orna-

mental tree

Major references Bolza & Keating, 1972;Chikamai at al. , undated; Katende, Birnie &Tengnas, 1995; Maundu & Tengnas (Editors),2005; SEPASA1, , 2011.

Other references Beentje, 1994; CoatesPalgrave, 2002; Green, 2003; Kokwar0, 1993;Kupicha, 1983b; Liben, 1973; Maundu at al. ,

SCHREBERA 571

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2001; Mbaya, 1976; Neuwinger, 2000; Palmer& Pitman, 1972-1974

Authors R. H. M. J. Lemmens

SCHREBERAARBOREAA. Chev.

Protologue Mein. SOC. Bot. France a 180(1912).

Family OleaceaeVernacular names Pau goiaba (Po)Origin and geographic distribution Schre-

bero orboreo Is quite widespread, from Senegaland Guinea Bissau eastwards to southern Su-dan and Uganda, and south to DR Congo.

Uses The wood is used for general construc-tion, railway sleepers, boat-keels, utensils, toolhandles, shafts and beehives. It is suitable forflooring, joinery, interior trim, mine props,vehicle bodies, furniture, sporting goods, toys,novelties, agricultural Implements, turnery,pattern making, veneer and plywood. It is alsoused as firewood and for charcoal productionIn traditional medicine, fresh twigs are an in-gredient of a preparation that is taken to treatskin diseases. Macerations of the twigs areadministered as purgative to treat abdominalpain. In Nigeria the seeds are applied to theheads of children to treat skin complaints. InUganda several plant parts are used as purga-tive

Production and international trade The

wood of Schreber0 o760reo is only used locallyand not traded on the international timbermarket

Properties The heartwood is pinkish toyellowish brown, sometimes with violet

streaks, and not distinctly demarcated from

the paler sapwood. The grain is straight towavy, texture usually fine. The wood is moder-ately heavy, with a density of 750-830(-930)kg/in' at 12% moisture content, tough andquite hard. It air dries well, but care should betaken to avoid surface checking. The rates ofshrinkage are rather high, from green to ovendry 4.7-5.6% radial and 7.8-8.6(-9.5)% tangen-tial. Logs may either be back-sawn or quarter-sawn after felling. Once dry, the wood is un-stable in service. At 12% moisture content, themodulus of rupture is (77-)108-132 N/min, ,modulus of elasticity 11,800 N/mm2, coinpres-sion parallel to grain 44-63 N/mm2 shear 12-14 N/min2, cleavage 14.5-23.5 N/mm andChalais-Meudon side hardness 2.7-3.6The wood is fairly easy to saw and work. Inplaning, picking-up of grain can be problemat-IC, but it can be polished to a smooth finish. Fornailing, pre-boring is recommended to avoidsplitting. The gluing properties are satisfacto-ry. The wood is easy to wax, paint and varnish.The heartwood is fairly durable, but occasional-Iy susceptible to wood-boring beetles. The sap-wood is liable to Lyetus attack. The heartwoodis moderately resistant to preservatives.

Description Deciduous, medium-sized treeup to 30(-40) in tall; hole branchless for up to20(-30) in, straight and cylindrical, up to 80 cmin diameter, with small buttresses or fluted atbase; bark surface smooth but flaking with thinscales, yellowish white to greyish brown withpaler patches, inner bark fibrous, inottledcream and orange, darkening upon exposure;crown rounded; twigs initially hairy but becoin-ing glabrous, with lenticels. Leaves opposite,simple and entire; stipules absent; petiole I-5cm long; blade elliptical to ovate, 5-15 cm x 3-10 cm, curieate to rounded at base, acute toshort-acuminate at apex, thinly leathery, gla-brous or nearly so, pinnately veined with 6-12pairs of lateral veins. Inflorescence a terminalor axillary cyme up to 10 cm long, few-flowered. Flowers bisexual, regular, (4-)5-8-meTous, heterostylous, sweet-scented; pedicel3-7 mm long; calyx campanulate, 3-5 mm long,slightly hairy or glabrous; corolla white togreenish white with purplish brown hairs atthe base of lobes, tube funnel-shaped, I-2 cmlong, lobes c. 6 min long, spreading; stamens 2,Inserted in the upper part of the corolla tube,with short filaments; ovary superior, c. I minin diameter, 2-celled, style slender, short orlong. Fruit a pear-shaped woody capsule up to6.5 cm x 3.5 cm, purplish brown with palerlenticels, dehiscing with 2 valves, usually with

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ders; similar to Intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumina 50-100 prn;48: 20-40 vessels per square minimetre. Tra-cheids and fibres: 61: fibres with simple to inI-nutely bordered pits; (62: fibres with distinctlybordered pits); 65: septate fibres present; 66:non-septate fibres present; 69: fibres thin- tothick-walled; 70: fibres very thick-walled. Axialparenchyma: 75: axial parenchyma absent orextremely rare; 78: axial parenchyma scantyparatrachea1; (92: four (3-4) cells per paren-chyma strand); 93: eight (5-8) cells per paren-chyma strand. Rays: 97: ray width I-3 cells;(100: rays with multiseriate portion(s) as wideas uniseriate portions); 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 mm. Miner-alInclusions: 136: prismatic crystals present;137: prismatic crystals in upright and/orsquare ray cells; (138: prismatic crystals inprocumbent ray cells); 151: styloids and/orelongated crystals(FD. Kamala, P. Baas & H. Beeckman)

Growth and development In Senegal thegrowth of planted seedlings was quite slow;they reached on average 2.6 in in height 5years after planting. In C6te d'Ivoire floweringtrees have mainly been recorded in October-November and ripe fruits in February-Marchand November. In Nigeria trees flower inMarch-May and fruitsripen in April-June andDecember. Fruits usually open while still at-tached to the tree. The seeds with their wingsare dispersed by wind; they spin during falling

Ecology Schreberu orboreo occurs mostly indry deciduous forest and semi-deciduous forest,up to 1500 in altitude. It is occasionally foundin thickets in savanna regions and at forestedges, and in Uganda it occurs in mixed rain-forest.

Propagation and planting Schrebero or-boreo is propagated by seed and wildlings. Ma-ture fruits should be collected from the tree

before dehiscence. They are usually dried inthe sun until they open to collect the seeds.There are about 10,000 seeds per kg. Seeds canbe stored for some time in a dry and coollocali-ty. It is recommended to sow seeds with wingsup; they can be sown without pre-treatment.They start germinating 2-4 weeks after sow-ing, with a moderate germination rate.

Management In most regions within thedistribution area of Schrebero orboreo, trees

.

351

Schrebero arbore0 - I, 60se of 601e, ' 2, 110u!eringtwig, ' 3, fruit, ' 4, seedRedrown und adopted by J. M. de Vries

4 fertile seeds. Seeds with a large wing, up to4(-5) cm x I cm. Seedling with epigeal germi-nation; hypocotyl c. 3 cm long, epicotyl c. I cmlong, glabrous; cotyledons fleshy, flattened, c.1.5 cm long; first leaves opposite.

Other botanical information Schrebero

comprises about 8 species, of which 5 occur intropical Africa including Madagascar, 2 in trop-ICal Asia and I in South America. It seems

most closely related to Coinoronthus fromComoros and Madagascar, which also has awoody capsule.Schrebero orboreo has been included in

Schrebero golungensis WeIw. , but the latter isprobably a synonym of Schrebero trichoclodoWeIw

Anatomy Wood-anatomical description (IAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(10: vessels in radial multiples of 4 or morecommon); 13: simple perforation plates; 22:intervessel pits alternate; (23: shape of alter-nate pits polygonal); 24: intervessel pits minute(S 4 prn); 30: vessel-ray pits with distinct bor-

SCHREBERA 573

4

3

a,

11,

I

574 TIMBERS2

occur scattered and are not common in the

forest. They are sometimes planted in bananaand cocoa plantations. Planted trees can bemanaged by coppicing and pollarding.

Genetic resources Schrebero arboreo is

widespread and harvested on a moderate scaleonly. Therefore, it is unlikely to be threatenedby genetic erosion notwlthstanding being un-common. It has been recorded as endangered inTogo and it Is rare in Ghana

Prospects Although Schrebero arboreo isconsidered a valuable timber tree of local jin-

portance, very little information is available onits growth rates and proper management prac-tices. It deserves further research as a poten-tial multipurpose species that could be usefulin agroforestry systems. In Uganda Schreberoo760reo has been recommended for planting asornamental tree and as shade tree in coffee,cocoa and banana plantations

Major references Aubr6ville, 1959c; Bolza &Keating, 1972; Burki11, 1997; ITvine, 1961;Katende, Birnie & Tengnas, 1995; Keay, 1989;Liben, 1973; Takahashi, 1978; Turrm, 1952;Vivien & Eaure, 1985.

Other references Adjanohoun at al. (Edi-tors), 1988; Adjanohoun at a1. , 1989; Ako6gni-nou, van der Burg & van der Maesen (Editors),2006; Aubr6ville, 1950; de in Mensbruge, 1966;Eggeling & Dale, 1951; Garzuglia, 2006; Green,1963; Hawthorne, 1995; Hawthorne & Jong-kind, 2006; Idu, Erhabor & Efijuemue, 2010;Kupicha, 1983b; Louppe, 1993; Neuwinger,2000; Normand & Paquis, 1976; Tailfer, 1989;Taylor, 1960; Wallander & A1bert, 2000

Sources of illustration Keay, 1989; Vivien& Faure, 1985.

Authors S. Pentsil & E. A. Obeng

.

SCO!opio inoddgoscoriensis - wild

Madagascar, possibly SCOlopio mudogoscori-errsis, has been tested. The heartwood is grey-ish to pale brown or slightly reddish, and dis-tinctly demarcated from the yellowish white,3-5 cm wide sapwood. The grain is straight tointerlocked, texture rather coarse. The wood isheavy, with a density of 890-950 kg/ms at 12%moisture content, and hard. It should be airdried slowly and with care to avoid seriousdegrade. The rates of shrinkage are moderateto high, from green to oven dry 5.5% radial and11.2% tangential. A board of 2.5 cm thick airdried in 3-4 months with little distortion. Once

dry, the wood is rather unstable in service. At12% moisture content, the modulus of ruptureis 189 N/mm2, modulus of elasticity 15,200N/min2, compression parallel to grain 77N/mm2, shear 21 N/mm2 and Chalais-Meudonside hardness 6.8-7.6. The wood is fairly easyto work and takes a good finish. It is moderate-Iy durable and rather difficult to treat withpreservatrves.

Botany Evergreen, small to medium-sizedtree up to 25 in tall, glabrous except for theinflorescence; hole up to 60(-100) cm in diame-ter; bark surface nearly smooth to scaly, grey-ish, with longitudinal rows of lenticels, innerbark reddish. Leaves arranged spiralIy, simpleand entire; stipules minute, caducous; petiole0.5-I cm long; blade elliptical to oblong-elliptical or obovate, 6-Ile14) cm x 2.5-7(-9)cm, base broadly curieate, apex acuminate,leathery, pinnately veined with many pairs oflateral veins and with 2 distinct basal lateral

veins ascending towards the upper part of theleaf. Inflorescence an axillary raceme I-2.5(-5)cm long, minutely hairy. Flowers bisexual,

SCOLOPIAMADAGASCARIENSIS SIeumer

Protologue Repert. Spec. Nov. Regni Veg.45: 14 (1938).

Family F1acourtiaceae (APG: Salicaceae)Origin and geographic distribution SCOlo-

pig mudogoscoriensts is endemic to Madagas-car, where Is widespread along the easterncoast.

Uses The wood, known as 'menavahatra'inMadagascar, is used for heavy construction,heavy flooring, joinery, panelling, mine props,vehicle bodies and railway sleepers, and also asfirewood. The fruit is eaten. A tea made from

the leaves is drunk against fever.Properties A sample of SCOlopio wood from

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regular, 4-5-merous, whitish; pedice1 3-5 minlong; sepals ovate, 1.5-2 mm long; petals simi-Iar but slightly narrower than sepals; disksmall, lobed; stamens 80-100; ovary superior,glabrous, I-celled, style 3-4 mm long, slender,stigma obscurely 3-4-lobed. Fruit a globose toellipsoid berry I-1.5 cm in diameter, fleshy, 2-5-seeded. Seeds rounded, flattened, c. 2 mm indiameter. Seedling with epigealgermination.SCO!opio inodoguscoriensis trees grow slowlyIn an enrichment planting, trees reached onaverage only I in tall 7 years after plantingYoung trees tolerate some shade. The floweringperiod is usually AprilJuneSCOlopio comprises about 40 species and occursfrom tropical Africa through tropical Asia toAustralia. In mainland Africa about 6 speciesare found, 14 in Madagascar and I in the Mas-carene islands. The wood of some other SCOlo-

pm spp. of the rainforest of eastern Madagas-car is used for similar purposes as that ofSCO!opto mudogoscoriensis.SCO!opio erythrocorpo H. Ferner is a largeshrub or small tree up to 8(-15) in tall. Itswood is used for similar purposes as that ofSCOlopio mudogoscoriensis. In traditional med-ICine its leaves are applied as a poultice againstrheumatic pain. Its fruits are eaten.SCO!opio her20mby H. Perrier is a small to me-dium-sized tree up to 20 in tall. Its wood isused for similar purposes as that of SCOlopiomudogoscoriensis. It has edible fruits.SCOlopio orientalis SIeumer is a smalltree upto 15 in tall. Its wood is used for similar pur-poses as that of SCOlopio ingdogoscuriensis.SCOlopio heterophyllo (Lain. ) SIeumer is ashrub or smalltree up to 7(-15) in tall, endem-IC to Mauritius, Reunion and Rodrigues. Itswood has been used for similar purposes asthat of SCOlopio mudogoscoriensis in Madagas-car. In Mauritius and Reunion leaf decoctions

are taken against fever. However, SCO!opioheterophylla has become rare because of forestdegradation

Ecology SCO!opio mudogoscoriensis occursin rainforest up to 1000(-1900) in altitude, inregions with about 2500 mm annual rainfall,usually on latentic soils

Management There are 25,000-33,000 seedsper kg. Fresh seeds have a high germinationrate, up to 95% in 3.5-15 weeks. They can bestored in a coollocality, but lose their germina-tion capacity after 4-6 months

Genetic resources and breeding SCOlopiomadogoscoriensis is widespread and althoughits habitat is shrinking, it is not yet listed as

threatened by genetic erosion.Prospects The wood of SCO!opio mudogos-

cone, Isis and some other SCOlopio spp. fromMadagascar is likely to remain of local jin-portance. However, trees seem to grow tooslowly to have good prospects for timber plan-tations.

