tions and as pulpwood is even possible, In viewof the current trends of uslrig bio-based andbiologically degradable materials Instead ofsynthetic ones. It is not clear to what extentand where exactly the tree occurs planted ornaturalized in tropical Africa, but it may havesome potential here for plantations. An Impor-tant advantage is its fast growth, as the woodcan be harvested at 7-8 years rotation in tim-her plantations

Major references Beentje & Smith, 2001;Farmer, 1972; Keating & Bolza, 1982; Viniers,1975b; Wiselius, 1998b

Other references Burki11, 1994; Chittenden& Palmer, 1990; CTFT, 1961b; Dahms, 1991;Lamprecht, 1989; Sallenave, 1964; Sallenave,1971; Villavelez & Meniad0, 1979; Webb at al. ,1984; Wiemann & Williamson, 1988.

Sources of illustration Wiselius, 1998bAuthors M. Brink

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

ately high: from green to oven dry about 4.2%radial and 8.0% tangential. Boards of 2.5 cmthick take about I month to dry to 30% ino1s-ture content. The wood is moderately stable inservice. At 12% moisture content, the modulusof rupture is 1/9 N/min2, compression parallelto grain 43 N/mm2, cleavage 16 N/mm andChalais-Meudon side hardness 2.7The wood is easy to saw and work with bothhand and machine tools. Sanding is sometimesneeded for good results in planing operationsThe wood is fairly easy to glue, paint and var-nish. It nails well, with good nail-holding ca-pacity. The peeling and slicing properties aregood. The moulding properties are satisfactoryand the wood is suitable for carving. The woodis often attacked by pinhole borers, but is inod-erately resistant to fungi, dry-wood borers,termites and marine borers. It is resistant toimpregnation by preservatives.

Botany Medium-sized tree up to 25 in tall;hole branchless for up to 16 in, but often bent,up to 80 cm in diameter, sometimes wLth but-tresses up to 1.5 in high; bark with numerouslenticels, with aromatic scent when cut; twigsslightly angular, densely powdery-hairyLeaves alternate, simple and entire; stipulesabsent; petiole I-2 cm long, grooved above;blade elliptical, 6-17 cm x 2-8 cm, curieate atbase, acuminate at apex, papery, glossy green,glabrous, pinnately veined with 5-8 pairs oflateral veins, with aromatic scent. Innores-cence an axillary or terminal cyme or panicleup to 6 cm long, densely short-hairy but gla-brescent, 3-20-flowered; peduncle I-3 cm longFlowers bisexual, regular, fragrant; pedice1 I-3min long; perianth lobes 6, elliptical, c. 3 minlong; stamens 9 in 3 whorls, arithers 4-celled,stamens of inner whorl with 2 glands at base,staininodes I-2, minute; ovary superior, ovoidto ellipsoid, I-1.5 min long, glabrous, I-celled,style c. I min long, stigma disk-shaped. Fruitan ovoid drupe-like berry up to 2 cm long, atbase enclosed in the cup-like enlarged recepta-cle I-1.5 cm long, I-seeded.Ocoteo cymoso trees grow slowlyThe estimated number of species in Ocoteoranges from 200 to 350, most of them in tropi-cal America. Mainland Africa has about 7 spe-cies, Madagascar about 35.In Madagascar several other Ocoteo spp. arevalued for their wood, which is quite similar tothat of Ocoteo cymoso, also known under thename 'varongy' and used for similar purposesThe most important of these are OcoteoIOUcherei(Danguy) Kosterm. , Ocoteo joeuis

OCOTEACYMOSA (Nees) Palackj'

Protologue Cat. PI. madagasc. 2: 9 (1907).Family LauraceaeSynonyms RODensoro topuk Bail1. (1885).Origin and geographic distribution Ocoteo

cymoso occurs throughout eastern MadagascarUses The wood of Ocoteo cymoso and several

other Ocoteo spp. , known as 'varongy', is inhigh demand for furniture, cabinet making,boat construction and mortars, but is also val-ued for construction, Joinery, vehicle bodies,interior trim and pattern making. It is suitablefor mine props, ship building, toys, novelties,

andboxes, crates, carvings, turnery, veneerplywood. It is less suitable for flooring becauseit is too soft

The leaves, bark and fruits are aromatic; theleaves and fruits are used as condiment, thebark is added to locally prepared alcoholicdrinks

Properties The heartwood is pale brownand usually indistinctly demarcated from thesapwood. The grain is usually interlocked, tex-ture medium. Radial surfaces often show amarked stripe or ribbon figure. Freshly sawnwood has an unpleasantsmellThe wood is moderately lightweight, with adensity of about 560 kg/ms at 12% moisturecontent. It air dries well with little degrade,but best results are obtained with quarter-sawn logs. The rates of shrinkage are moder-

OCOTEA 401

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402 TIMBERSl

Kosterm. , Ocoteo inocrocorpo Kosterm. , Ocoteoplotydisco Kosterm. , Ocoteo rocemoso (Dan-guy) Kosterm. , Ocoteo thouuenotii(Danguy)Kosterm. and Ocoteo trichophlebio BakerThese all occur in eastern Madagascar buthave smaller distribution areas than Ocoteocymoso, except Ocoteu joeuis and Ocoteo ro-cemoso, which occur throughout eastern Mada-gascarlike Ocoteo cymosoOcoteo coinoriensis Kosterm. is endemic to theComoros. Its wood is also similar to that ofOcoteo cymoso and used for similar purposes. Abark decoction is used to treat headache, un-nary disorders and stomach complaints. Ocoteocoinoriensis showed in-vitro antimalarial activ-ityOcote0 o6tusoto (Nees) Kosterm. is a shrub orsmalltree up to 15 in tall, endemic to Reunionand Mauritius; in Reunion it occurs locallyfrequently, but in Mauritius it is rare. Thewood, known as 'bois de canelle', is used forfurniture. Oil extracted from the fruits is usedfor lighting

Ecology Ocoteo cymoso occurs in forest andcoastal dunes, from sea-level to 1250(-1900) inaltitude. It is locally common. It has been clas-SIfied as a semi-hellophyte, but trials showedthat it is extremely shade tolerant.

Management The growing stock of Ocoteocymoso has been estimated at 42 million m'in1990 and 40 million in' in 2000. Although thewood is suitable for peeling, it is often difficultto obtain logs that are large enough; moreover,larger-diameter logs are commonly hollow.

Genetic resources and breeding Ocoteocymoso is widespread in eastern Madagascarand is locally common. There are no indicationsthat it is in Immediate danger of genetic ero-SIon, but the popularity of its wood and theongoing forest fragmentation are reasons for

Prospects Although Ocoteo cymoso belongsto the more highly esteemed timber species ofMadagascar, very little research has been doneOn It

Major references Bolteau, Bolteau & A1-10rge-Bolteau, 1999; Bolza & Keating, 1972;Gu6neau, Bedel & Thie1, 1970-1975; Koster-mans, 1950; Sallenave, 1955.