Major references Blaser at a1. , 1993; Boi-teau, Bolteau & NIOrge-Bolteau, 1999; Brownat a1. , 2009; SIeumer, 1972b; Styger at a1. , 1999.

Other references Gu6neau, 1971a; Gunb-Fakim, 2007; Gunb-Fakim & Brendler, 2004;Gunb-Fakim, Gu6ho & Bissoondoya1, 1996;Lavergne, 2001; Parant, Chichignoud & Rako-tova0, 1985; Ferner de in Bathie, 1946c; Rako-tovao at al. , en preparation; Sallenave, 1971;SIeumer & Bosser, 1980.

Authors L. FA. Oyen

SCOLOPiA 575

SCOLOPIAZEYHERl(Nees) Harv.

Protologue F1. cap. a 584 (1862).Family F1acourtiaceae (APG: Salicaceae)Vernacular names Thorn pear(En). MgOvi-

govi(Sw)Origin and geographic distribution SCOlo-

pig gayheri occurs from Rwanda, Kenya andUganda southward to Angola, Botswana, Zim-babwe, Mozambique, eastern South Africa andSwaziland. It has also been reported fromGarnero0n

Uses The wood of Solopio zeyheriis used inhouse construction and for poles, tool handlesand spoons, and also as firewood and for char-coal production. Because of its strength andhardness, it has been used for teeth of millwheels and for axles, felloes and spokes in

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SCOlopio zeyheri- wild

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wagon-making.Ripe fruits are fleshy and sweet, and eaten rawas an occasional snack. In East Africa SCOlopiogayheriis an important bee plant. In SouthAfrica the foliage is a preferred browse ofgoats. Maasai people use the roots against ve-nerealdiseases

Properties The wood is very strong, hardand rather heavy, with a specific gravity (inois-ture content O%) of 0.69. It is difficult to saw,but polishes well.

Botany Much-branched shrub or smalltreeup to 15(-25) in tall, glabrous except for theinflorescence; hole often branching fairly low,up to 60 cm in diameter, sometimes withbranched spines; bark surface fissured, darkgrey to brownish, inner bark rather thin;branches and twigs often with strong, axillary,simple, straight spines up to 20 cm long.Leaves arranged spiralIy, simple; stipules inI-nute, caducous; petiole up to 1.5 cm long; bladeobovate to oblanceolate or elliptical, 2-10 cm xI-6 cm, base curieate, apex acute to obtuse orrounded, margin entire, wavy or bluntlytoothed, leathery, pinnately veined with 3-6pairs of lateral veins. Inflorescence an axillaryraceme I-5(-6) cm long, short-hairy. Flowersbisexual, regular, (3-)4-5(-6)-meTous, scented;pedice1(I-)2-5(-10) mm long; sepals ellipticalto ovate, I-1.5 mm x I min; petals smallerthan sepals or absent; disk small, lobed; sta-mens (20-)30-40; ovary superior, glabrous, I-celled, style 1.5-2.5 min long, stout, stigmaobscurely 2-3-lobed. Fruit a globose berry 7-8min in diameter, fleshy, purple-red when ripe,2-3-seeded. Seeds angular.SCOlopio comprises about 40 species and occursfrom tropical Africa through tropical Asia toAustralia. In mainland Africa about 6 speciesare found, 14 in Madagascar and I in the Mas-carene islands. In East and southern Africa,several other SCOlopio spp. are used for similarpurposes as SCOlopiu gayheriSCO!opio inundii(ECkl. & Zeyh. ) Warb. is asmall tree up to 7(-20) in tall, occurring in dryevergreen forest and wooded grassland in Zim-babwe and South Africa. Its pale brown, heavy,hard and strong wood was formerly consideredvaluable for furniture and wagon-making, butlarge specimens are now rare. The bark is usedin traditional Zulu medicine in the treatment

of heart problems, but has been implicated incases of lethal poisoning.SCOlopio rhomniphyl!o GIIg is a shrub or smalltree up to 15 in tall, locally common in rainfor-est or dry evergreen forest from Garnero0n east

to Kenya and south to Tanzania and Angola.The wood is hard and used for poles, tool han-dles and yokes, and also as firewood and forcharcoal production. The fruits are edible andthe flowers are a source of nectar for honeybees. A vapour bath from the leaves is taken asemetlc.

SCOlopio stolzii GIIg ex SIeumer is a small,much-branched tree up to LOG15) in tall, oc-curring in dry evergreen forest or woodedgrassland from Cameroon, DR Congo and Ken-ya south to Zimbabwe, Mozambique and SouthAfrica. The wood is hard and used for poles,tool handles and yokes, and also as firewoodand for charcoal production. The fruits are edi-ble. In Tanzania women drink a root decoctionto treat boils on the breasts.

SCOlopio theifolio GIIg is a much-branchedshrub or smalltree up to 15 in tall with bole to25 cm in diameter, occurring in dry evergreenforest and wooded grassland in southern Su-dan, Ethiopia, Kenya, Uganda, Tanzania andnorthern Malawi. The wood is hard and used

for poles and toolhandles, and also as firewoodand for charcoal production. The fruits are edi-ble. The tree is used for shade. Maasai peopleof Kenya use a root preparation to treat vene-real diseases.

Ecology SCOlopio zeyheriis found in dryevergreen forest, riverme forest and woodedgrassland, up to 2400 in altitude, often openrocky and sandy sites

Genetic resources and breeding SCOlopiozeyheriis common in East Africa and SouthAfrica, but rare in southern tropical Africa.There are no indications that it is in danger ofgenetic erosion.

Prospects The wood of SCOlopto zeyheri, likethat of other SCO!opio spp. , is likely to remainof local importance when hardness and tough-nes are more importantthan workability

Major references Coates Palgrave, 2002;Heme & Heme, 1988; Ruffo, Birnie & Tengnas,2002; SIeumer, 1972b; SIeumer, 1975.

Other references Beentje, 1994; Breebaart,Bhikraj & 0'Connor, 2002; Burring, 2005;Campbell & Mon, 1977; Kitula, 2007; Lovett ata1. , 2007; Neuwinger, 2000; Roux, 2003; Wild,1960

Authors L. P. A. Oyen

SCORODOPHLOEUSZENKERIHarms

Protologue Bot. Jabrb. Syst. 30(I): 78 (1901).Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Vernacular names Garlic tree, divida (En).

Arbre a ail, divida (Fr).Origin and geographic distribution SCOro-

dophioeus zenheriis distributed in Cameroon,Equatorial Guinea, Gabon, Congo, DR Congoand Cabinda (Angola).

Uses The wood, traded as 'divida', is suitablefor poles and piles in construction, flooring,joinery, mine props, ship building, vehicle bod-Ies, furniture, railway sleepers, ladders, agri-cultural implements and tool handles. In DRCongo it is used for making charcoal.The wood, bark, young leaves and seeds arelocally used as condiments, giving a garlic-liketaste to food. The young leaves are eaten as avegetable after cookingIn traditional medicine in Gabon, bark mru-sions are taken for the treatment of constipa-tion. Mixed with chillies and bitter eggplantfruits, they are taken to cure colds and cough.The bark is used in fumigations against rheu-matism, and the pounded bark is taken for thetreatment of headache. In Congo bark decoc-tions are drunk, used in vapour baths or sitzbaths, or vaginalIy administered for the treat-merit of bronchitis and genito-urinary problemsand as an aphrodisiac. In DR Congo they areapplied in an enema against the complicationsof measles, whereas decoctions of the bark orleaves are used as an urinary antiseptic andare drunk as arithelmintic. The bark adminis-

tered as an enema is a purgative, and a macer-ation of the inner bark is drunk for the same

Bark macerations are also used forpurpose

wound care, taken for the treatment of an en-larged spleen and used in nasal instillationsagainst headache. Unspecified bark prepara-tions are used for the treatment of hernia.

Production and international trade The

wood is only used IOCaUy and nottraded on theinternational market. The bark and seeds are

sold on markets in Cameroon, EquatorialGuinea, Gabon and DR Congo

Properties The heartwood is yellowish withpurplish brown streaks, the sapwood is whiteThe grain is often irregular or interlocked, tex-ture fine. The wood has an attractive figure. Itemits a garlic-like smellThe wood is heavy, with a density of 765-930kg/in' at 12% moisture content. It has a ten-dency to distort and check during drying, andmild kiln-drying schedules must be used. Theshrinkage rates are high, from green to ovendry 6.0% radial and 10.1-12.0% tangential.Once dry, the wood is unstable in service. At12% moisture content, the modulus of ruptureis 130-199 N/min", modulus of elasticity13,800-17,100 N/min', compression parallel tograin 59-90 N/min', shear 10.5-15.5 N/min',cleavage 23-26 N/min and Chalais-Meudonside hardness 6.3-80

The wood is difficult to work and tends to pickup during planing. It takes a good polish andturns well. It splits easily upon nailing, makingpre-boring necessary. It glues satisfactorilyThe wood is durable, being resistant to fungalattack but liable to termite attack. The sap-wood is susceptible to Lyetus attacks.Wood fibres of a sample from DR Congo wereon average (0.70-)1.26(-1.74) mm long and(IIJ20(-33) prn wide, with a cell wall thick-ness of (I-)5(-11.5) pin and a lumen width of(2-)Toe25.5) prn. The wood contains 42-51.5%cellulose, 29-33% 11gnin, 11-17% pentosan,0.7-0.8% ash and little silica. The solubility is1.2-1.6% in alcohol-benzene, 0.9-1.6% in hotwater and 12.5-14.7% in a I% NaOH solution

Pulplrig experiments gave paper of mediocrequality onlyThe garlic-hke scent and taste of the bark andother plant parts is due to the presence of SUI-phur containing compounds. The main constit-uents of essential oil from the bark are 2.4,5.7-tetrathiaoctane, 2.3,5-trithiahexane, 2.3,4.6-to-trathiaheptane and 2.4,5.6,8-pentathianonaneEssential oil from the bark and several of its

constituents have shown activity against vari-ous fungi and bacteria. Methanol extracts ofthe bark and, to a lesser extent, the seeds have

SCORODOPHLOEUS 577

.

SCOrodophloeus genhert - wild

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578 TIMBERS2

shown anti-oxidant activity. Stem bark ex-tracts have shown in-vitro antiprotozoal activi-ty against fryponosom0 67ucei and Letshmo-itto infontum, without showing cytotoxicityThe powdered stem bark has shown a dose-dependent protective effect against damage tostored beans by the bean beetle ACUntho-seendes o6tectus. Seeds from DR Congo con-tamed 12.1% fat, with as main constituentspalmitic acid (49%) and oleic acid (46%)

Description Medium-sized to large tree upto 40 in tall; hole branchless for up to 25 in,straight and cylindrical but sometimes irregu-Iar, up to 100 cm in diameter, without but-tresses but fluted at base; bark surface initiallysmooth, later scaly, yellowish grey, inner barkbrittle, yellowish; all plant parts, but especiallythe bark, with a garlic-like scent. Leaves alter-nate, imparipinnately or panpinnately coin-pound with (3-)5-10 pairs of leaflets; stipulesfree, linear-lanceolate, caducous; petiole andrachis together up to 10 cm long, grooved,slightly winged; leaflets alternate, sessile, ob-long or oblong-lanceolate, 1.5-5.5 cm x 0.5-2.5cm, base obliquely truncate, apex rounded orobtuse, margins entire, finely hairy below, pin-

nately veined. Inflorescence an axillary or ter-minal, dense raceme 5-8 cm long, hairy; bractslinear-lanceolate, 4-5 min long. Flowers bisex-ual, nearly regular; pedice1 I-7 mm long, within the lower half 2 caducous bracteo}es 2-3 mm

long; sepals 4, free, ovate or oblong, 4-6 min x2-3.5 min, reflexed; petals 5, free, nearly equal,obovate, 7-11 mm x 2-3 mm, hairy at margins,white; stamens 10, free, alternating longer andshorter, I-2 cm long; ovary superior, ellipsoid,hairy, with distinct stipe, I-celled, style c. I cmlong. Fruit an oblong pod 7.5-14 cm x 3-5 cm,flattened, with 0.5-I cm long stipe, pointed atapex, smooth, brown, dehiscent with 2 curlingvalves, I-2-seeded. Seeds ovoid to disk-shaped,c. 2 cm in diameter, flattened, shiny, brown.Seedling with epigeal germlnation; hypocotylI-4 cm long, epicoty1 4-14 cm long, hairy; coty-ledons rather thin, green; first 2 leaves alter-nate with nearly opposite leaflets

Other botanical infonnation SCOrodophloeuscomprises 3 species, and is restricted to tropi-calAfrica.

SCOrodophloeus lischeri(Taub. ) J. Leonard (Swa-hill names: ingodoma, inhande) is a medium-sized tree up to 30 in tall, distributed in coastalforests in Kenya and Tanzania, possibly also innorthern Mozambique. Its wood is used forpoles in construction, furniture and tool han-dles. It is also used as firewood. Twigs are usedfor tying. Root decoctions are taken as arithel-mintic. The flowers are visited by honey bees.SCOrodophloeus torrei Lock is a shrub or smallto medium-sized tree up to 20 in tall, endemicto coastal forest and thickets in Mozambique.

Anatomy Wood-anatointcal description (IAWAhardwood codes):Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: SLmple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 25: intervessel pits small(4-7 prn); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 42: mean tan-gential diameter of vessellumina 100-200 prn;47: 5-20 vessels per square minimetre; 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; 70:fibres very thick-walled. Axial parenchyma:(76: axial parenchyma diffuse); 78: axial paren-chyma scanty paratrachea1; 79: axial paren-chyma vasicentric; (83: axial parenchyma con-fluent); 85: axial parenchyma bands more than

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three cells wide; 86: axial parenchyma in nar-row bands or lines up to three cells wide; 89:axial parenchyma in marginal or in seeminglymarginal bands; 93: eight (5-8) cells per paren-chyma strand. Rays: 97: ray width I-3 cells;104: all ray cells procumbent; (106: body raycells procumbent with one row of uprightand/or square marginal cells); 1/5: 4-12 raysper mm; 1/6: a 12 rays per mm. Mineral inclu-SIons: 136: prismatic crystals present; 137:prismatic crystals in upright and/or square raycells; 138: prismatic crystals in procumbent raycells; 142: prismatic crystals in chamberedaxial parenchyma cells(S. N'Danikou, P. E. Gasson & E. A. Wheeler)

Growth and development Seedling growthis fast. The tree has a deep taproot

Ecology SCOrodoph!oeus zenheri occurs inmixed evergreen forest on well-drained localI-ties. It does not tolerate waterlogging. It islocally common and sometimes gregarious

Propagation and planting SCOrodoph!oeuszenheriis easily propagated from seed. Seedscan be sown in nursery beds at a depth of 2-3cm, in loose and deep soil. Germination is usu-ally rapid, and the germination rate highSeedlings have been planted out when theywere 2-3 years old and 1.5-2 in tall; the use ofsupports for seedlings has been recommendedSeeds can also be sown directly in the field.