Other references Decary, 1946; de Gouve-nain, 2001; FAO, 2008; Gunb-Fakim &Brendler, 2004; Kostermans, 1982; Parant,Chichignoud & Rakotova0, 1985; Sarrailh ata1. , 2007; Schatz, 2001; Takahashi, 1978

Authors R. H. M. J. Lemmens

OCOTEAKENYENSIS (Chiov. ) Robyns &R. Wilczek

Protologue Bull. Jard. Bot. Etat 20: 212(1950).

Family LauraceaeSynonyms Ocoteo uiridis Kosterm. (1938).Vernacular names Bastard stinkwood,

northern stinkwood, Transvaalstinkwood (En)Origin and geographic distribution Ocoteo

henyensis occurs from southern Sudan andsouthern Ethiopia, through eastern DR Congo,Rwanda, Kenya, Uganda and Tanzania, toZimbabwe, Mozambique, Swaziland and east-ern SouthAfrica

Uses The wood is used for flooring, panel-ITng, furniture and carving. It is suitable forlight construction, joinery, interior trim, vehi-cle bodies, handles, ladders, sporting goods,toys, novelties, turnery, veneer and plywood. Itis also used as firewood and for charcoal pro-duction

A bark decoction is used in traditional medi-cine as an antitussive. The bark is chewed totreat diarrhoea

Properties The heartwood is pale goldenbrown to dark brown, with blackish markings;it is lustrous. The grain is often wavy, texturemoderately fine. The wood is moderatelyheavy, with a density of about 750 kg/ina at12% moisture content. It air dries slowly andshould be dried with care to avoid serious de-

grade. Drying of thick boards may be problem-atIC. The rates of shrinkage are moderatelyhighThe wood saws and works satisfactory withboth hand and machine tools, but picking up ofgrain at the surfaces may occur; sharp cutting

concern.

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edges are recommended to obtain surfaces witha nice finish. The gluing properties are satis-factory. The wood slices and peels well. It isfairly durable and moderately resistant to ter-mite and marine borer attacks, but susceptibleto Lyetus attack. The heartwood is moderatelyresistant to impregnation by preservatives.

Description Evergreen, small to medium-sized tree up to 30(-40) in tall; hole usuallystraight, up to 100(-150) cm in diameter; barksurface greyish brown to reddish brown, rough,scaling off in irregular flakes, with conspicu-

scattered lenticels, inner bark greenishOUS,

white to yellowish brown or orange, turningdark grey to dark brown upon exposure; crownrounded, much-branched, dark green; brancheswith distinct leaf scars, twlgs angular, short-hairy to nearly glabrous. Leaves alternate,simple; stipules absent; petiole 0.5-1.5(-2) cmlong; blade ovate to elliptical, (4.5-)6.5-22 cmx 2-10 cm, curieate to rounded at base, acute toacuminate at apex, margins entire or wavy,leathery, glossy dark green above, paler below,often reddish at margins, red when young, gla-brous, pinnately veined with 8-10 pairs of Iat-eral veins, with aromatic scent. Inflorescencean axillary or terminal cyme or panicle up to 7

cm long, densely greyish short-hairy, few-flowered; peduncle 05-I(-2.5) cm long. Flow-ers bisexual or unisexual, regular, fragrant;pedice1 I-5 min long, thickening in fruit; perl-arith lobes 6, ovate-erriptical to orbicular orobovate, 2-3.5 min long, greenish yellow orwhitish yellow, short-hairy; stamens (7-)9 in 3whorls, arithers 4-celled, stamens of innerwhorl with 2 glands at base, staininodes form-ing a fourth whorl around the ovary or absent;ovary superior, ovoid to globose, c. I min long,glabrous, I-celled, style I-1.5 mm long, stigmadisk-shaped; male flowers with reduced ovary,female flowers with reduced stamens. Fruit anoblong-ellipsoid to ovoid-ellipsoid drupe-likeberry 1.5-2.5 cm long, olive-green, at base en-closed in the cup-like enlarged receptacle c. Icm long, I-seeded

Other botanical information The estima-ted number of species in Ocoteo ranges from200 to 350 species, most of them in tropicalAmerica. Mainland Africa has about 7 species,Madagascar about 35Ocoteo bulloto (Burch. ) Baill. is endemic toeastern and southern South Africa. It stronglyresembles Ocoteo kenyensis, but differs in Ltsleaves having blisters in the axils of the lowerlateral veins. Its yellowish grey to dark brown,often nicely figured and lustrous wood, knownas 'stinkwood', is already for centuries in highdemand, especially for furniture, small orna-merits and turnery, and is highly prized. Meth-ods of sustainable production of timber havebeen developed recently, which, however, allowonly very little production. The wood is alsoused as firewood. The bark is Important in tra-ditional medicine, particularly to treat head-ache, urinary and nervous disorders, stomachcomplaints and diarrhoea. Ocoteo bulloto isplanted as ornamental tree in gardens andparks

Anatomy Wood-anatomical description OAWAhardwood codes)Growth rings: a growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 27: intervessel pits large (Z 10 prn); 31:vessel-ray pits with much reduced borders toapparently simple: pits rounded or angular;(35: vessel-ray pits restricted to marginalrows); 41: mean tangential diameter of vesselInmina 50-100 prn; (42: mean tangential di-ameter of vessellumina 100-200 prn); 47: 5-20vessels per square minimetre; 56: tyloses coin-mon. Tracheids and fibres: 61: fibres with SIm-

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PIe to minutely bordered pits; 65: septate fibrespresent; 69: fibres thin- to thick-walled. Axial

parenchyma: 78: axial parenchyma scantyparatrachea1; 92: four (3-4) 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. Secretory elements and cambial van-ants: 124: oil and/or mucilage cells associatedwith rayparenchyma.(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development In general, thetree grows fairly rapidly. The fruits are eatenby birds, which may serve as seed dispersers

Ecology Ocoteo henyensis occurs in ever-green rainforest at 1100-2400(-2600) in alti-tude. The mean annual rainfall in the area ofdistribution ranges from 1500 mm to 2200 mmIn the mountains of southern Sudan it occurstogether with Podocorpus andAlbiaio spp. , andin Kenya with Podocorpus andMocorongo spp

Propagation and planting Fresh seedshould be used for sowing. Seed is sensitive todesiccation, but can be stored for a short periodin moist saw dust.

Propagation by root suckers is easy; these areoften produced abundantly. Under naturalconditions, regeneration of Ocoteo kenyensis byseed in Ethiopia was found to be satisfactory

Management Ocoteu kenyensis trees can bemanaged by coppicing.

Diseases and pests Fruits are often heavilyattacked by insects

Genetic resources In many regions Ocoteohenyensis has been heavily exploited for itsvaluable timber. In southern Africa it shows avery scattered distribution pattern, with oftenonly small groups of trees or individual treesgeographicalIy isolated from other stands. It iscriticalIy endangered in Zimbabwe. Ocoteokenyensis Is included in the TUGN Red list asvulnerable

Prospects Although Ocoteo kenyensis pro-vides valuable timber and has been over-exploited, very little research has been done onits wood properties, growth rates and propaga-tion methods. The findings of such research areneeded to be able to develop methods of sus-tainable exploitation. This would probably re-SUIt in very low production levels, but the ex-ample of Ocoteo buildtu in South Africa showsthat this might still be a valuable option forsuch high-quality timbers

Major references Bekele-Tesemma, 2007;Bolza & Keating, 1972; Diniz, 1997; Prtis,

1992; Maundu & Tengnas, 2005; Palmer &Pitman, 1972-1974; Sommerlatte & Sommer-Iatte, 1990; Strahm, 1998

Other references Beentje, 1994; Bekele-Tesemma, Birnie & Tengnas, 1993; CoatesPalgrave, 1983; Dale & Greenway, 1961; Friis,2000; Hamillet a1. , 2000; InsideWood, undated;Robyns & Wilczek, 1951; Troupin, 1982; vanWyk & Gencke, 2000; Verdcourt, 1996; Wim-bush, 1957

Sources of illustration Hutchinson &Moss, 1930; Maundu & Tengn6. s, 2005; Palmer& Pitman, 1972-1974

Authors F. S. Mairura & R. H. M. J. Lemmens

OCOTEAUSAMBARENSIS Engl.

Protologue Abh. Konigl. Preuss. Akad. WISSBerlin 1894, I: 51, 54 (1894)

Family LauraceaeVernacular names East African camphor

wood (En). Camphrier (Fr). Mkulo, inukongo(Sw)

Origin and geographic distribution Ocoteousumborensis occurs in eastern DR Congo,Rwanda, Kenya, Uganda, Tanzania, northernMalawi and northern Zambia. It is planted ona smallscale in Kenya and Tanzania

Uses The wood, often traded as 'camphor' isvalued for Joinery, panelling, poles for building,doors, window frames, shutters, furniture,cabinet work, vehicle bodies, sliced veneer andplywood. It is used for flooring of local housesand for Implements. The wood is suitable forconstruction, ship building, boxes, crates, vats,matches and pulpwood. It is less suitable fordraining boards and kitchen utensils because

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of the camphor-like smell. It is also used asfirewood and for charcoal productionBark and roots are used in traditional medi-cine. The pounded bark of roots, hole andbranches is applied to swellings, boils andwounds. A bark decoction Is given to treatwhooping cough and measles. Bark powder istaken against stomach-ache. Roots steeped inwater are taken to treat malaria and back

Ocoteo usQinborensis is occasionallypain

planted as an ornamental shade tree, but itscrown is too dense to be useful for agroforestrypurposes

Production and international trade Ocoteuusumborensis is traded internationally in 11m-ited amounts, but there are no statistics onproduction and trade of the timber. In2000-2001 in Kenya it was the most highlypriced timber species. Exploitation of the barkfor medicinal purposes Is considerable, butthere is no information on amounts.

Properties The heartwood is pale yellowishbrown when freshly cut, darkening to deepbrown upon exposure, and sometimes lustrous;it is not distinctly demarcated from the slightlypaler sapwood. The grain Is usually inter-locked, texture even and moderately fine tofine. Quarter-sawn surfaces show a stripe orribbon figure. The wood has a distinct cam-phor-like sineUThe wood is medium-weight, with a density of450-640 kg/in3 at 12% moisture content. It airdries slowly with little degrade, although se-vere case-hardening may occur. Drying of thickboards may be problematic. Boards 2.5 cmthick take about 3 months to air dry, boards 5cm thick about 6 months. The rates of shrink-

age are moderately low, from green to oven dry2.5-4.1% radial and 5.0-6.7% tangential. Oncedry, the wood is quite stable in serviceAt 12% moisture content, the modulus of rup-ture is 75-92 N/mm2, modulus of elasticity9900-11,100 N/mm2, compression parallel to

37-53 N/min2, shear 11-15 N/min2,graln

cleavage about 11 N/mm, Janka side hardness2840-4310 N and Janka end hardness5070-5120 N

The wood is easy to saw, and it works satisfac-tonly with both hand and machine tools; theblunting effect on cutting edges is small. A 15-20' cutting angle is recommended in sawingand planing to avoid picking up of interlockedgrain. In nailing the wood has a tendency tosplit at the edges or ends, but the nail-holdingpower is satisfactory. Gluing and staining donot give problems, but the use of a filler Is re-

quired for optimum polishing results. It israrely peeled because logs a rarely sound at thecentre. The steam bending properties are inod-erate. The wood is moderately durable to dura-ble. It is very resistant to decay but can be sus-ceptible to termite and marine borer attacks,and the sapwood to powder-post beetle attackThe heartwood is very resistant to Impregna-tion by preservatives, the sapwood moderatelyresistant

In in-vitro tests, methanol extracts of the rootbark showed pronounced cytotoxic actLvityagainst several human cancer celllines at 100pg/inI. Bark extracts showed antifungal activ-ity against Clodosporium cueumerinum. Thisactivity was linked to the presence of the 11g-nan (+)-piperito1 and an essential oil, in whichnerolidol, or-terpineol, orcopaen-11-o1 and anunidentified sesquiterpene alcohol were themain active compounds.

Description Evergreen, smallto fairly largetree up to 35(-45) in tall; hole usually straight,often slightly fluted, branchless for up to15(-20) in, up to 200(-300) cm in diameter,with small buttresses at base; bark surfacegreyish brown to reddish brown, fissured, scal-ing off in round or square flakes, inner bark

OGOTBA 405

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406 TIMBERSl

white to pale pink, with sweet, camphor-likescent; crown dense, spreading; twigs slender,angular, hairy. Leaves alternate to opposite,simple and entire; stipules absent; petiole

long; blade ovate to elliptical,0.5-2 cm

(2.5-)4-17 cm x (2-)2.5-9 cm, curieate torounded or truncate at base, usually acuminateat apex, thinly leathery, glossy dark greenabove, silvery white below, slightly hairy atleast on veins, pinnately veined with 6-8 pairsof lateral veins, with camphor smell. Innores-cence an axillary panicle up to 13 cm long,densely yellowish brown hairy; peduncle 2-8cm long. Flowers bisexual or unisexual, regu-Iar; pedice1 I-4 min long; perianth lobes 6,ovate-elliptical, 2-3 mm long, greenish yellowor whitish yellow, short-hairy; stamens 9 in 3whorls, arithers 4-celled, stamens of innerwhorl with 2 glands at base, staininodes form-ing a fourth whorl around the ovary; ovarysuperior, ovoid, c. I min long, glabrous, I-celled, style c. I min long, stigma disk-shaped;male flowers with reduced ovary, female flow-ers with reduced stamens. Fruit an ellipsoid tonearly globose drupe-like berry c. I cm long,green to orange-brown, at base enclosed in thecup-like enlarged receptacle c. 3 min long, I-seeded.