Management In forest near Kisangani inDR Congo, SCOrodoph!oeus genkeriis the mostabundant tree species with an average densityof 23 trees with a bole diameter of more than

10 cm per ha. In forest in Gabon, a mean woodvolume of 0.4 mama has been reported. InCameroon trees are protected when forest landis cleared, and the species is sometimes plant-ed in coffee and cacao plantations, for whichwildlings or seedhngs from nurseries are used.Seedlings can be pruned and lopped to obtainstout holes and straight branches, which willmake harvesting of the bark easier

Harvesting The bark is harvested through-out the year, mainly by women. Staggered re-inoval of bark pieces, for instance every 2-3years, gives thick bark of good quality. Often,however, too much bark is removed, resultingin the death of the tree.

Handling after harvest Before being usedas condiments, the seeds and bark are driedand ground into powder or paste.

Genetic resources Although SCOrodoph!oeuszenkeriis locally overexploited, it is currentlynot considered a threatened species

Prospects The wood of SCOrodoph!oeus zen-

heriis heavy, hard and durable, but difficult towork and unstable in service, and very littleinformation is available on its uses. The tree is

a popular source of garlic-like condiments, ob-tamed from various plant parts. As the speciesis locally overexploited, it may be worthwhileto promote its planting. It is considered to havepotential for planting in tree plantations aswell as in crop fields, because it has a deep rootsystem and its open crown does not give toomuch shade

Major references Agbor at a1. , 2005; Au-br6ville, 1970; Bolza & Keating, 1972; Dupriez& De Leener, 1989; Eyog Matig at al. (Editors),2006; Kouokam, Jahns & Becker, 2002; Kouo-kam, Zapp & Becker, 2001; Sallenave, 1955;Vivien & Faure, 1985; Wilczek at a1. , 1952

Other references Allen & Allen, 1981;Beentje, 1994; Brenan, 1967; Burgess &Clarke, 2000; Fouarge, Quoilin & Roosen, 1970;1stas, Raekelboom & Heremans, 1959; Kawuk-pa & ingoy0, 1994; Konda ku Mbuta at al. ,2010; Koona at a1. , 2007; Lovett at a1. , 2007;Mukaya at a1. , 2003; Musuyu Muganza at al. ,2012; Neuwinger, 2000; Pakia & Cooke, 2003;Raponda-Walker & Sillans, 1961; Sindani &Ndjele, 1982; Takahashi, 1978; Ternu, 1990;Termote, Van Darnme & Dhed'a Djai10, 2010;Weiss, 1973.

Sources of illustration Aubr6ville, 1968;Wilks & TSSemb6, 2000.

Authors M. Brink

SCOTTELLIA 579

SCOTTELLIAKLAINEANA Pierre

Protologue Bull. Mens. SOC. Linn. Paris,ser. 2, 14: 1/3 (1899)

Family F1acourtiaceae (APG: AChariaceae)Synonyms Scottellio homerunensis Gilg

(1908), Scottellio cheuolieri Chipp (1923), SCOt-tento congceo Achev. ex Hutch. & Dalziel

(1927)Vernacular names Akossika (Fr)Origin and geographic distribution Scottel-

lad hloineono occurs from Sierra Leone east to

the Central African Republic and south to DRCongo and northern Angola.

Uses The wood of Scottel!to hloineono, trad-ed as 'akossika' or 'odoko', is used for posts inhouse building, light flooring, joinery, interiortrim, ladders, sporting goods, toys, novelties,food containers, spoons, ladles, combs, toolhandles, draining boards, carving and turneryIt is suitable for furniture, cabinet making,veneer and plywood. It is also used as firewood

580 TIMBERS2

I.

Scottellio Momeono - wild

Roots and bark are used in traditional medi-cine. Root decoctions are taken as diuretic and

purgative, and to treat oedema, and bark de-coctions to treat stomach-ache. In Nigeria, thewood ash is used by the Yoruba people in soapmaking.

Production and international trade The

export of 'akossika' wood from C6te d'TVoirewas 58,000 in31n 2004, 28,000 main 2005 and16,000 ina in 2006

Properties The heartwood is pale yellow topale yellowish brown, occasionally with darkerstreaks, and indistinctly demarcated from theup to 10 cm wide sapwood. The grain is usuallystraight, sometimes interlocked, texture usual-Iy fine. Quarter-sawn surfaces show an attrac-tive silver grain figure. The wood is fairly Ius-trous, odourless and tasteless when dried.The wood is medium-weight, with a density of590-750 kg/in' at 12% moisture content, inod-erately hard and tough. It air and kiln driessatisfactorily, but is liable to checking, splittingand warping, especially in thick boards. Therates of shrinkage are moderately high, fromgreen to oven dry 4.1-5.4% radial and 9.2-11.6% tangential. During drying, the wood issusceptible to blue stain attack. Once dry, it ismoderately stable in service.At 12% moisture content, the modulus of rup-ture is 82-140 N/mm2, modulus of elasticity8000-14,700 N/min2, compression parallel tograin 45-70 N/min2, shear 8-15 N/min2 cleav-age 17-25 N/min, Janka side hardness 4840-7480 N, Janka end hardness 7200-8110 N andChalais-Meudon side hardness 2.5-4.3.

The wood saws and works well with both ina-

chine and hand tools. It takes a good finish and

polishes well. It is slightly brittle and quarter-sawn surfaces tend to flake. It has a tendencyto split upon nailing; pre-boring is advised. Ithas good screw-holding properties. The woodpaints and varnishes easily. The gluing proper-ties are good. Slicing and peeling should bedone with care because the veneer has a ten-

dency to break across. The wood is not durable,being liable to fungal, termite, pinhole borers,Lyetus and marine borer attacks. It is permea-ble to impregnation with preservatives.

Description Evergreen, medium-sized treeup to 30(-35) in tall; hole branchless for up to20 in, usually straight, up to 100(-130) cm indiameter, slightly fluted at base or with smallbuttresses; bark surface fairly smooth to sright-Iy scaly, greenish grey to dark brown, innerbark thin, granular, orange brown to reddishbrown, rapidly darkening to brown upon expo-sure, with colourless exudate and almond-likesmell; crown often small, narrow and dense;twigs glabrous, with lenticels. Leaves alter-nate, simple; stipules absent; petiole 0.5-I(-2)cm long, rather slender; blade obovate to obo-vate-oblong or elliptical, 5-16 cm x 3-9 cm,broadly curieate at base, usually acuminate atapex, margins entire to wavy or shallowlytoothed, leathery, glabrous, pinnately veinedwith 5-8 pairs of lateral veins. Inflorescence anaxillary raceme 4-10 cm long, slightly hairy.Flowers bisexual or male, regular, 5-meTous,creamy white, scented; pedice1 3-5(-8) minlong; sepals free, rounded to elliptical, 3-4 mmlong; petals oblong-obovate, (4-)5 min long,hairy at margins, at the base inside with ascale-like hairy appendage; stamens alternat-ing with petals, 3-5 mm long; ovary superior,glabrous, I-celled, style short, 3-lobed at apex.Fruit a nearly globose capsule (3-)4-8(-9) minin diameter, finely warty, dehiscing with 3valves, I-2(-3)-seeded. Seeds globose-angular,up to 5 mm in diameter, smooth, covered by athin orange-red anI. Seedling with epigealgermination; hypocoty1 4-5 cm long, epicotyl0.5-I cm long, hairy; cotyledons leafy, rounded,I-2 cm long; first leaves alternate.

Other botanical information Scotte!lid

comprises 3 species and is restricted to Westand Central Africa. It seems to be related to

DOSylepis, Erythrospermum and Rowsonio.Scottellio Ieonensis 011v. , a small to medium-sized tree up to 20 in tall with hole up to 30 cmin diameter, occurring in coastal swamp forestand gallery forest from Guinea Bissau east toLiberia, differs from Scottellio Maimedrid in itsthicker and often larger leaves with thicker

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mean tangential diameter of vessellumina 50-100 pin; 42: mean tangentialdiameter of vesselIumina 100-200 prn; 47: 5-20 vessels persquare minimetre; 48: 20-40 vessels per squareminimetre. Tracheids and fibres: 61: fibres

with simple to minutely bordered pits; 62: fi-bres with distinctly bordered pits; 65: septatefibres present; 66: non-septate fibres present;69: fibres thin- to thick-walled. Axial paren-chyma: 75: axial parenchyma absent or ex-tremely rare. Rays: 98: larger rays commonly4- to 10'seriate; 102: ray height > I min; 103:rays of two distinct sizes; 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; 110: sheath cellspresent; 1/5: 4-12 rays per min. Mineral inclu-SIons: 136: prismatic crystals present; 137:prismatic crystals in upright and/or square raycells; (140: prismatic crystals in chamberedupright and/or square ray cells); (159: silicabodies present); (160: silica bodies in ray cells)(L. Awoyemi, P. E. Gasson & E. A. Wheeler)

Growth and development The growth isslow. In C6te d'Ivoire an investigation of growthrings showed a mean annual hole diametergrowth of 4.5 mm for the first 30 years, 6 minfor trees of 30-80 years old and 5 min for treesof 80-140 years old. In field trials the meanannual diameter growth was only 2-3 minAlthough Scotte!Ito Hornedito is usually ever-green, it is reported to be deciduous for up to 2months (October-November) in Sierra LeoneIn West Africa, flowering has been recorded inFebruary-August and fruits ripen in May-De-cember. In Gabon trees flower in December-

January and have ripe fruits in January-AprilThe seeds with their red anIs are probablydispersed by birds, but monkeys also eat theanIs

Ecology Scottel!to hloineono most common-Iy occurs in lowland evergreen forest, but canalso be found in semi-deciduous forest. It often

occurs in small stands along rivers andstreams, but may also be common in rocky 10-canties. It is sensitive to forest fires

Propagation and planting There are about25,000 seeds per kg. Although seedlings areoften common in the understorey close tomother trees, the germination rate of seed hasbeen reported to be low, only 15-25% 8-12days after sowing. Saplings and young treesare fairly shade tolerant, but older trees arereported to be light demanding. Regenerationis commonly observed in undisturbed forest.

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Scotte!Jin kiwineon0 - I, 60se of 601e, . 2, flower-ing twig, . 3, flower, ' 4, fruit, ' 5, seedRedrawn grid udopted by W We88el-Brond

5

SCOTTELLIA 581

I

petioles. Its wood is possibly used for similarpurposes

Scotte!jig orientalis GIIg, a medium-sized tofairly large tree up to 40 in tallwith hole diam-eter up to 100 cm, occurring from Nigeria eastto southern Sudan and south to DR Congo, isdifficult to distinguish from Scottellio h!ot-neono, except by Its larger fruits. Its wood isprobably used for similar purposes.Scotte!Ito foldineono is variable, with 2 varie-ties being distinguished, connected by transi-tions

Anatomy Wood-anatomical description CAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(12: solitary vessel outline angular); 14: scalari-form perforation plates; 15: scalariform perfo.ration plates with s 10 bars; (16: scalariformperforation plates with 10-20 bars); 22: inter-vesselpits alternate; 23: shape of alternate pitspolygonal; 26: intervessel pits medium (7-10pin); 27: intervessel pits large ^ 10 prn); 31vessel-ray pits with much reduced borders toapparently simple: pits rounded or angular; 41:

2

582 TIMBERS2

Management Scotte!lid Maineono usuallyoccurs scattered in the forest or in smallgroupsof trees. In forest in western Gamero0n, a meandensity of 0.15 tree with a hole diameter above60 cm has been recorded per ha, with an aver-age wood volume of 0.9 in31ha. In C6te d'Ivoire7-27 trees with a hole diameter of more than

10 cm have been recorded per ha. Moderatethinning resulted in better growth of youngtrees.

Handling after harvest After harvest, logsshould be removed rapidly from the forest ortreated with preservatives because the wood issusceptible to blue stain attack. Fresh logsfloat in water and can thus be transported byriver.

Genetic resources Scottellio k!Qineo, IQ is

locally common and in most regions within itsdistribution area there are no indications of

overexploitation. Therefore it is unlikely to beliable to genetic erosion. However, in someregions, e. g. in C6te d'Ivoire, exploitationseems to be considerable and may cause sen-ous pressure on populations of the species

Prospects Presently, Scottellio hidineonohas limited importance on the internationaltimber market, but its multipurpose wood mayhave good prospects for further commercialIza-tion. However, information on available stands,growth rates, silviculture and propagation isvery limited and much research is still neededto be able to develop sustainable managementsystems for this poorly known species

Major references ATIBT, 1986; Bolza &Keating, 1972; Burki11, 1994; CIRAD ForestryDepartment, 2008; Cooper & Record, 1931;Guiscafre, 1977; Hawthorne, 1995; 0teng-Amoako (Editor), 2006; Savill & Fox, 1967;SIeumer, 1972c.

Other references Aubr6ville, 1959c; Barnps,1968; Bertault at a1. , 1999; Blackett & Gar-dette, 2008; Chase at a1. , 2002; de in Mens-bruge, 1966; Derienne, 1974; Hawthorne &Gyakari, 2006; HUI, 1995; ITvine, 1961; Keay,1954b; Keay, 1989; Neuwinger, 2000; Normand& Paquis, 1976; Raponda-Walker & Sillans,1961; Sallenave, 1955; Takahashi, 1978; Vivien& Faure, 1985; White & Abernethy, 1997;Wilks & Issemb6, 2000.

Sources of illustration Aubr6ville, 1959c;HUI, 1995; Wilks & TSSemb6, 2000.