Other botanical information The esti-

mated number of species in Ocoteo ranges from200 to 350 species, most of them in tropicalAmerica. Mainland Africa has about 7 species,Madagascar about 35

A11atomy Wood-anatointcal description CAWAhardwood codes):Growth rings: a 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); (16: scalariform perfora-tion plates with 10-20 bars); 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 27: intervessel pits large (Z 10 prn); 32vessel-ray pits with much reduced borders toapparently simple: pits horizontal(scalariform,gash-like) to vertical(palisade); 42: mean tan-gential diameter of vessellumina 100-200 pin;47: 5-20 vessels per square minimetre; 56tyloses common. Tracheids and fibres: 61: fi-bres with simple to minutely bordered pits; 66:non-septate fibres present; 69: fibres thin- tothick-walled. Axial parenchyma: 78: axial pa-renchyma scanty paratrachea1; 92: four (3-4)cells per parenchyma strand. Rays: 97: raywidth I-3 cells; 106: body ray cells procumbentwith one row of upright and/or square marginal

cells; (107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells); 1/5: 4-12 rays per min. Secretoryelements and cambial variants: 124: oil and/or

mucilage cells associated with ray parenchyma;(125: oil and/or mucilage cells associated withaxial parenchyma)(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development Records of growthrates of Ocoteo usQinborensis are contradictory.Growth rates of up to 2 in/year have been re-corded for young trees, but this seems to beexceptional. In a 75-year-old plantation at 2450in altitude in Kenya, trees were 15-29 in tall,with a hole diameter of 19-51 cm. The meanannual diameter increment was 6.2 mm until

18 years after planting, but thereafter de-creased gradually to 4.4 mm at 75 years old. Itwas suggested that the initialspacing (1.5 in x1.5 in) was too close to enable good growth. InTanzania 49-year-old trees were 15-24.5 intall, with a mean hole length of 10 in and meanhole diameter of 40-49 cm. It was recorded that

90% of the trees showed heart rot, rangingfrom 4-24% of the log volume, but in anotherstudy 60% of the trees was recorded to be freeof heart rot.

Trees may produce fruits in large amounts, butusually only once in 3-4 years, so-called mastyears. They often develop root suckers, butthese are often eaten by large animals such aselephants. Regeneration by suckering and cop-picing is high after clear felling.

Ecology Ocoteo usQinborensis occurs in rain-forest at 900-2600(-3000) in altitude. Themean annual rainfallin the area of distributionranges from (1200-)1600 min to 2450 min, withoften 2-3 dry months, and a mean annual tern-perature of (12-)16-20G26)'C. Ocoteo usQinbo-rensis may be dominant in mountain forest. Itprefers deep, fertile soils with good drainage

Propagation and planting The 1000-seedweight is about 150 g. Fresh seed should beused for sowing. The germination rate is oftenlow, up to 45%. Seeds are often heavily at-tacked by insects. They usually start germinat-Ing in 30-45 days, but germination may takeup to 90 days. Seeds should be cleaned fromthe surrounding pulp before sowing by rubbingin water. Seed Is sensitive to desiccation, butcan be stored for a short period in moist sawdust.

Propagation by root suckers Is easy; these areoften produced abundantly. Under naturalconditions, Ocoteo t'somborensis regeneratesmainly by suckers because undamaged seed is

.

uncommon. After natural breakdown of an oldtree, the gap is first filled by fast-growing PIO-neer species, in the shade of which the Ocoteousermburensis suckers can establish, and afterdeath of the pioneer species they can developInto new trees.

Management Ocoteo usumborensisis mainlyharvested from naturalstands, and the extentof plantations is very limited and confined toKenya and Tanzania. Large-scale logging leadsto secondary forest types in which Ocoteousumborensis does not play a significant rolebecause o11ack of regeneration.In plantations a heavy first thinning Is recoin-mended 15-20 years after planting, reducingthe stock to about 700 treesA1a, with subse-

quent thinnings at intervals of 7-10 years.Rotation cycles of 60-70 years have been prac-tised in Tanzania, butthese may be reduced to50 years with proper thinning regimes to fi-nany 220 treesA1a. Trees can be managed bycoppicing, to which they respond well at anyage

Diseases and pests Fruits are often heavilyattacked by insects. Standing older trees oftenshow heart rot caused by fungi such as Guno-dermo OPPlonotum and Fomes spp. The bark ofyoung trees is stripped by squirrels and treehyraxes and the leaves are browsed by ele-phants.

Handling after harvest The logs of largetrees often have rotten cores, and holes aresometimes ill-shaped or have much twistedgrain. This makes selection of logs before saw-ing for different purposes important

Genetic resources In Kenya and TanzanlaOcoteo usumborensis was dominant in manymountain forests, the so-called 'ocotea forest,but it occurs more sparsely in Uganda. How-ever, it has become rare throughout East of-rica due to over-exploitation and is officiallyprotected in Kenya and Tanzania. Illegal log-ging is still widespread, and conservatlonmeasures seem to be necessary.

Prospects Ocoteo usumboreirsisseemsto havegood prospects as a plantation timber tree,providing wood of exceUent quality. Although itis considered valuable and has been over-

exploited, very little research has been done onits growth rates and propagation methods. Thecommon use of the bark in local medicine war-rants more research on the pharmacologicalactivities and on sustainable collection of thebark

Major references Bolza & Keating, 1972;Diniz, 1997; Farmer, 1972; Hines & ECkman,

1993b; rugom0, 1987; Maundu & Tengnas,2005; Mbuya at a1. , 1994; Takahashi, 1978;Tanzania Forest Division, 1965; World Agro-forestry Centre, undated

Other references Beentje, 1994; Bryce,1967; Bussmann, 2001; CAB International, 2005;Chudnoff, 1980; Dale & Greenway, 1961; Dick,1969; F1euret, 1980; InsideWood, undated;minuhabwa, Nshimo & dewitte, 2000; Katende,BITnie & Tengn6. s, 1995; minariy0, 1971; Kok-war0, 1993; Mugasha, 1978b; Mugasha, 1980;Ndangalasi, Bitariho & Dovie, 2007; Neu-winger, 2000; Terreaux at a1. , 1994; Troupm,1982; Verdcourt, 1996; Wimbush, 1957

Sources of illustration Diniz, 1997; Katen-de, BITnie & Tengnas, 1995

Authors J. M. Okeyo

OCTOLOBUS 407

OCTOLOBUSSPECTABILISWelw

Protologue Trans. Linn. SOC. London 27: 18(1869)

Family Sterculiaceae (APG: Malvaceae)Synonyms Octolobus origustotus Hutch

(1937)Origin and geographic distribution Octolo-

bus spectobilis is distributed from Sierra Leoneto DR Congo and Angola; also in Tanzania

Uses In Nigeria the sterns of Octolobus spec-tobi!is are made into spear shafts. The seedsare said to be edible. A maceration of the rootbark is taken against sexual asthenia.

Properties The wood of Oct0!o6us specterbi-lis is yellowish white, with brownish necrosisin the centre. The texture is fine. The wood isheavy and rather hard

Botany Shrub or smalltree up to LOG15) intall; hole up to 15 cm in diameter; branchletsdensely stellate-hairy but glabrescent. Leavesalternate, simple and entire; stipules attenu-ate, 4-12 min long, usually less than I minwide; petiole up to 7 cm long, pubescent, rustybrown; blade obovate-elhptical to obovate-oblong, 6-24(-30) cm x 1.5-8(-11) cm, basecurieate to rounded, apex acuminate, papery,light brown, with (5-)7-11 pairs of lateralveins. Flowers solitary, male or bisexual, yel-low, sessile, surrounded at base by an Involucreof 7-20 ovate bracts; calyx 8-lobed, 1.5-3.5 cmlong, with tube c. 1.5 cm long and oblong-lanceolate lobes I-1.5 cm long, with crisp mar-ginal fringes, hairy outside; corona absent;male flowers with an androphore 5-7 min long,carrying a whorl of numerous connate stamensc. 4 mm long; bisexual flowers with numerous

.

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carpels in 2-4 Imbricate rows. Fruit consistingof 10-65 follicles; follicles almost globose, up to5 cm x 3.5 cm, with stipe 0.5-1.5 cm long andpointed beak 2-5 min long, scabrous, denselyand shortly hairy, reddish brown, 2-7-seededSeeds compressed, plano-convex, diameter c. 15mm, black, shiny, surrounded by mucilageSeedling with epigealgerminationOctolobus comprises 3-4 species. It is closelyrelated to Co!0, but differs in its 8-lobed calyxand numerous carpels. Octolobus specter bins ishighly variableIn Ghana Octolobus spectobilis flowers inJanuary-February; fruiting is in February

Ecology In West Africa Octolobus spectobi-lis occurs in the undergrowth of forest, scat-tered but common in dry forest types. In Cen-tral Africa its habitats are dense humid semi-deciduous forest and gallery forest. In Tanza-nia Octolobus spectobilis is found in forest atround 1000 in altitude

Genetic resources and breeding In viewof its wide distribution and common occur-rence, e. g. in Nigeria, Octolobus spectobilis isnotthreatened by genetic erosion.

Prospects Little is known on the timber prop-erties of Oct0!o6us spectobilis, but its small sizeand hole diameter are serious limitations to its

wood.

A flour is prepared from the seeds that is usedas a condiment in sauces. Fat extracted from

the seeds can be eaten raw and is also madeinto various medicines. A mixture of the

pounded leaves with palm oilis rubbed on thehead against head lice, or the head is washedwith a maceration of the bark for the same

purpose. A decoction of the bark is takenagainst stomach-ache, bronchitis and lungcomplaints. The ground bark mixed with palmo111s applied as a salve to treat psoriasis. Theflowers attract honeybees.

Properties The heartwood is pale yellowand lustrous; it is not demarcated from thesapwood. The grain is fairly regular, sometimesirregular, texture medium to coarse; quarter-sawn surfaces occasionally have a inottled fig-ure. The wood is lightweight with a density ofabout 380 kg/in3 at 12% moisture content.Shrinkage rates are 3.4% radial and 5.7% tan-gential from green to oven dry. The wood sea-sons rapidly with little splitting. The wood issoft and brittle. At 12% moisture content, themodulus of rupture is about 55 N/mm2modulus of elasticity 5690 N/mm2 compressionparallel to grain 28 N/mm2 and shear 2.2N/min2

Seasoned wood saws easily and planes wellScrews and nails are applied easily, with littletendency for the wood to split, but it does nothold nails well. The wood peels and slices eas-11y, but as the logs are deeply fluted peeling isnotrecommended

The wood has a low durability, being liable tofungal attack and susceptible to termites, POW-der-post beetles (Lyetus) and dry-wood borersThe heartwood and sapwood are fairly perme-able for preservatives and can be treated usingpressure methods.

The ultimate fibres are 1.1 min long and 27 pinwide, with a lumen diameter of 19 pin and acell wanthickness of 7 pin. The wood can bepulped satisfactorily using the kraft or SUIphiteprocesses

Cytotoxic quassinoids have been isolated fromthe bark

Botany Medium-sized to fairly large tree upto 40 in tall; hole branchless for up to 24 in, upto 200 cm in diameter, usually straight, deeplyfluted over Its entire height; outer bark grey togreyish green, finely longitudinalIy cracked,inner bark thick, fibrous, pale yellow; crownspherical; branches thick, cylindrical, blackish.Leaves arranged spiralIy, Imparipinnatelycompound with 4-6 pairs of leaflets, glabrous;

use

Major references Aubr6ville, 1959b;Burki11, 2000; Cheek & Frimodt-Muller, 1998;Germain & Barnps, 1963; ITvine, 1961

Other references Hall6, 1961; Keay, 1958e;Marshall at a1. , 2001; Neuwinger, 2000; Nor-mand, 1955; Wilkie at a1. , 2006

Authors M. Brink

ODYENDYEAGABONENSIS (Pierre) Engl

Protologue Nat. Pfianzenfam. 300: 215 (1896)Family SimaroubaceaeSynonyms Quossiogobonensis Pierre (1896).Origin and geographic distribution Odyen-

dyeo gobonensis is distributed in Cameroon,Equatorial Guinea, Gabon and Congo

Uses The wood of Odyendyeo gobonensisOrade name: onzan, onzang, inbanko) is usedfor making musical Instruments, mallets tobeat bark for house construction, spoons, andbells for hunting dogs. In general it is consid-ered suitable for ship and boat building, turni-ture and cabinet work, frame moulding, lightboxes and crates, Interior trim, matches, toysand novelties, veneer and plywood, hardboardand particle board, wood-wool, and as pulp-

.

stipules absent; rachis 30-50 cm long, chan-nelled at base; petiolules c. 3 mm long, chan-nelled;leaflets oblong, up to 12 cm x 5 cm, basecurieate or rounded, apex truncate or slightlynotched, leathery, pinnately veined with c. 10pairs of hardly visible lateral veins almost per-pendicular to inIdrib. Inflorescence a terminalpanicle shorter than the leaves, branches gla-brous. Flowers unisexual or bisexual, regular,4(-5)-meTous; pedicel c. 3 min long; calyx cu-pule-shaped, with short obtuse lobes; petals c5 min long, glabrous outside, pubescent inside;stamens 8(-Ion, c. 8 mm long; ovary superior,consisting of 4 fused carpels. Frult an obovoiddrupe up to 7 cm long, grooved at one side,becoming red when ripe; pyrene with thickhard wallhaving a longitudinal crest, I'Seeded.Kernelthick, brownIn Gabon Odyendyeo gobonensis flowers inSeptember. The fruits are eaten by porcupines,which reportedIy makes the meat of these anI-inals bitter

Odyendyeo comprises only ISPeciesEcology Odyendyeogobonensis occurs main-

Iy in rainforest, in primary as well as in secon-dary forest, often on sandy soil. Although itnowhere occurs abundantly, It is locally coin-

Management To prepare flour, the seeds areboiled for 2-3 hours, placed in running waterfor one week to eliminate bitterness, kept in asmokehouse to dry for one month, and crushedto obtain the flour

Genetic resources and breeding AlthoughOdyendyeo gobonensis is not mentioned in theIUCN red list of threatened species, it has arestricted distribution and does not occurabundantly. Therefore, the species may becomeendangered as a result of ongoing logging ac-tivities.