AuthorsA. Addo-Danso

SCYPHOCEPHALIUMMANNll(Benth. & Hook. f. )Warb.

Protologue Notizbl. K6nigl. Bot. Gart. BerlinI: 98 (1895).

Family MyristicaceaeSynonyms Scyphocepholium ochocoo Warb

(1897).Origin and geographic distribution Scypho-

00pholittm monnii occurs from south-easternNigeria to Gabon and Congo

Uses The wood, known as 'sorro' or 'OSsoko'in Gabon, is used for interior joinery, furniture,cabinet work, veneer and plywood, and tradi-tionaUy for canoes. It is suitable for light con-struction, light flooring, interior trim, toys,novelties and turnery. It is also used as fire-wood and for charcoal productionThe seeds contain an edible fat, and they areused as condiment or spice in fish dishes andsauces. They are also used as bait in traps forporcupines and squirrels. The bark is used intraditional medicine. Bark decoctions are ad-

ministered to treat gonorrhoea, sterility inwomen and cough. The bark is also an ingredi-ent of a preparation to treat convulsions. Driedbark mixed with clay is applied to stop exces-SIve flow of milk

Production and international trade The

wood of Scyphocepholium monniiis rarelytraded on the international market;it is almostexclusively used locally

Properties The heartwood is brown to or-ange-brown towards the centre of the hole,with some greyish streaks, but it has an up to10 cm wide band of yellowish white or palegreyish brown wood outwards. It is indistinctlydemarcated from the narrow sapwood. The

.

Scyphocepho!turn indrinii- wild

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grain is straight, sometimes interlocked, tex-ture moderately coarse. The wood is lustrous,and greasy to the touch. It is lightweight tomedium-weight with a density of 460-650(-740) kg/ina at 12% moisture content. It air drieseasily and fast, without serious defects. Boardsof 2.5 cm thick take about one month to air dryfrom 87% to 16% moisture content. The rates of

shrinkage are fairly small, from green to ovendry 2.5-4.2% radial and 4.8-7.8% tangential.Once dry, the wood is moderately stable inservice

At 12% moisture content, the modulus of rup-ture is 84-124 N/min2, modulus of elasticity(6660-)9310-14,110 N/min2, compression par-allel to grain 29-54 N/min2, shear 4-8 N/min2,cleavage 9-20 N/min and Chalais-Meudon sidehardness (1.4-)2.9-5.8.The wood is easy to saw and work with bothhand and machine tools, with slight bluntingeffectto saw teeth. It can be planed and sandedto a smooth surface. The nailing and gluingproperties are satisfactory. The wood varnisheswell, but the heartwood near the centre of thehole may be somewhat difficult to paint. It hasgood slicing and peeling characteristics, alt-hough the logs often have a poor form. Thewood is not durable, being susceptible to ter-mite and borer attacks, butthe heartwood nearthe centre of the hole is resistant to dry-woodborers. The heartwood is moderately resistantto impregnation with preservatives, the sap-wood is permeable. The wood contains 38-41%ceUulose, 33% 11gnin, 11.5% pentosan, 0.3% ashand 0.1% silica. The solubility is 11.0% in alco-hol-benzene, 3.1% in cold water and 21.4% in aI% NaOH solution.

The aromatic seeds are rich in lipids. Hexaneextracts showed the presence of triglycerides ina solid form. Two cyclohgnans have been iso-

of whichlated from leaf extracts, one

(ocholignan A) showed significant in-vitro anti-bacterial activity against methicillin-resistantStophylococcus oureus

Adulterations and substitutes The wood

resembles that of Pycnonthus origolensis(WeIw. ) Warb. and is used for similar purposes.

Description Evergreen, dioecious, medium-sized to fairly large tree up to 35(-45) in tall;hole branchless for up to 20 in, straight orslightly twisted, up to 140(-300) cm in diame-ter, often fluted at base or with small buttress-es; bark surface scaly with large irregularscales, greyish brown with reddish markings,with lenticels, inner bark slightly fibrous, pink-ish, rapidly becoming reddish, with a reddish

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SCYPHOCEPHALIUM 583

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Scyphocepholium monnii - I, 60se of 601e, ' 2,port o1/10uiering twig, . 3, mole flower, 4, fernoleflower, ' 5, port of twig with fruits.Redrown grid odopted by Ishoh Syomsudtn

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layered, with horizontalexudate; crown

branches; young twigs reddish brown short-hairy. Leaves alternate, simple and entire;stipules absent; petiole c. I cm long; blade ob-long-lanceolate, 20-25 cm x 4-8 cm, cordate atbase, acuminate at apex, thin-leathery, reddishbrown short-hairy below but becoming gla-brous, pinnately veined with 18-25(-35) pairsof lateral veins. Inflorescence an axillary cymeup to 4(-6) cm long, short-hairy, with flowersin heads, each head with 2 bracts and up to50(-100)-flowered. Flowers unisexual, regular,up to 5 min long, reddish, with short pedicel,perianth cup-shaped, 3-5-lobed, reddish brownshort-hairy; male flowers with 6-10 fused sta-mens, filaments merged into a column; femaleflowers with superior, ovoid, slightly hairy, I-celled ovary, stigma 2-lobed. Fruit an irregulardepressed-globose drupe 2.5-5 cm x 3-6(-7.5)cm, in bunches of up to 4, yellowish to orangewhen ripe, brown short-hairy, flesh abundant,resinous, dehiscent with 2 valves, I'SeededSeed flattened, c. 3 cm in diameter, with entirearlI.

I

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584 TIMBERS2

Other botanical information Scyphoce-phonum comprises a single species, possibly 2because Scyphocepholium chrysothrix Warbhas been described based on a single specimencollected over 100 years ago in Gamero0n; thelatter species seems to have larger fruits.

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; (12:solitary vessel outline angular); 13: simple per-foration plates; (14: scalariform perforationplates); (15: scalariform perforation plates withS 10 bars); (16: scalariform perforation plateswith 10-20 bars); 22: intervessel pits alternate;23: shape of alternate pits polygonal; 27: inter-vessel pits large ^ 10 pin); 31: vessel-ray pitswith much reduced borders to apparently sim-PIe: pits rounded or angular; 32: vessel-ray pitswith much reduced borders to apparently SIm-PIe: pits horizontal(scalariform, gash-like) tovertical (palisade); (33: vessel-ray pits of twodistinct sizes or types in the same ray cell); 42:mean tangential diameter of vessellumina100-200 pin; 46: S 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:78: axial parenchyma scanty paratrachea1; 79:axial parenchyma vasicentric; (89: axial paren-chyma in marginal or in seemingly marginalbands); (92: four (3-4) cells per parenchymast"and); 93: eight (5-8) cells per parenchymastrand; (94: over eight cells per parenchymastrand). Rays: 97: ray width I-3 cells; (100:rays with multiseriate portion(s) as wide asuniseriate portions); 102: ray height > I min;107: body ray cells procumbent with mostly 2-4rows of upright and/or square marginal cells;108: body ray cells procumbent with over 4rows of upright and/or square marginal cells;(109: rays with procumbent, square and up-right cells mixed throughout the ray); 115: 4-12 rays per min. Mineral inclusions: 159: silicabodies present; 160: silica bodies in ray cells(R. Shanda, E. Uetimane, P. Baas & H. Beeck-man)

Growth and development In Gabon treesflower mainly in October-December and fruitsripen in January-April. The seeds are occa-SIonally eaten by gorillas.

Ecology Scyphocepholium monniiis wide-spread in lowland rainforest up to 550 in alti-tude, but is most common in secondary forest

It usually occurs on well-drained soils.Management In Gabon extensive stands of

Scyphocepholtum monnii have been recordedin evergreen forest, with a estimated standingvolume in 1973 of 20-30 million ms. In the1960s, the average wood volume was estimatedat 3.2 in8tha. In Equatorial Guinea an averagedensity of I tree of more than 70 cm in holediameter per ha has been recorded. In forest inwestern Cameroon the average density of treeswith a hole diameter of more than 60 cm is 0.4

per ha, with an average wood volume of 5.4in8tha. The tree can be coppiced

Harvesting Larger holes may have brittleheart, especially when they are more than 90cm in diameter, and very large holes are oftenhollow. In felling, trees with their comparative-Iy lightweight wood may have difficulty to fallwhen there are lianas in the crown

Handling after harvest Boles are suscep-tible to splitting and they should be well sup-ported when they are cross-cut into logs afterfelling. Logs should be removed from the forestrapidly or treated with preservatives to avoidattacks by fungi and borers. They float in wa-ter and thus can be transported by river.Boards and planks should be treated with afungicide immediately after sawing and beforedrying.

Genetic resources Scyphocepho!turn mon-nitis quite widespread in different forest typesincluding secondary forest, and is locally coin-mon. There are no indications that it is threat-ened by genetic erosion

Prospects Although the wood of Scyphoce-phonum monniiis not considered of premiumquality, it has a decorative two-coloured ap-pearance and has good prospects for applica-tions such as panelling and moulding. It maybe interesting for further commercialization,providing timber that, although not durable,can be used for various purposes includingveneer and plywood production. Its commonoccurrence in secondary forest formations mayimply good prospects for sustainable produc-tion in natural forest, but research on growthrates, regeneration and proper managementsystems are needed. The often more or lessfluted hole is a drawback, especially for rotaryveneer production.Further phytochemical and pharmacologicalinvestigations of the bark and leaves are rec-ommended to assess the value of their applica-tions in traditional medicine and antibacterialactivity. The seed may have commercial jin-portance for the production of edible fat.

Major references Bolza & Keating, 1972;Burki11, 1997; CIRAD Forestry Department,2008; CTFT, 1971; Fouilloy, 1974; Nziengui,2001; Sallenave, 1971; Takahashi, 1978; Vivien& Faure, 1985; Yvon, 1973

Other references Bouquet, 1969; Chudnoff,1980; de Saint-Aubin, 1963; Foullloy, 1965; Huat a1. , 2005; Keay, 1989; Neuwinger, 2000;Normand & Paquis, 1976; Painbou Tchivoundaat a1. , 1992; Raponda-Walker & Sillans, 1961;Sallenave, 1955; Sallenave, 1964; Senterre &Nguema, 2001; Tailfer, 1989; White & Aber-nethy, 1997; Wilks & Issemb6, 2000.

Sources of illustration Fouilloy, 1965;Tailfer, 1989; White & Ahemethy, 1997

Authors R. H. M. J. Lemmens

SCYTOPETALUMTiEGHEMii(Achev. ) Hutch.& Dalziel

such as tympanites, and as purgative. Decoc-tions of the twigs are added to a bath to treatskin diseases including leprosy.

Production and international trade The

wood of Soytopetolttm tieghemiiis only usedlocally and not traded on the internationalmarket

Properties The heartwood is yellowish topinkish, turning yellowish brown to greyishbrown upon exposure, and indistinctly demar-cated from the pinkish sapwood. The grain isstraight to interlocked, texture moderatelycoarse. The wood has little lustre

The wood is medium-weight to fairly heavy,with a density of 710-770 kg/ms at 12% ino1s-ture content, moderately hard but tough andstrong. It should be air dried with care to avoidserious checking and end splitting. The rates ofshrinkage are high. At 12% moisture content,the modulus of rupture is 123-148 N/min2,modulus of elasticity 13,430 N/min2, coinpres-SIon parallel to grain 55-59 N/min2, cleavage26.5 N/mm, Janka side hardness 7310 N,Chalais-Meudon side hardness 2.3 and Janka

end hardness 7910 N

The wood is rather difficult to saw and work. It

finishes well, but surface's may be woolly due tothe presence of interlocked grain. It holds nailswell, but pre-boring is recommended. The woodglues satisfactorily, and is suitable for veneerand plywood production. It is not durable, be-ing liable to fungal and termite attacks. Theheartwood is resistant to impregnation withpreservativesLeafy twigs contain saponosides, tannins andsterols.

Adulterations and substitutes The wood

of Soytopetolum tieghemiiis similar to that ofOubonguio spp

Description Evergreen, small to medium-sized tree up to 25(-40) in tall; bole branchlessfor up to 15(-20) in, usually straight and cylin-drical, up to 60(-100) cm in diameter, some-times with smallbuttresses; bark surface scaly,dark grey, inner bark rather thick, spongy-fibrous, pinkish, with unpleasant smell; crownnarrow and dense, with drooping branches;twigs slender, glabrous. Leaves distichouslyalternate, simple and entire; stipules absent orminute and early caducous; petiole c. 2 minlong; blade elliptical to narrowly elliptical, 4.5-13 cm x 2-4 cm, curieate at base, distinctlyacuminate at apex, papery to thin-leathery,glabrous, pinnately veined with 4-5 pairs oflateral veins, and with numerous very fine,transverse, parallel veinlets. Inflorescence a

Protologue F1. W. trop. Mr. I: 238 (1927)Family Scytopetalaceae (APG: Lecythidaceae)Chromosome number2n=22

Vernacular names Sourwood (En). Mous-sangou6 (Fr)

Origin and geographic distribution Soyto-petalurn tieghemiiis restricted to West Africa,where it occursfrom Sierra Leone to Ghana.

Uses The wood is used in Sierra Leone and

Ghana for poles in house building, and also fortelegraph poles. It is suitable for joinery, mineprops, ship building, railway sleepers, ladders,sporting goods, agricultural implements, toolhandles, veneer and plywood.The gum from the wood has been used as glueThe fruits are edible. Bark preparations aretaken to treat cough, intestinal complaints

SCYTOPETALUM 585

I.