Prospects The wood of Odyendyeo gobonen-sis is soft and not durable, and the logs are toomuch fluted to be useful for rotary peelingTherefore, its importance is unlikely to in-crease beyond being used locally. Its applica-tions in food and medicine deserve more atten-tion.

Major references Aubr6ville, 1962c; Bolza& Keating, 1972; Raponda-Walker & Sillans,1961; Takahashi, 1978

Other references CTFT, undated; de Saint-Aubin, 1963; Neuwinger, 2000; Nooteboom,1962; Nziengui, 2001; Petroff, Doat & Tissot,1967; Waterman & Ampof0, 1984; Wilks &TSSemb6, 2000

Authors M. Brink

OMPllALOCARPUMELATUMMiers

Protologue Trans. Linn. SOC. London, Botser. 2, I: 16 (1875).

Family SapotaceaeChromosome number2n=26

Synonyms Qinpholocorpum onorentrumPierre ex Eng1. (1904)

Originand geographic distribution Ompho-IOCorpum elotum occurs from Sierra Leone eastto the Central African Republic and south toGabon and Congo

Uses The wood is used for planks, imple-merits such as mortars and bowls, handles,seats and drums. It is also used for dugoutcanoes. The latex has been used as an adulter-ant of rubber. In Nigeria a bark decoction Isused in traditional medicine to treat constipa-

tion. In Cameroon the Bata Pygmies use a barkdecoction together with fruits of GOPsic!, in on-ituum L. and Solanum origuiui Lain. to treatmalaria, whereas a decoction or maceration ofthe bark is taken in case of lactation failure,and a decoction of the young leaves for thetreatment of cough. In C6te d'Ivoire a mixtureof the seeds and crushed bark, diluted in palmwine, is used as a purgative in case of poison-ing and to treat scrotal elephantiasis. In SierraLeone the seeds are used in the treatment ofyaws. The seeds are used for decorative pur-poses, e. g. for necklaces.

Properties The heartwood is pale reddishbrown or pale brown with a pinkish tinge, andindistinctly demarcated from the whitish sap-wood. The grain is straight or srightly wavy,texture is moderately fine. Fresh wood has afoetid smell, but dry wood is odourless. In atest in Liberia, the wood had a density of 640kg/in3 at 12% moisture content, Janka sidehardness was 5840 N and Janka end hardness6800 N. The wood is fairly easy to work andfinishes smoothly. It is only moderately dura-ble

In tests the bark showed high in-vitroarithelmintic activity against Hoemonchuscontortus larvae. The seeds contain alkaloidsand saponins

Botany Evergreen medium-sized tree up to30 in tall, with latex; hole straight and cylin-drical but often slightly fluted, up to 80(-180)cm in diameter, without buttresses; bark sur-face scaly, brown, with lenticels in longitudinalrows, inner bark reddish brown, finely fibrous;crown compact; young branches glabrous.Leaves arranged spiralIy, clustered at ends ofbranches, simple and entire; stipules absent;

OMPHALOCARPUM 409

mon.

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petiole short, up to I cm long; blade oblong-obovate to oblanceolate, 12-25 cm x 4-8 cm,curieate at base, rounded to shortly acuminateat apex, glabrous, pinnately veined with 10-15pairs of lateral veins. Inflorescence a fascicleon hole or older branches. Flowers bisexual orunisexual, regular; pedicel c. I cm long; sepals5, free, thick; corolla up to 2 cm long, white,with c. 6 min long tube and 5(-7) ellipticallobes hairy at margins; stamens inserted atapex of corolla tube, in bundles of 5-6 oppositeeach corolla lobe, about as long as lobes, infemale flowers rudimentary, bundles alternat-ing with large, petalold, fringed staininodes;ovary superior, glabrous, up to 30-celled, stylecylindrical, included. Fruit a large depressedglobose berry up to 15(-20) cm in diameter,with greyish brown, woody wall and whitishpulp, many-seeded. Seeds flattened ellipsoid, c4 cm x 2.5 cm, black, with linear scar. Seedlingwith epigeal germination; hypocoty1 6-10 cmlong, epicotyl c. 2 cm long; cotyledons leafy,broadly elliptical, c. 6.5 cm x 4.5 cm.Ompholocorpum comprises about 7 species,and is most closely related to the genusTridesmostemon from Central Africa. It ischaracterized by the large fruits attached tothe hole. The wood of Ompholocorpum ohioA. Chev. , occurring from Sierra Leone toGhana, is used in Ghana for house buildingand carpentry, whereas its bark is used to treatstomach-ache and rheumatism; the seeds haveornamental value. Ompho!ocorpum ohio haslarger leaves than Ompholocorpum elotum (upto 45 cm x 18 cm). In Central Africa the woodof several other Ompholocorpum spp. and thatof Qinpholocu, pum elotttm is probably usedindiscriminately. Ompholocorpum @10tum canbe found flowering and fruiting throughout theyear, but in C6te d'TVoire it flowers mainly inJune-July. The fruits take about one year toripen. They are eaten by elephants, which arethe only animals able to crack open the hard,thick shell. The elephants swallow the pulptogether with the seeds, which are deposited inthe dung and thus dispersed. It seems that thepresence of elephants is needed for successfulseed germination under natural conditions,because the seeds of rotten fruits on the groundare destroyed by insects and seedlings are notfound under trees in regions where elephantshave disappeared

Ecology Ompholocorpt, in e!otum occurs scat-tered in evergreen forest, often in humid localI-ties

Management The 1000-seed weightis about

2.5 kg. Seeds that have passed through an ele-phant's gut germinate after about 2 weeks;seeds extracted from the fruits take 3-9 weeks

The seedlings are probably shade tolerantGenetic resources and breeding Ompho-

locumum elotum is fairly widespread and 10-cally not uncommon, and consequently it doesnot seem to be endangered at present. How-ever, as Its regeneration seems to be dependentfrom forest elephants, it may become threat-ened in the near future in regions where theelephant populations are under much pressure,as is especially the case in WestAfrica

Prospects There is very little informationon many aspects of Ompholocorpum elotum,and an evaluation of its potential as a timbertree in managed natural forest and/or timberplantations Is recommended.

Major references Aubr6ville, 1964; Burkill,2000; Cooper & Record, 1931; Normand &Paquis, 1976; Takahashi, 1978

Other references Betti, 2004; Bouquet &Debray, 1974; de in Mensbruge, 1966; Diehl ata1. , 2004; Hawthorne, 1995; Nchanji &Plumptre, 2003; Pennington, 1991; Saville &Fox, 1967; Vivien & Faure, 1985; Wilks & Is-semb6, 2000.

Authors R. H. M. J. Lemmens

OREOBAMBOSBUCHWALDllK. Schum.

Protologue Notizbl. Bot. Oart. Berlin-DahlemI(5): 178 (1896)

Family Poaceae (Gramineae)Vernacular names Large green bamboo

(En)Origin and geographic distribution Oreo-

bombos buchwoldiiis restricted to eastern andsouth-eastern Africa, where it occurs scatteredin eastern DR Congo, Burundi, western Kenya,Uganda, Tanzania, Malawi, Zambia and Zim-babwe; it may also be present in South Africa(Northern Province)

Uses In Malawithe stems are used to makefences for livestock and flat, plate-like basketsIn Tanzania and northern Zambia they areused for construction including scaffolds andfencing, and less commonly for mats and has-ketry. People from Northern Provinces (SouthAfrica) fashion ritual flutes from the sterns of abamboo species, possibly Oreobombos 614ch-couldit.

Production and international trade Oreo-bombos buchwoldii sterns are only traded 10-cally; the economic value of international trade

.

is negligible, as is the case for other Africanbamboo products

Botany Bamboo growing in small densepatches or solitary clumps, with woody, green,hollow stems (CUIms) up to 20 in tall and up to10 cm in diameter, usually spreading or droop-ing. Leaves alternate, simple; sheath initiallycovered with appressed, stiff, brownish hairs;blade lanceolate or oblong-lanceolate, 10-35 cmx 2.5-6 cm, acutely acuminate, pale green toslightly bluish green, with parallel veins. Inflo-rescence a large, loose raceme with alternateclusters of spikelets; bracts ovate-elliptical,involucre-like, up to 1.5 cm long. Spikeletlanceolate to oblong-lanceolate, 12-15long, brown, 2-flowered with both norets bisex-ual;lower glume absent, upper glume 9-11mmlong, 11-18-veined; lemma broadly ovate toelliptical-ovate, 10-14 mm long, 11-23-veined;palea narrower, 8-12 min long, 5-11-veined;stamens 6; ovary superior, hairy at tip andhaving a single stigma. Fruit a caryopsis(grain), with a tuft of silky hairs on apexOreobombos comprises a single species, andseems most closely related to the large genusBombuso, which has its origin in tropical Asiaand America. It differs from Bombuso mainlyin its spikelet clusters surrounded by Involu-cre-like bracts

Plants do not flower for many years after es-tablishment. Gregarious flowering has beenrecorded in the Shire Highlands in Malawi, butin the Usambara Mountains in Tanzanla flow-

ering plants can reportedIy be found nearlyevery year. The plants die after flowering.

Ecology Oreobombos buchwoldii occurs inforest clearings and swampy forest, and alongstreams, at altitudes of 300-2000 in

Genetic resources and breeding Oreo-bombos buchwoldii occurs scattered and is, asfar as known, not planted, and might be liable

In Zambia it has been ex-to genetlc eroslontensively over-utilized and appears to be al-ready extinct in some localities. The total areacovered by Oreobumbos buchwoldii in Tanza-nia has been estimated at 19,000 ha. Inclusionin germplasm collections is desirable.

Prospects Oreobombos buchwo!dtiis one ofthe very few indigenous African bamboos. Verylittle is known about it. More research isneeded on cultivation techniques and potential

at a1. , 2000; Chihongo at a1. , 2000; ingomo,1990a

Authors R. H. M. J. Lemmens

ORMOCARPOPSISITREMOENSIS DUPuy &Labat

ORMOCARPOPSIS 4/1

Protologue Novon 6(I): 54, fig. I (1996)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Origin and geographic distribution Ormo-

corpopsis itreinoensis is only known from theItreino Massif, a smallregion in central Mada-gascar.

Uses The hard wood is used for house con-struction, carts and toolhandles.

Botany Deciduous shrub up to 3 in tall;stems with numerous robust lateral shortshoots. Leaves alternate, pinnately compoundwith 5-13 leaflets; stipules narrowly triangu-Iar, 4-5 mm long, persistent and closely spacedon short shoots; petiole and rachis groovedabove, pubescent; leaflets alternate, oblong-erriptica1, 9-11 min x 5 mm, wlth a sinaUinucro at apex, thinly leathery, glabrous above,shortly hairy below. Flowers clustered at theends of short shoots, bisexual, papilionaceous;pedicel short, elongating up to 1.5 cm in fruit;calyx c. 5 min long, with 5 slightly unequallobes; corolla yenow, with erect, rounded stan-dard c. I cm long, oblong wings, keel as long aswings; stamens 10, fused into 2 groups of 5;ovary superior, long-stalked, glandular hairy,I-celled, style slender, curved. Fruit an oblong-ellipsoid pod 2-5 cm long, with stipe up to I cmlong, straw-coloured, reticulately veined, inde-hiscent, (I-)2-4-seeded. Seeds ellipsoid, I-2 cmlong, pale brownOrmocorpopsis comprises 6 species and is en-demic to Madagascar. It is related to Ormocur-PMm, which differs in its segmented pods, andPeltiero, which differs in its I-seeded, dehis-cent pods.Ormocorpopsis itreinoensis grows slowly, withgrowth almost restricted to the slow extensionof the short shoots which produce flowers andleaves at their tips. Flowering is in Septemberwhen the plantis leafless

Ecology Ormocorpopsis itreinoensis occursin dry woodland on thin soil above marblerocks at 1300-1400 in altitude.

Genetic resources and breeding Ormo-corpopsis itreinoensis is restricted to a smallarea, which is under heavy pressure because ofmining for marble and forest degradation. It is

min

uses

Major references Bystriakova, Kapos &Lysenk0, 2004; Clayton, 1970; Coates Palgrave,1983; Launert, 1971; Williamson, 1955

Other references Beentje, 1994; Bingham

4/2 TIMBERSl

classified as criticalIy endangered in the JUGNred list of threatened species.

Prospects Protection of this rare and slow-growing species Is urgently needed, and har-vesting of its timber should be stopped jinme-diately to save it from extinction.

Major references du Puy at a1. , 2002Other references du Puy & Labat, 1998f;

Labat & du Puy, 1996; Schatz, 2001Authors R. H. M. J. Lemmens

OXYTENANTHERAABYSSINICA(ARIch. )Miniro

Protologue Trans. Litin. SOC. London 26: 127(1868)

Family Poaceae (Gramineae)Chromosome number2n= 72

Synonyms OryteiLortthero macrothyrsusK. Schum. (1895), Orytenonther0 670unii Pilg(1907), Orytenonther0 6072iiMattei(1909)

Vernacular names Savanna bamboo, Bin-dura bamboo, West African bamboo (En).Bambu africano (Po). Mwanzi(Sw)

Origin and geographic distribution Oryte-norither0 o6yssinicu is distributed throughouttropical Africa outside the humid forest zone,from Senegal east to ETitrea, and south to An-gola, Mozambique and northern South Africa.It is often planted