Scytopetolum tieghemii - rolld

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586 TIMBERS2

4

purposes as that of Soytopetolum tieghemii.Soytopetulum Alumeonum Pierre is a medium-sized to large tree up to 40 in tall with holebranchless for up to 25 in and up to 100 cm indiameter, occurring from Cameroon south toCabinda (Angola). Its yellowish white wood ismoderately heavy and moderately hardSoytopetolumpterreonum (De Wild. ) Tiegh. is asmallto medium-sized tree up to 30 in tallwithhole up to 80 cm in diameter, occurring inCameroon, Central African Republic, Congoand DR Congo.

anatomy Wood-anatointcal description (IAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; (14: scalariformperforation plates); (15: scalariform perforationplates with s 10 bars); 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;25: intervessel pits small (4-7 pin); 26: inter-vessel pits medium (7-10 pin); 31: vessel-raypits with much reduced borders to apparentlysimple: plts 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 prn; 47: 5-20 vessels per square inil-limetre. Tracheids and fibres: 61: fibres with

simple to minutely bordered pits; 66: non-septate fibres present; 70: fibres very thick-walled. Axial parenchyma: 77: axial parenchy-ina diffuse-in-aggregates; 78: axial parenchymascanty paratrachea1; 86: axial parenchyma innarrow bands or lines up to three cells wide;(88: axial parenchyma scalariform); 93: eight(5-8) cells per parenchyma strand; 94: overeight cells per parenchyma strand. Rays: 98larger rays commonly 4- to 10'seriate; 102: rayheight > I mm; (103: rays of two distinct sizes);106: body ray cells procumbent with one row ofupright and/or square marginal cells; 107: bodyray cells procumbent with mostly 2-4 rows ofupright and/or square marginal cells; 1/5: 4/2

Mineral inclusions: 136rays per mm pris-

matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells.(E. E. Mwakalukwa, P. E. Gasson & E. A. Wheeler)

Growth and development Scytopetolumtieghemiiis classified as a shade bearer. Treesflower in February-April and fruits mature inMay-July. The flowers open in the morningand petals and stamens already shrivel andfall in the evening of the same day. The fruits

2

Scytopetolum tieghemii - I, leaf und old milo-rescence, 2, inflorescence with old flowers gridflower bud, 3, longitudino! section o1/10coer, 4,fruiting twig.Redrouin und odopted by G. WE. Don den Berg

@b

short axillary raceme, glabrous. Flowers bisex-ual, regular, whitish, fragrant; pedicel flat-tened on one side, c. 0.5 cm long, elongating toc. 1.5 cm in fruit; calyx cup-shaped, small, with2 minute teeth; petals 5-8, slightly unequal,oblong to lanceolate, 5-7.5 min long; stamensnumerous, about as long as petals and fused atbase wlth these; ovary superior, ovoid, usually6-celled, style short, stigma head-shaped. Fruitan ovoid drupe up to 2 cm x I cm, acute withpersistent style, red when ripe, I-seeded. Seedswith ruininate endosperm. Seedling with epi-geal germination; hypocoty1 5-6 cm long, epi-cotyl c. I cm long, glabrous; cotyledons leafy,remaining within the fruit stone; first 2 leavesopposite

Other botanical information The so-called

petals of the flower should probably be consid-ered as staininodes, while true petals are lack-Ing

Scytopetolum comprises 3 species and is restrict-ed to West and Central Africa. The wood of the

2 Central African species is suitable for similar

3

are eaten by birds such as turacos and horn-bills, which are probably the main seed dis-persers, but also by monkeys. However, it hasalso been reported that the fruits fall in quan-tity to the ground below the mother tree.

Ecology Soytopetolum tieghemii occurs inforest and moist semi-deciduousevergreen

forest, usually in the middle storey. In Ghanait is reported to prefer acid, base-poor soils.

Propagation and planting There are about2000 fruit stones per kg, which take 2-5months for germination. The germination rateis low, 20-25%. Seedlings develop a taprootwith short lateral roots.

Management Soytopetolum tieghemtioccurslocally abundant, e. g. in Liberia and southernC6te d'Ivoire, but is in other regions less coin-mon, e. g. in Sierra Leone and Ghana. In forestsin western Gabon, the average wood volume ofSoytopetolum hloineonum is 4 ina per ha.

Handling after harvest Freshly felled logsshould be removed from the forest rapidly or betreated with preservatives because the wood issusceptible to blue-stain fungal attack

Genetic resources Soytopetolum tieghemiiis fairly widespread in West Africa, locallycommon and currently not heavily exploitedTherefore, there are no reasons to consider itliable to genetic erosion

Prospects In regions where it occurs abun-dantly, Soytopetolum tieghemii has been con-sidered promising as commercial timber pro-ducer for special applications such as construc-tion work where durability is not required.Locally in Gabon Soytopetolum Momeonumoccurs in fair densities and may be of interestfor commercial exploitation. However, data ongrowth rates and regeneration are needed toassess the possibilities for sustainable coinmer-cialproduction

Major references Appal, 1996; Aubr6ville,1959a; Bolza & Keating, 1972; Burki11, 2000;Cooper & Record, 1931; de in Mensbruge, 1966;Hawthorne & Jongkind, 2006; Kryn & Fobes,1959; Takahashi, 1978; Taylor, 1960

Other references Adjanohoun & Ak6 Assi,1979; Bouquet & Debray, 1974; Breteler, 2002;Breteler, 2005; de Saint-Aubin, 1963; Germain,1963; Hawthorne, 1995; Irvine, 1961; Keay,1958k; Letouzey, 1978a; Letouzey, 1978b;Neuwinger, 2000; Normand, 1955; Normand &Paquis, 1976; Sallenave, 1955; Savill & Fox,1967; Vivien & Faure, 1985

Sources of illustration Appe1, 1996; Au-br6ville, 1959a; Hawthorne & Jongkind, 2006.

Authors R. H. M. J. Lemmens

SEARSiALANCEA (L. f. ) FA. Barkley

Protologue Luridell, F1. Texas a 104 (1943)Family AnacardiaceaeChromosome number2n=30

Synonyms Rhus loneeo L. f. (1781)Vernacular names Karee, African sumac,

willow rhus (En).Origin and geographic distribution Seorsio

loneeo is indigenous in Zambia, Namibia, Bot-swana, Zimbabwe, South Africa and Lesotho. Itis planted in hedges and as ornamental tree,e. g. in South Africa, but also in desert regionsof the United States and Mexico, where it islocally naturalized

Uses The tree holes are considered valuable

as fence poles, and the wood is used for imple-merit handles and wagon parts. Branches havebeen used for bows and for spars in hut build-ing. Young twigs are occasionally used for bas-ketry. The fruits are edible; they are poundedwith water and produce a beer after fermenta-tion, and they are also used to produce curdledmilk. Searsio loneeo is commonly planted inhedges, as an ornamental tree in gardens andparks, as a roadside tree and in live fences andwind breaks. Roots, leaves and stem bark areused to treat skin diseases. A root infusion is

taken to treat abdominal and chest complaintsand diarrhoea, and the roots are chewedagainst stomach-ache. Leaf decoctions or mru-SIons are taken to treat measles and pustules,

are inhaled to curewhereas leaf vapours

cough. The bark and wood have been used fortanning, and the bark gives a brown dye. Theleaves are browsed by livestock, especially inthe dry season.

Properties The wood is reddish brown, with

SEARSiA 587

.

Seorsio lanced - wild

L.

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588 TIMBERS2

a fine texture, and it is hard, tough and dura-ble. It is heavy, with a density of 890-970kg/in3 at 10% moisture content. The rates ofshrinkage are moderate, from green to ovendry about 3.1% radial and 7.2% tangential. Thewood works well and takes a nice polish. It hasa sweetish and spicy smellBark extracts showed in-vitro antibacterial

activity against a wide range of gram-positiveand gram-negative bacteria. Leaves yield 0.2%of essential oil, with orpinene, benzene and 6-3-carene as main constituents. The oil showed

remarkable anti-oxidant activity and dose-dependent antibacterial and antifungal activi-ties. These activities may be associated withthe high concentration of orpinene in the oil(87%). The flavour of cattle milk is affectedwhen large quantities of Seorsio loneeo leavesare browsed.

Botany Evergreen, dioecious shrub or smalltree up to 9(-12) in tall, glabrous; hole oftentwisted and crooked; bark surface dark grey todark brown, rough, often irregularly fissured;twigs reddish brown, angular or slightlygrooved, pendulous. Leaves arranged spiralIy,3-foliolate; stipules absent; petiole 2-7 cm long,slender but slightly winged near apex, groovedabove; leaflets sessile, narrowly lanceolate,often slightly sickle-shaped, 7-24.5 cm x 0.5-3cm, curieate at base, acute at apex, marginsentire, thinly leathery, pinnately veined withnumerous lateral veins. Inflorescence an axil-

Iary or terminal, much-branched panicle up to9 cm long, lax. Flowers unisexual, regular,usually 5-merous, greenish yellow; pedice1 2-3mm long; calyx segments c. 0.5 mm long, ob-tuse; petals free, oblong, c. 1.5 mm long; sta-mens usually 5, free; disk cup-shaped; ovarysuperior, globose, I-celled, styles 3, thick, re-curved; male flowers without ovary or withstrongly rudimentary ovary, female flowerswith rudimentary stamens. Fruit a depressed-globose drupe 4-6.5 mm in diameter, oftenslightly asymmetric, dull yellow to greyish orbrown, with fleshy pulpSearsio lanced grows fairly fast, up to 80cm/year in height for young trees. It flowersfrom April to September(-January) and ripefruits can be found from September to Febru-

The flowers have a sweet smell and areary

visited by insects such as bees, which serve aspollinators. The fruits are eaten by birds suchas guinea-fowl, francolins and bulbuls, whichare probably the main seed dispersers.Seorsto comprises approximately 110 speciesand occurs in southern Europe, Asia and Mri-

ca. Southern Africa is by far richest in species.Until recently, most authors did not separateSeorsio from Rhus, although the separationhas already been proposed at the beginning ofthe 1940s. Recent phylogenetic research usingDNA and gene spacer data confirmed thatSearsio is distinct from Rhus sensu stricto,which is limited to the Northern Hemisphere.Several other Seorsio spp. produce wood that isused for similar purposes as that of Seorsioloneeo. The wood of Searsio gluttnoso (Hochstex ARIch. ) Moffett (synonyms: Rhus o6ysstnico011v. , Rhus gluttnoso Hochst. ex A. Rich. ), ashrub or smalltree up to 10 in tallfrom Sudan,Eritrea and Ethiopia, is used for agriculturalimplements and tool handles, and as firewoodThe leaves are used in traditional medicine to

treat influenza, the roots in veterinary medi-cine to treat udder complaints. They have beenused for tanningThe reddish, hard and tough wood of Searsiogueirtzii (Sond. ) FA. Barkley (synonym: Rhusgueinzii Sond. ), a shrub or smalltree up to 8 intall occurring in Zimbabwe, southern Mozam-bique and eastern South Africa, is used for walllaths in house building. Branches are used totreat eye complaints; they are made into a 10-tion, or smoke from burning twigs is applied tothe eyes. A root infusion is taken to treat schis-tosomiasis and a leaf decoction against gallsickness.

The stems of Seorsio lucido (L. ) FA. Barkley(synonym: Rhus lucido L. ), a shrub or smalltree up to 7 in tall from Zimbabwe and SouthAfrica, are valued for fence posts. The bark hasbeen used for tanning. Ground bark is appliedto ring worm infections.The stems of Seorsio pendulino (Jacq. ) Moffett(synonyms: Rhus pendulino Jacq. , Rhus Dimi-nons auct. non Aiton nec Vahl), a smalltree upto 10 in tall occurring in Namibia and SouthAfrica, are used as fence posts and in hutbuilding, whereas the twigs are used for fishtraps. The bark yields a reddish brown dyeThe fruits are edible and have been used in

beer making. Seorsio pendulino is planted asornamental tree.

The wood of Seorsio retinorrhoeo (Steud. ex011v. ) Moffett (synonym: Rhus retinorrhoeoSteud. ex Onv. ), a shrub or smalltree up to 6 intall from Sudan, Eritrea, Ethiopia, Somaliaand Arabia, is used for agricultural imple-merits and walking sticks, and as firewood. Theleaves serve for local mattresses. Several na-

vonoids have been isolated from the leaves,some of which showed weak antibacterial and

antimalarial activities in vitro

Ecology Seorsi0 10nceo occurs in grasslandand open woodland, often on river banks andtermite mounds. It Is considered as an Indica-

tor of underground water. It seems to prefersoils rich in lime. Once established, the tree isdrought and frosttolerant

Management Searsio juncea is easy to pro-pagate by seed, cuttings, truncheons and airlayering. Ripe seeds should be sown in seedlingtrays and can be transplanted in containerswhen the seedlings have 2 leaves. Cuttingstaken from young shoots often root successful-Iy.The tree holes are often crooked, and straightpieces of wood of fair length are usually diffi-cult to obtain. Moreover, the hole of older treesis reportedIy often hollow. A disease, causingleaves at the extremities of young branches tobecome deformed, has been recorded from thecentral highlands of Namibia. In South Africathe fungus Muribosidiosporo indied was Iden-tified as the causal organism associated withleaf spots, a serious disease in several Andcor-didceoe

The fruits are usually rubbed between thehand palms to remove the tough skin beforethey are being eaten, or are soaked in milk.

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

Prospects The holes of Searsio loneeo andother Seerrsio spp. are too small and poorlyshaped to yield timber of sufficient dimensionsand quality for the international market, butlocally the wood will continue to be valued fore. g. fences and implements. The tree has con-siderable ornamental value, and has a non-aggressive root system and provides shadethroughout the year with its evergreen, wide-spreading crown. The pharmacological proper-ties of the leafoil deserve more attention.

Major references Gundidza at a1. , 2008;Moffett, 1993; Palmer & Pitman, 1972-1974;Stern, 2002; van Wyk & Gencke, 2000.

Other references Bekele-Tesemma, 2007;Coates Palgrave, 1983; Orous, Groenewald &Carr011, 2003; Fernandes & Fernandes, 1966;Moffett, 2007; Neuwinger, 2000; Obi at al. ,2003; van Vuuren, Banks & Stohr, 1978.

Authors R. H. M. J. Lemmens

SENNAIACTEA 01atke) DU Puy

Protologue Kew Bull. 50(3): 581 (1995)Family Caesalpiniaceae (Leguminosae - Cae-

salpinioideae)Synonyms Cussio IOCteo Vatke (1880)Origin and geographic distribution Senno

IOCteo is endemic to northern Madagascar, southto Marianara.

Uses The wood is used for furniture, and alsoas firewood and for charcoal production. SennoIOCteo is planted as a shade tree in coffee plan-tations

Properties The wood is reportedIy yellowishand strong.

Botany Deciduous small to medium-sizedtree up to 30 in tall; bole up to 150 cm in diam-eter; bark surface grey, often inottled; twigsshort-hairy. Leaves alternate, panpinnatelycompound with (IIJ15-22(-25) pairs of leaf-lets; stipules awl-shaped, 5-7 mm long, earlycaducous; petiole 2.5-4 cm long, rachis up to 30cm long, grooved; petiolules 2-3 mm long; leaf-lets opposite to alternate, oblong, 2-5.5 cm x0.5-1.5 cm, rounded to notched at apex, gla-brous or sparsely hairy, pinnately veined. In-norescence a terminal or axillary raceme 5-12cm long, often combined into a lax panicle atends of shoots, short-hairy. Flowers bisexual,slightly zygomorphic, 5-meTous; pedice1 12-18min long; sepals free, unequal, 3-8 mm long,reflexed, tinged red; petals free, nearly equal,oblong-elliptical, 10-11 mm long, white, some-times tinged pink; stamens 10 of which 7 fertileand up to 9 min long, 3 rudimentary and up 5mm long; ovary superior, 6-7 min long, style 5-6 min long, curved. Fruit a strap-shaped, flatpod 16-20 cm x 3-4 cm, with transverse lines

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between the seeds, glabrous, reddish brown toblackish, dehiscing with 2 thin valves remain-ing fused at one margin, up to 22-seeded. Seedsoblong, flat, c. I cm x 0.5 cm, smooth, brown.Senno IOCteo can be found flowering from Aprilto AugustUntilthe early 1980s, Cussio was considered avery large genus of about 550 species, but wasthen split into 3 genera: Cossio s. s. with about30 species, Chomoecristo with about 250 spe-cies and Senno with about 270 species. Sennois very similar to Cossio, but is distinguishedfrom it by the possession of 3 adaxial stamenswhich are short and straight, and the pedicelswhich have no bracteoles

Madagascar has 9 endemic Senno species. Ad-ditional to Senno IOCteo, some of these havedocumented uses of their wood. Sen, to mendi-

orions (R. Vig. ) DU Puy is a shrub or smalltreeup to 5 in tall restricted to south-western Ma-dagascar. The branches are used as poles inbuilding local houses. Senno Diguierel!0 (Ghesq. )DU Puy is a shrub or smalltree up to 8 in talloccurring in southern Madagascar. Its wood isalso used in house construction.

Senno stdineo (Lain. ) H. SITwin & Barneby, asmallto medium-sized tree up to 20 in tallorig-mating from tropical Asia, is planted in west-ern Madagascar and also in mainland tropicalAfrica. Although it is most commonly used forfirewood and as ornamental, it is sometimesalso planted for poles because of its durablewood.

Ecology Senno IOCteo occurs in humid ever-green forest from sea-level up to 1600 in alti-tude, often along rivers, but it can also befound in degraded vegetation.

Genetic resources and breeding A1houghSenno IOCteo has a comparatively small area ofdistribution, it has not been reported to be en-dangered. It has been recorded as locally coin-mon as an adventive in degraded vegetationand thus may survive in areas subject to forestdegradation caused by human intervention.

Prospects Too little is known about SennoIOCteo to judge its prospects as a timber tree ofmore commercial importance. It may have POS-sibilities as an auxiliary plant and may be in-teresting for planting in agroforestry systemsHowever, much research is still needed, e. g. onwood properties, growth rates and propagation

Major references du Puy at a1. , 2002Other references Capuron, 1968.Authors R. H. M. J. Lemmens

SINDORAKLAiNEANA Pierre ex Penegr.

Protologue Bull. Mus. natn. Hist. nat. , Par-is 25: 653 (1919).

Family Caesalpiniaceae or. ,eguminosae - Cae-salpinioideae)

Origin and geographic distribution Sindo-ro hloineono has a small distribution area,restricted to western Equatorial Guinea, west-ern Gabon and south-western Congo.

Vernacular names Banda rouge(Fr)Uses In Gabon the wood, known as 'n'gom6'

or 'rigom', is used for joinery and that of thebuttresses for implements. It is suitable forlight construction, interior trim, furniture,boxes, crates, draining boards, veneer, ply-wood, hardboard and particle board.

Production and international trade The

wood is only used locally and not traded on theinternational timber market.

Properties The heartwood is pinkish brownto pinkish grey, darkening upon exposure witha coppery tinge, and indistinctly demarcatedfrom a pale reddish intermediate zone, which ismore distinctly demarcated from the whitishyellow sapwood. The grain is variable, texturemedium. Wood surfaces show a decorative fig-ure of dark and pale stripes. The wood containsa resinous exudate becoming dark upon drying.It is medium-weight, with a density of 610-690kg/ina at 12% moisture content, and moderatelyhard. It air dries slowly but well, with littletendency to splitting and distortion. The ratesof shrinkage are moderate. Once dry, the woodis moderately stable in service. At 12% ino1s-ture content, the modulus of rupture is 103-128 N/min2, compression parallel to grain 46-49 N/min2, cleavage 12 N/mm and Chalais-

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The wood saws and works well with both hand

and machine tools, with little blunting effect onsaw teeth and cutting edges. It usually produc-us a nice finish except when interlocked grainis present. The resin in the wood may interferewith gluing, polishing and painting. The nail-Ing and screwing properties are good, withsatisfactory holding power. The wood peelswell, producing good-quality Theveneer

heartwood is fairly durable, but slightly sus-ceptible to termite attack, whereas the sap-wood is liable to Lyetus attack. The heartwoodis resistant to impregnation with preserva-tives, the sapwood moderately resistant

Description Medium-sized to large tree upto 40(-45) in tall; hole branchless for up to 30in, straight and cylindrical, up to 130(-200) cmin diameter, without buttresses but slightlythickened at base; bark surface fairly smoothbut finely fissured, greenish grey, inner barkmoderately thick, fibrous, pinkish brown, withsome transparent sticky exudate; twigs short-hairy, becoming glabrous. Leaves alternate,imparipinnately compound with 6-10 leaflets;stipules absent or early caducous; petiole and

rachis together 6-14 cm long; petiolules c. 0.5cm long; leaflets alternate, oblong to elliptical,5-9 cm x 3-4 cm, usually rounded at base, ob-tuse to acute at apex, leathery, glabrous, withmany translucent dots, pinnately veined withmany lateral veins, inIdrib below with gland atdistal end. Inflorescence an axillary or termi-nal panicle up to 20 cm long with branches upto 10 cm long, densely yellowish brown short-hairy. Flowers bisexual, zygomorphic, yellow-ish white; pedicel c. 2.5 mm long; sepals 4,fused at base, oblong-lanceolate, unequal, c. 8mm long, one larger than other 3, brown short-hairy outside; petal I, elliptical, c. 6 mm x 4mm, brown hairy outside, often 4 additionalminute petals present; stamens 2, with 8 rudi-meritary stamens, all fused but I rudimentarystamen free; ovary superior, obliquely ovoid,slightly hairy, with short stipe, style slender,glabrous. Fruit a flattened nearly round pod 5-7 cm in diameter, glabrous, transverselyveined, dehiscing with 2 woody valves, withshort stipe, I(-2)-seeded. Seeds flattened ovoid,c. 4 cm x 3 cm, glossy black. Seedling with epi-geal germination; hypocoty1 11-13 cm long,epicoty1 11-15.5 cm long, finely hairy; cotyle-dons fleshy and thick; first leaves alternate,with 2-4 leaflets

Other botanical information Stridoro

comprises about 20 species, which are allfoundin tropical Asia except Stridoro h!giftedrid. Sin-doro timber from tropical Asia, traded as 'sepe-tir', is important on the international market.Stridoro is related to Stridoropsis and Tess-monnio, which differ in having I petal and 10fertile stamens, and 5 petals and 10 fertilestamens, respectively.

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 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 prn; 46: S 5vessels per square minimetre; (47: 5-20 vesselsper square minimetre); (58: gums and otherdeposits in heartwood vessels). Tracheids andfibres: 61: fibres with simple to minutely bor-dered pits; 66: non-septate fibres present; (68fibres very thin-walled); 69: fibres thin- to

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thick-walled. Axial parenchyma: (79: axial pa-renchyma vasicentric); 80: axial parenchymaallform; 81: axial parenchyma lozenge-allform;83: axial parenchyma confluent; 89: axial pa-renchyma in marginal or in seemingly margin-at bands; (91: two cells per parenchymastrand); 92: four (3-4) cells per parenchymastrand; (93: eight (5-8) cells per parenchymastrand). Rays: 97: ray width I-3 cells; 98: Iarg-er rays commonly 4- to 10'seriate; (104: all raycells procumbent); 106: body ray cells procum-bent with one row of upright and/or squaremarginal cells; 1/5: 4-12 rays per mm. Secre-tory elements and cambial variants: 129: axialcanals diffuse; 131: intercellular canals oftraumatic origin. Mineral inclusions: 136prismatic crystals present; 142: prismatic crys-tals in chambered axial parenchyma cells(E. K. AChi, S. N'Danikou, P. E. Gasson & P.Baas)

Ecology Sindoro hmmeono occurs in low-land rainforest near the coast and along inari-groves, sometimes at the edge of small naturalsavanna areas, up to 100 in altitude

Harvesting The minimum hole diameterallowed for harvesting in Gabon is 70 cm. Theresin present in the wood may cause difficultiesIn sawing by gumming-up saw teeth.

Genetic resources Stridoro Mumeono has a

small area of distribution and is there not

common, which makes it easily liable to genetic

Prospects Stridoru h!umeo, IQ provides a ge-neral-purpose timber of fair quality, with asonly drawback the presence of resin. However,the amounts available are too limited to have

prospects for commercial exploitation, and re-search could better focus on protection of thisspecies, which is so strangely isolated fromother species of the genus.

Major references moriymous, 1957b; Au-br6ville, 1968; Bolza & Keating, 1972; de Saint-Aubin, 1963; Leonard, 1994; Normand & Pa-quis, 1976; Raponda-Walker & Sillans, 1961;Sallenave, 1955; Takahashi, 1978; Tucker, 2003

Other references Aubr6ville, 1970; Sambasat a1. , 1993; Tailfer, 1989

Sources of illustration Aubr6ville, 1968;de Saint-Aubin, 1963.

Authors R. H. M. J. Lemmens

SINDOROPSIS LE-TESTUl(Fellegr. ) J. Leonard

Protologue Mein. ACad. Roy. Belg. , Cl. Sci.s6r. 2, 30(2): 82 (1957)

Family Caesalpiniaceae or. ,eguminosae - Cae-salpinioideae)

Synonyms GOPoifero Ie-testui(Penegr. ) Pel-legr. (1947).

Origin and geographic distribution Sindo-Fopsis Ie-testui has a small distribution area,restricted to Equatorial Guinea and Gabon.

Uses The wood, known as 'gh60mbi', is suit-able for construction, flooring, joinery, interiortrim, mine props, ship building, vehicle bodies,furniture, toys, novelties, boxes, crates, veneerand plywood.Bark decoctions are used in traditional medi-

cine; they are applied as enema to treat intes-tinal complaints and externalIy to treat head-ache and as stimulant, whereas they are ad-ministered orally as arithelmintic. Pulverizedbark is rubbed in to treat scabies. Bark ex-

tracts have been used to dye Rophio fibresblackish. The seed oilis applied as skin careproduct and perfume.

Production and international trade Thewood is traded on the international timber

market in small amounts. In 1997 Gabon ex-

ported nearly 4000 ms of logs, but in 1999 theamount had declined to about 500 in3 and to

100 ina in 2001. However, it increased again to2750 ing in 2005.

Properties The heartwood is pinkish brownto reddish brown, darkening upon exposurewith a coppery shine and slightly darkerstreaks, and distinctly demarcated from thewhitish, 8-10 cm wide sapwood turning grey-ish to brown upon drying. The grain is straight

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to slightly interlocked, texture medium tocoarse and even. The wood contains a resinous

exudate. It is medium-weight to fairly heavy,with a density of 630-810 kg/ms at 12% ino1s-ture content, and fairly hard. It air dries quitewell but slowly, with some tendency of distor-tion and checking. Air drying to 15-20% ino1s-ture content is recommended before kiln dry-ing, especially for thin boards. The rates ofshrinkage are moderate, from green to ovendry 3.0-4.8% radial and 5.5-9.0% tangentialOnce dry, the wood is moderately stable to un-stable in service. At 12% moisture content, themodulus of rupture is 1/5-201 N/mm2, inodu-Ius of elasticity 10,700-20,200 N/min2, coin-pression parallel to grain 55-70 N/mm2, shear7.5-12.5 N/mm2, cleavage 13-24 N/min andChalais-Meudon side hardness 4.5-8.8.

The wood saws and works well with both hand

and machine tools, but with some bluntingeffect on saw teeth and cutting edges; stellite-tipped saw teeth and tungsten-carbide cutting

recommended. Clogging of sawedgesblades and tools may occur due to the presenceof resin. The wood planes to a smooth finishexcept when interlocked grain is present. Thenailing and screwing properties are good, withsatisfactory holding power. The resin in thewood may interfere with polishing and paint-ing. The wood glues well. The peeling and SIIc-ing characteristics are satisfactory after steam-ing. The heartwood is moderately durable, be-ing moderately resistant to fungal, termite,borer and marine borer attacks. The sapwoodis liable to Lyetus attack. The heartwood ismoderately resistant to impregnation withpreservatives, the sapwood is permeable.The resin has been reported to be very inflam-inable

Botany Medium-sized to very large tree upto 60 in tall; hole branchless for up to 30 in,straight and cylindrical, up to 130(-200) cm indiameter, thickened at base or with roundedbuttresses; bark surface fairly smooth but fine-Iy fissured, dark grey, inner bark moderatelythick, fibrous, reddish brown, with some darkresinous exudate; twigs short-hairy, becomingglabrous. Leaves alternate, imparipinnatelycompound with 6-1/1eaflets; stipules absent orearly caducous; petiole and rachis together 8-10 cm long; petiolules c. 0.5 cm long; leafletsalternate, oblong to ovate or elliptical, 4-6 cmx 2-3 cm, usually rounded at base, short-acuminate at apex, leathery, glabrous, withmany translucent dots, pinnately veined withmany lateral veins, with a gland near attach-

merit of petiolule and another gland at distalend of inIdrib below. Inflorescence an axillaryraceme up to 10 cm long, densely yellowishbrown short-hairy. Flowers bisexual, zygomor-phic, white; pedice1 5-7 mm long, jointed atbase; sepals 4, oblong-lanceolate, unequal, 5-7mm long, one slightly shorter than other 3,short-hairy; petal I, elliptical, up to 5 min long,glabrous; stamens 10, allfused but I free, up toI cm long but alternately long and short; ovarysuperior, obliquely ovoid, hairy at margins,with short stipe, style slender, glabrous. Fruita flattened oblong to lanceolate pod 7-10 cmlong, distinctly pointed at apex, glabrous,smooth, dehiscing with 2 thin-leathery valves,with short stipe, I-2-seeded. Seeds flattenedoblong, glossy black. Seedling with epigealgermination; hypocoty1 8-10 cm long, epicotyl4-5 cm long, finely hairy; first 2 leaves OPPo-site, with 4 leaflets.Trees flower at the end of the dry season andfruits ripen around January, when they splitopen suddenly dispersing the seeds over shortdistances

Stridoropsis comprises a single species. It isrelated to Stridoro and ressmonnio, whichdiffer in having I petal and 2 fertile stamens,and 5 petals and 10 fertile stamens, respective-Iy.

Ecology Sindoropsis Ie-testui occurs in low-land rainforest up to 700 in altitude, often scat-tered but sometimes locally abundant in pri-mary forest

Management The minimum hole diameterallowed for harvesting in Gabon is 70 cmFreshly felled logs should not be left in theforest too long or treated with preservativesbecause they are susceptible to fungal and in-sect attacks. They do not float in water andthus cannot be transported by river. The green-ish resin present in the wood may cause diffi-culties in sawing by gumming-up saw teeth

Genetic resources and breeding Sindo-ropsis Ie-testui has a small area of distributionmainly in undisturbed forest, which makes iteasily liable to genetic erosion. Monitoring ofstandsis recommended.

Prospects Stridoropsis Ie-testui provides ageneral-purpose timber of fair quality, with asdrawback the presence of resin. However, theamounts available in its small distribution

area are probably too limited to have prospectsfor more extensive commercial exploitation.Knowledge on growth rates and regenerationare lacking

Major references Aubr6ville, 1968; Bolza &

are

SINDOROPSIS 593

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594 TIMBERS2

Keating, 1972; CIRAD Forestry Department,2008; de Saint-Aubin, 1963; Sallenave, 1955

Other references Aubr6ville, 1970; Christyat a1. , 2003; Maisonneuve & Manfredini(Edi-tors), 1988c; Neuwinger, 2000; Normand &Paquis, 1976; Raponda-Walker & Sillans, 1961;Sallenave, 1971; Tailfer, 1989; Takahashi,1978; White & Ahemethy, 1997.

Authors R. H. M. J. Lemmens

SLOANEARHODANTHA (Baker) Capuron

Protologue Adansonia, ser. 2, 12(3): 386(1972)

Family ElaeocarpaceaeSynonyms Eloeocorpus rhodonthus Baker

(1883), Eloeocorpus quodrilobus Jum. & H. Per-rier (1916).

Origin and geographic distribution 8100-neo rhodontho is endemic to Madagascar,where it is widespread in the central and east-ern parts of the island

Uses The wood, known as 'voanana' is tradi-tionally used for construction and planks inhouse construction, for joinery and furnitureKitchen utensils such as plates, holes andspoons are produced from the buttresses,whereas the hole is used to make dug-out ca-noes. The wood is suitable for panelling, ceil-Ing, Iathing, poles, piles, interior trim, agricul-turalimplements, containers, matches, turn-ery, veneer, plywood and particle board. Woodpulp is used for paper making and the wood forcharcoal production. The fruits are used inritual ceremonies.

Production and international trade Thewood is only used locally and not traded on the

International timber market

Properties The heartwood is purplish red topurplish brown and distinctly demarcated fromthe 5-10 cm wide, yellowish white sapwoodThe grain is variable but often wavy to inter-locked, texture medium to coarse. The wood ismedium-weight, with a density of 520-690kg/ms at 12% moisture content. The rates ofshrinkage are rather high, from green to ovendry 2.8-4.0% radial and 7.0-10.2% tangential.Partial air drying before kiln drying is recoin-mended. At 12% moisture content, the modulusof rupture is 108-137 N/min2, modulus of elas-ticity 8600-10,980 N/min2 compression paral-Iel to grain (37-)46-54 N/min2, shear 4-6N/mm2, cleavage 13-17 N/min and Chalais-Meudon side hardness 1.9-3.4

The wood saws and works easily with bothhand and machine tools, but in planing thepresence of Interlocked grain may be problem-atic. The bending properties are good. Thewood is not durable, being susceptible to fungaland insect attacks. The sapwood is liable toblue stain attack and to Lyetus borersRoot bark extracts showed weak inhibitoryactivity against strains of Plusmodium foieipo-ruin. Several phenolic compounds have beenisolated as active compounds

Description Medium-sized to large tree upto 40 in tall; hole branchless for up to 20 in, upto 120(-200) cm in diameter, with rather thinand spreading buttresses up to 2 in high; twigsglabrous to short-hairy. Leaves arranged spi-rally, more or less clustered near apex of twigs,simple; stipules minute, peg-like to linear, ca-ducous; petiole 0.5-3.5(-6) cm long, usuallyjointed at apex; blade obovate, 3.5-15(-18) cmx 1.5-9.5(-11) cm, curieate to rounded at base,obtuse to rounded or short-acuminate at apex,margins entire to slightly toothed, papery toleathery, glabrous to slightly hairy, pinnatelyveined with 3-7 pairs of lateral veins. Flowerssolitary in leaf axils of twigs, pendant, bisexu-al, regular, 4-6(-8)-meTous, large and showy;pedice1 2-4.5 cm long; sepals free, triangular,0.5-I cm long, hairy; petals free or vanablyfused, 1.5-3 cm long, irregularly lobed ortoothed at apex, dark pink to red; stamens nu-meTous, 0.5-I cm long, yellow; disk c. I cm indiameter, flat, hairy; ovary superior, conical, c0.5 cm long, hairy, (3-)4-5-celled, with a nar-rowly conical style up to I cm long. Fruit anovoid-oblong capsule 6-TICm x 3-7 cm, smoothbut with numerous small whitish lenticels,brown, with cushion-shaped persistent disk atbase, dehiscent with (3-)4-5 woody valves,

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pits horizontal(scalariform, gash-like) to verti-cal(palisade); 42: mean tangential diameter ofvessellumina 100-200 pin; 47: 5-20 vesselsper square minimetre. Tracheids and fibres:61: fibres with simple to minutely borderedpits; 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: 75:axial parenchyma absent or extremely rare; 89axial parenchyma in marginal or in seeminglymarginal bands; 92: four (3-4) cells per paren-chyma strand. Rays: 98: larger rays commonly4- to 10'seriate; 102: ray height > I min; 103:rays of two distinct sizes; 106: body ray cellsprocumbent with one row of upright and/orsquare marginal cells; 107: body ray cells pro-cumbent with mostly 2-4 rows of uprightand/or square marginal cells; (110: sheath cellspresent); 115: 4-12 rays per mm. Secretoryelements and cainbial variants: 131: intercellu-Iar canals of traumatic origin. Mineral inclu-sinns: 136: prismatic crystals present; 137:prismatic crystals in upright and/or square raycells; 154: more than one crystal of about thesame size per cellor chamber.(E. A. Obeng, P. E. Gasson & E. A. Wheeler)

Growth and development Trees flowerfrom October to March

Ecology SIoo, ,eo rhodontho occurs in dense,humid primary forest at (300-)400-2000 inaltitude. It is most common in inId-elevationforest at 800-1200 in altitude.

Management 8100neo rhodontho is locallycommon; up to 650 trees with a hole diameterof more than 10 cm have been recorded per haHowever, in general natural regeneration israted as quite poor.

Genetic resources 8100neo rhodonthu is

quite widespread in Madagascar and locallycommon. However, the on-going fragmentationof forest may make it liable to genetic erosionin the future because it is characteristic forundisturbed forest.

Prospects Research on propagation and SII-viculture is recommended to ensure sustaina-

ble management in the future for this locallyimportant but relatively poorly known species

Major references Bolza & Keating, 1972;CaO at a1. , 2006; Gu6neau, Bedel & Thiel,1970-1975; Gu6neau & Gu6neau, 1969; Sal-Ienave, 1971; Schatz, 2001; Takahashi, 1978;Tire1, 1985; Wiselius & SOSef, 1998

Other references Bolteau, Bolteau & NIOr-ge-Bolteau, 1999; Brown at a1. , 2009; Capuron,1966e; Goode, 1983; Decary, 1946; Gu6neau,1971a; Gu6neau, 1971b; Gu6neau, 1972; Mian-drimanana, 2008

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8100neo rhodontho - I, flowering twig, . 2, de-hisced fruit, . 3, seed.Redrown und udopted by J. M. de Vries

each cellwith I-10 seeds. Seeds oblong, c. I cmlong, black, with fleshy, irregularly lobed, or-ange-red anI

Other botanical information 8100neo coin-

prises about 100 species and occurs in antropi-cal and subtropical regions except mainlandAfrica. Only SIooneo rhodontho is found inMadagascarSIooneu rhodontho is variable, particularly inleaf size and shape, in hairiness of the leaves,in shape of petals and in number of stamens.Two varieties have been distinguished; varrhodontho has glabrous leaves below, var.dolechompioides (Baker) Capuron short-hairy

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: I: growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; 13: sim-PIe perforation plates; 21: intervessel pits op-POSite; 22: intervessel pits alternate; (23: shapeof alternate pits polygonal); 27: intervessel pitslarge (^ 10 prn); 31: vessel-ray pits with muchreduced borders to apparently simple: pitsrounded or angular; 32: vessel-ray pits withmuch reduced borders to apparently simple:

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Sources of illustration Tire1, 1985.Authors S. Rakotonandrasana

SONNERATIAALBASm.

Protologue Rees, Cyc1. 33(I): Sonneratia no.2 (1816)

Family Sonneratiaceae (APG: Lythraceae)Chromosome number it = 11.12Vernacular names Red-brown

(En). Mlilana, inpia (Sw)Origin and geographic distribution Sonne-

roti0 o160 Is extremely widespread. It occursalong the east coast of tropical Africa from So-mana south to Mozambique and in Comoros,Mayotte, the Seychelles and Madagascar. It isalso found throughout tropical Asia, and innorthern Australia, Vanuatu and New Galedo-nia

Uses The wood of Sonneroti0 o160 is used inKenya and Tanzania for carpentry, canoes,boat ribs, paddles, masts, floats, and windowand door frames. It is further used as firewoodand for charcoal production. In Madagascar thewood Is preferred for making paddles. In tropi-cal Asia the wood is used for house building,ship building, piles of bridges, sleepers, pavingblocks, flooring, furniture and sporting goodsSulphate pulp of the wood is suitable for inak-Ing paper

The sour fruits are eaten raw or cooked andused to make vinegar. Leaves are used forcamelfodder in Kenya. The bark is locally usedas a brownish dye and for tanning leather andfishing nets

Production and international trade In

tropical Africa there is no significant trade in

Sonnerotio timber. For Asia trade statistics arenot readily available and refer to timber of allSonnerotio species together. In 1992 about5100 in3 of logs and 1700 ina of sawn timberwere exported from Sabah at prices of aboutUs$ 731m3 and Us$ 1501m3, respectively. Verysmall amounts of Sonnerotio timber are ex-ported from Papua New Guinea.

Properties The heartwood is brown to red-dish brown and distinctly demarcated from thepaler sapwood. The wood is moderately heavy,hard and durable. It contains small amounts ofsalt, making it resistant to wood borers. It cor-rodes metal and therefore special nails andscrews are needed.

The wood is of moderate quality for firewoodAlthough it produces a lot of heat, it producesmuch ash and salt. The bark of Sonnerotiu

o160 contains 9-12% tannin based on dryweight

Botany Evergreen shrub or smalltree up to15(-20) in tall; hole often crooked and fluted atbase, without buttresses, surrounded by aerialroots arising vertically from long horizontalroots; bark surface initially smooth, greyish,becoming irregularly fissured and dark grey-brown; twigs distinctly jointed above the nodesand 4-angled. Leaves opposite, simple and en-tire; stipules absent; petiole 3-15 min long;blade elliptical to ovate, 3-13 cm x 2-9 cm,curieate at base, rounded or notched at apex,leathery, pinnately veined with 11-14 pairs oflateral veins. Flowers solitary or 3 together atthe apex of twigs, bisexual, regular, 6-8-merous; calyx tube c. 1.5 cm long, thick, Ieath-ery, persistent, lobes I-2 cm long; petals strap-shaped, I-2 cm long, caducous; stamens nu-merous, showy, white; ovary superior, 12-20-celled, style 4-6 cm long, slender. Fruit a de-pressed-globose berry, 2-3 cm x 2.5-4.5 cm,indehiscent, crowned by the style base, many-seeded. Seeds irregularly angular, c. 12 mmlong. Seedling with epigeal germination; hypo-cotylelongated; cotyledons emergentSonneroti0 o160 trees develop according toRauh's architectural model, characterized by amonopodialtrunk which grows rhythmicalIyand so develops tiers of branches. It flowers atnight and petals and stamens drop withinhours from arithesis

Sonnerotio comprises about 5 species.Ecology Sonneroti0 o160 is found at the sea-

ward edge of mangrove because it does nottolerate wide fluctuations in salt concentration

A tidal range of at least I in is necessary and itgrows along seashores and at the mouth of

mangrove

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tidal creeks on sandy, rocky or muddy soils,and also on coral terraces. It may act as a PIO-neer, colonizing newly formed sandy mud flatsin sheltered situations. It is often gregarious,but usually does not form dense stands.

Management Sonneroti0 o160 can be prop-agated by seed as well as by air-layering. It isless suited for replanting on sites where sedi-meritation is high as the sediment will coverthe pneumatophores and hence killthe plants

Genetic resources and breeding Sonnero-ti0 o160 is widespread and common in manyregions and does not seem vulnerable, alt-hough the rate of destruction of mangrove hab-itats is alarming with an estimated annual lossof I-2% worldwide

Prospects In mangrove forest Bruguieroand Rhizophoro are usually economicalIy moreimportant timber species than Sonnerotto.Replanting in damaged and logged mangrovevegetations is usually done with Auicennio andRhizophoro species and thus the importance ofSon, Ierotio for its wood is likely to decrease.However, it has very good potential as a valua-ble source of raw material for kraft pulp whichwarrants serious consideration.

Major references Boer & Lemmens, 1998b;BOSire at a1. , 2008; Duke & Jackes, 1987; Fer-nandes, 1978; Kairo at a1. , 2001.

Other references Alongi, 2008; Dabdouh-Guebas at a1. , 2000; Ellison, 1998; FAO, 2007;Nshubemuki, 1993; Schatz, 2001; Shi at al. ,2000; Williams Sangai, 1968.

Authors C. H. BOSch

.

SOYAUxlA 597

SOYAm"ACRANDIFOLLA Gilg & Stapf

Soyouxiogrondifo!to - wild

Protologue Journ. Linn. SOC. , Bot. 37: 102(1905).

Family MedusandraceaeOrigin and geographic distribution Soyou-

xio grandifolio is found in Liberia, C6te d'I-voire and Ghana

Uses The wood of Soyauxio grandifolio,known as 'abotesima' in Ghana, is mainly usedas posts for construction of huts and houses.Twigs are used as chewing-sticks for cleaningteeth

Properties The wood is dull purple-brown,hard, strong and flexible. It has a fine texture,is easy to work and finishes smoothly.

Botany Smallto medium-sized tree up to 25in tall; hole usually crooked, up to 25 cm indiameter, buttresses absent; bark surface grey-brown, thin, flaking; crown spreading; twigs

flat and with lenticels. Leaves arranged spiral-Iy, simple and entire; stipules paired, oblong, c.10 mm x 2 min; petiole short; blade oblong tolanceolate, 12-32 cm x 4-11 cm, obtuse atbase, acuminate to acute at apex, leathery,almost glabrous, pinnately veined with morethan 15 pairs of distinct lateral veins. Inno-rescence a dense raceme 6-12 cm long. Flowersbisexual, regular, 5-merous; pedicel very short;sepals c. 5 mm long, persistent in fruit, rustyhairy; petals free, whitish; corona short, tubu-Iar; stamens many, filaments long, arithers 4-celled; ovary superior, I-celled, styles 3, longand exserted. Fruit a sessile, 3-valved capsuleup to 3 cm x 2.5 cm, red-brown to purple, I-seeded. Seed trigonous, c. 2 cm in diameterSoyouxio comprises about 6 species. It hasbeen placed in Flueourtioceoe and Possi/lord-cede and resembles some Euphorbiaceoe. Morerecently it has been included in Medu-sondroceoe, but the most modern approach isto place it in Pertdiscoceoe, together with 2genera from tropical America

Ecology Soyouxio grondifolto is found inhumid, evergreen forest, sometimes in second-ary forest, on river banks and in coastal sa-vanna. It prefers regions with high annualrainfall(above 2500 mm), but can also be foundin regions with a lower mean annual rainfall,from 1500 min onwards. In Ghana it is onlycommon on very acid soils, but elsewhere it hasbeen recorded on sandy, loamy as well as later-itic soils

Management Soyouxio grandifolio regen-erates in the shade of the forest canopy

Genetic resources and breeding AlthoughSoyauxio grondifolio is not widely distributed,

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598 TIMBERS2

no serious threats are envisagedProspects The wood of Soyouxto grandifolio

is not of interest for commercial exploitationbut will continue to play a role for local uses

Major references Aubr6viUe, 1959c; Burkill,1997; Holingren at a1. , 2004; Keay, 1958f

Other references Bancroft, 1935; Bayer,2007; Brenan, 1953; Hawthorne, 1995; Haw-thorne & Jongkind, 2006; Normand & Paquis,1976

Authors C. H. BOSch

SPIROSTACHYSAFRICANA Sond

Protologue Linnaea 23: 106 (1850)Family EuphorbiaceaeVernacular names African sandalwood,

Cape sandalwood, headache tree, tamboti(En).Sandalo africano (Po). Msarakana (Sw)

Origin and geographic distribution Spi-rostochys @1ncono occurs from south-easternKenya south to Namibia, Botswana, Zimbabwe,Mozambique, South Africa and Swaziland

Uses The wood is in high demand for decora-tive joinery, furniture and cabinet work, andfor carvings and turnery. It is also appreciatedfor use in house construction for posts, beamsand roof laths, and to make scented beads fornecklaces. It is suitable for heavy flooring,mine props, shlp building, toys, novelties, agri-cultural implements and musical instrumentsIt is also used as fuelwood, but should not beused for cooking because of the toxic smoke;the branches have been used for making scent-ed torches. The wood is not appreciated forcharcoal production.The roots, bark and latex are widely used in

traditional medicine. Root decoctions are taken

to treat malaria, constipation but also diar-rhoea, cough, gonorrhoea and headache, andthey are dropped into the eyes to treat oph-thalmia. Bark decoctions and infusions in

small dosage are used as purgative to treatconstipation but also diarrhoea, and to treatstomach ulcers, kidney complaints, cough andeye complaints, and to purify the blood. Driedbark is applied to rashes in children. Powderedbark is taken as arithelmintic. Latex diluted in

water Is taken as emetic and purgative, andthe latex is administered against toothacheand as anodyne. It is also applied to sores incattle to kill maggots, whereas the wood servesas insect repellent. Extreme caution is neededwhen bark or latex is administered for medici-

nal purposes. Several complications caused byan overdose and resulting in the death of pa-tients have been recorded. Extracts are used in

the treatment of opportunistic oralinfectionssuch as candidiasis in HIV-infected patients,and although they have potent antifungal ac-tivity, they should be used with care becausethey may have interactions with antiretroviralagents. The bark has been used as fish poison,and the latex as hunting poison for arrowheads. Leaf decoctions are applied to the eyesto treat ophthalmia. In Namibia the powderedoily wood, mixed with fat, has been rubbed intothe hair, and it has also been used as perfume.

Production and international tradeAstudyin 2003 in Maputo (Mozambique) showed thatthe wood of Spirostochys of neono is commonlyused by wood artists. The annual volume wasestimated at 360 in3. In general, it has beenreported that about 40 in3 of Spirostochys ofri-cong wood per month is available from sus-tainably managed sources

Properties The heartwood is brown to darkbrown with darker markings and streaks, anddistinctly demarcated from the whitish to paleyellow, 2.5-5 cm wide sapwood. The grain isstraight to slightly wavy, texture moderatelyfine to fine, and even. The wood has a beautifulbanded figure and a satin-like lustre, with anoily surface. It has a fragrant, spicy smell, re-sembling that of EastTndian sandalwood (Son-toIum o16um L. ), which can persist for manyyears

The wood is very heavy, with a density of 910-1090 kg/in3 at 12% moisture content, and hardIt air dries slowly. Boards of 2.5 cm thick takeabout 7 months to air dry and boards of 10 cmthick about one year. End checking is common,but the wood is usually not very liable to dis-

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tortion. The rates of shrinkage are low, fromgreen to oven dry 2.1-3.5% radial and 4.0-6.7%tangential. Once dry, the wood is moderatelystable in service.

At 12% moisture content, the modulus of rup-ture is 102-108 N/mm2, modulus of elasticity8600-9210 N/min2, compression parallel tograin 57-60 N/mm2, shear 16 N/mm2 andJanka side hardness 8940 N.

The wood is difficult to saw, particularly greenwood, but it is more easy to work. It rapidlyblunts saw teeth. The saw dustis gummy andsticks to the teeth. The wood near the centre of

the log may be extremely hard to saw. It planessmoothly and moulds well. Drilling and mortis-ing require considerable effort, and nailing isonly possible after pre-boring. Turning givesexcellent results. Sanding is difficult because ofthe oily wood surface. The wood is durable andresistant to fungi, termites and wood borers. Itis very resistant to preservatives. The sawdustis highly irritant to eyes and respiratory tracts.The wood is not wellsuited as firewood because

the smoke is irritant, causing headache andcomplaints of the respiratory organs.The latex is irritant and may cause innamma-tion and blistering of the skin. It is particularlyharmful when in contact with the eyes. It maycause severe purging and vomiting. It containsa complex mixture of diterpenes. Bark extractsshowed antibacterial activity and genotoxicityin tests. Experiments showed that powderedbark suppressed populations of the cowpeaweevil (Collosobruchus ingculotus) in storedcowpea seeds. The bark contains diterpenoids,triterpenoids, tannins, arithocyanins and sapo-rims. Several diterpenoids have been isolatedfrom the heartwood.

Description Deciduous, dioecious or monoe-CIOus shrub or small tree up to 15(-20) in tall,with white latex; bole branchless for up to 7 in,straight or bent, up to 50(-90) cm in diameter,more or less fluted near base; bark surfacerough and flaking in rectangular scales, darkgrey to dark brown or black, inner bark fibrous,dark red, exuding white latex; crown rounded,heavily branched; twigs glabrous, with lenti-cels. Leaves arranged spiralIy, simple; stipulestriangular-ovate, c. I min long, caducous; peti-o1e 0.5-Tel. 5) cm long, channelled above, with2 small glands at apex; blade ovate to elliptical,(1.5-)2-7 cm x (0.5-)I-3.5 cm, usually roundedat base, obtuse to rounded or slightly short-acuminate at apex, margins obtusely toothed,papery to thinly leathery, glabrous, pinnatelyveined with 6-14 pairs of lateral veins. Trillo-

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SPIROSTACHYS 599

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rescence an axillary spike I-2.5 cm long,densely flowered, with only male flowers orwith many male flowers and I-3 female now-ers at base; bracts arranged spiralIy, broadlyovate, c. 1.5 mm broad. Flowers unisexual,regular, sessile or nearly so; petals and diskabsent; male flowers with (2-)3 0bovate torounded sepals c. I mm long, stamens 3, fusedat base into a tube; female flowers with 5 tri.angular-ovate sepals c. I mm long, ovary supe-nor, c. I mm in diameter, 3-10bed and 3-celled,styles 3, c. 1.5 min long, fused at base, with redstigmas. Fruit a 3-10bed capsule c. I cm in di-ameter, smooth, becoming yellowish brown,falling apart into 3 parts, each part 2-valvedand I-seeded. Seeds ovoid to globose, c. 4 minlong, smooth, pale brown to yellowish brownwith darker streaks

Other botanical information Spirostochyscomprises 2 species and is closely related toEtcoecorio. Spitostochys o17icono closely re-sembles Etcoecorio stintt (Kuntze) Pax, whichis restricted to South Africa and differs in more

sharply toothed leaves without glands at peti-o1e apex.

600 TIMBERS2

Spirostochys uenentfero (Pax) Pax is a shrub orsmall tree up to 15 in tall, occurring in south-ern Somalia, Kenya and eastern Tanzania. Itswood is used for construction, door frames,ceiling beams, Implements, tool handles andcarvings, and also for charcoal production. Thewood is yellowish brown to pale brown, withbrown markings, medium-weight with a densi-ty of about 750 kg/ing at 12% moisture content,and quite durable. The bark and latex are usedas fish poison. Tests in mice showed jinmuno-suppressive activity of bark extracts. Spirosto-chys Denenifero is highly poisonous.

Anatomy Wood-anatomical description CAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;(10: vessels in radial multiples of 4 or morecommon); 13: simple perforation plates; 22:intervessel pits alternate; (23: shape of alter-nate pits polygonal); 26: intervessel pits medi-urn (7-10 prn); 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumina 50-100 ILm;48: 20-40 vessels per square minimetre; 49:40-100 vessels per square minimetre; 58: gumsand other deposits in heartwood vessels. Tra-cheids and fibres: (60: vascular/vasicentric tra-cheids present); 61: fibres with simple to ini-nutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled. Axialparenchyma: 76: axial parenchyma diffuse;(77axial parenchyma diffuse-in-aggregates); (86axial parenchyma in narrow bands or lines upto three cells wide); 92: four (3-4) cells per pa-renchyma strand; 93: eight (5-8) ceUs per pa-renchyma strand. Rays: 96: rays exclusivelyuniseriate; 104: all ray cells procumbent; (106:body ray cells procumbent with one row of up-right and/or square marginal cells); (109: rayswith procumbent, square and upright cellsmixed throughout the ray); 1/3: disjunctive rayparenchyma cell walls present; 1/6: z 12 raysper min. Mineral inclusions: 136: prismaticcrystals present; 137: prismatic crystals in up-right and/or square ray cells; 138: prismaticcrystals in procumbent ray cells(P. Mugabi, P. E. Gasson & E. A. Wheeler)

Growth and development Young treesmay have spiny branches. The trees grow veryslowly. They are usually deciduous, but some-times can be nearly evergreen, shedding theold leaves when young leaves develop. Insouthern Africa, trees usually start to flower inSeptember before new leaves develop, and

flowers may be present until January. Fruitsripen I-2 months after flowering.Although the tree is poisonous, several animalsfeed on it; birds such as guinea-fowl, francolinand doves eat the fruits, and large herbivoressuch as antelopes, elephants and rhinocerosyoung foliage. Squirrels, often nesting in thehollow hole, feed on the young leaves, bark andseeds.

Ecology Spirostochys @1nconudeciduous woodland and wooded grassland,often on termite mounts and stony slopes,sometimes in thickets, up to 1350 in altitudeThe largest trees are found near streams andseasonal watercourses. Spirostochys o17iconoprefers well-drained sandy or sandy loam soils.In southern Africa, it is often associated withGolophospermum mopone (Benth. ) J. LeonardandAcocio, Brochystegio and Combretum spp

Propagation and planting In Botswanafresh seeds showed a germination rate of 55-80% I-3 weeks after withoutsowing, pre-

treatment. They can be sown in containersfilled with river sand, with the seeds pushedinto the sand untillevel. They should be keptmoist. Seeds should be selected carefully be-cause they are often infested with moth larvae.

Management Trees can be coppiced, buttheaverage number of shoots per stump is small(0.4-I).

Diseases and pests The seeds may be in-fested with the larvae of a moth (Binporio me-10nobosis), and these may cause the seed tojump into the air for several centimetres

Handling after harvest The holes are oftenhollow, and larger dimensions of wood aretherefore difficult to obtain. Star shakes often

develop during log drying. In an Inventory inKwaZulu-Natal (South Africa), thelength of posts used in house building was 2-2.5 in and the mean diameter 7.5-9.5 cm

Genetic resources There are no indications

that Spirostochys of neono is currently threat-ened; it is both widespread and locally coin-mon. However, In many regions its wood isgreatly appreciated and sought for, and regula-tion of the yield is there needed to avoid toomuch pressure on the populations. A study in2007 in southern Tanzania showed that the

trees were harvested on a large scale below thelegally prescribed minimum hole diameter. InSouth Africa the bark is locally in high demandfor medicinal purposes, but it seems to be stillreadily available.

Prospects Spirostochys of neono producesone of the most beautiful African woods. In

occurs ln

mean