Uses The stems are widely used for con-struction, fencing, furniture and fish-trapsThey are also used for stakes, trellises, toolhandles, household implements and arrowshafts. The use of dry stems as fuelis wide-spread and they are sometimes made intocharcoal. The stems have some potential asraw material for paper making. Split stems are

used for basketry. Sap from the plant is col-Iected for wine making in Tanzania and Ma-lawi, the fresh or dried leaves are used as fod-der, and the seeds and young shoots as faminefood. Orytenonthero abysstnico is used inshelterbelts and windbreaks, and as a coinple-meritary crop in plantations of Cordi0 o17iconuLain. , Eucalyptus microtheco F. Muell. andKhoyo senegalerrsis A. Juss. in Sudan, for ero-sion control in land rehabilitation in Sudan

and Tanzania, and as an ornamental plantThe rhizome is used in the treatment of dysen-tery and the leaves are marketed for treatingdiabetes, colic and rheumatism. In Ethiopia therootis applied in the treatment of skin diseaseson the head. In Senegal leaf decoctions aretaken to treat polyuria, oedema and albuminu-ria

Production and international trade Onlylimited production information is available andmost estimates of the amount of bamboo pro-duced in Africa do not separate Orytenontheroo6yssinico from other species. Orytenontheroo6yssinico populations extend over 850,000 hain western Ethiopia, over 44,000 ha in Tanza-nia, at least 10,000 ha in Malawi, and 20,000ha in Senegal. Ethiopian Orytenonther0 o6ys-sinico stands possibly constitute over half thetotal area under bamboo in Africa. These

stands reportedIy comprise approximately 5300living stems (and 2700 dead stems) per ha. Ithas been calculated that the Ethiopian stand-ing crop (dry weight above-ground for livingstems) is 16.6 million t, and that there is po-tential for sustainably exploiting 5.5 million tannually. There Is no exporttrade in Orytenon-ther0 o6yssinico and it is used mostly nearwhere it grows. In the past, however, there hasbeen transportation of stems over long dis-tances from southern Sudan to supply demandin Khartoum, and formal commercialization ofharvests in Senegal. Price information forOrytenonther0 o6yssinicu materials, or prod-ucts derived from them, is not available

Properties The air-dry density of the stemwants 0.7-0.9 g/cm3. At 47% moisture content,the modulus of rupture is 82 N/min2 modulusof elasticity 14,600 N/mm2, compression paral-Iel to grain 49 N/min2 and shear (split stem) 11N/min2. Dried stems and fences made from thestems are susceptible to termite and borer at-tacks.

The stem contains approximately: holocellulose53-60%, pentosans 12-33%, 11gnin 15-27% andash I-4%. Solubilities are 6.5% in hot water,2.7% in alcohol-benzene and 27.6% in I%

"

* a^gt.

Oxytenonther0 o6yssinico- wild

a-

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,

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Q

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.

NaOH. Stem fibre cells of Oxytenonther0 o6ys-sinico have an average length of 2.0-2.8 min,an average diameter of 12-17 pin, a lumenwidth of 3-5 prn and a cell wanthickness of 5pin. In trials with different alkaline pulpingmethods, chlorine-free bleached pulps wereobtained of 82% ISO brightness and suitablefor writing and printing paper grades. TheOrytenonthero abyssinico pulps were similar tohardwood kraft pulpLeaves and twigs contain per 100 g dry mate-rial: crude protein 12.8-14.2 g, ether extract2.8-3.0 g, crude fibre 28.0 g, N-free extract36.4-40.0 g, ash 14.6-18.6 g, P 0.11-0.13 g, K0.60-0.66 g, Ca 0.25-0.40 g and Mg 0.32-0.35g. DigestIble protein and net energy levels areestimated, respectively, at 8.2 g and 3.7 MJ perkg dry matter. The fodder quality is low, due tolow energy value and high silica content

Description Clump-forming bamboo with arobust rhizome up to 10 cm in diameter; clumpdense, typically consisting of 20-100 stems;stems (CUIms) erect, ascending or leaning out-wards, 5-LOG15) in tall and 3-8(-10) cm indiameter, internodes 15-30(-40) cm long, thebasal ones solid, the distal ones thick-walled,

glabrous at maturity; young shoots grey-green,densely silky hairy. Leaves alternate, simple;sheath up to 15 cm long, with 2-5 min longbristles at top; 11gule short, c. 0.5 mm long;blade linear-lanceolate to oblong, 5-20(-26) cmx I-5 cm, base tapering into a short false peti-o1e, apex long-acuminate and pungent, some-what glaucous, with numerous longitudinalveins. Inflorescence a dense star-shaped cluster4-9 cm in diameter, with 10-20 spikeletsSpikelets sessile, narrowly lanceolate, 1.5-4.5cm long, pungent, I-4-flowered with uppernoret bisexual and lower florets male or sterile;lower glume 5-8 mm long, upper glume 8-10min long, lemma narrowly lanceolate, the low-est 12-20 min long, the uppermost about aslong as spikelet, tapering into a rigid spine upto 7 min long, palea narrowly lanceolate,somewhat shorter than lemma; noret with 6stamens, filaments united into a tube, and aglabrous ovary extending into a hollow styleterminating in 3 stigmas. Fruit a spindle-shaped caryopsis (grain) 10-15 mm long. Seed-ling with a short meSOCotyl and a loose coleop-tile, the first leaves without lainlna; primaryroot a pale taproot bearing short lateral roots

Other botanical information The dellmi-tation of Orytenonthero is unclear. Sometimesit is considered to comprise a single species, butsome authors include up to 15 species, most ofthem described from tropical Asia. It has alsobeen stated that Orytenonther0 o6yssinicotogether with another African bamboo, Oreo-bombos buchwoldii K. Schum. , should be trans-forred to Dendrocolomus, a large Asian genus.

Growth and development A single shoot,which may reach I in in height, is produced inthe first year from a rhizome. From the thirdseason onwards, several shoots are producedannuaUy. Rhizomes penetrate 30 cm in 3 yearsStems reach 1.2 cm in diameter and 1.8-3.0 in

in height within a few years of germination,reaching fullheight and diameter in 4-8 yearsNew stems break through the soil surface inthe rainy season. Extension growth slows downafter 3-4 weeks, and ceases after 2-4 monthsBranches develop from the upper nodes fromabout the fifth week of active stem growth.Foliage is mostly shed in the late dry seasonThe stems mature in 3 years and may survivefor 8 years, but they are over-mature and un-suitable for harvesting from 6 years of age on-wards. Clump diameters range from I in to 8in, and clumps may contain 20-100 (exception-ally up to 200) stems. New shoots appear at theperipheries of clumps. A clump longevity of 30

OXYTENANTHERA 4/3

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14'/

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I

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3

Orytenonther0 o6yssinicu - I, piece of stem, . 2,flowering broilchlets, . 3, spikeletRedrown und odopted by AChmod Sotiri Nur-homerit

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4/4 TIMBERSl

years has been estimated for Oxytenontheroo6yssinico in Sudan, but is less when massflowering occurs and rhizomes die with thestems. Mass flowering of Orytenunther0 o6ys-81n!co occurs every 7 years (Uganda), 14 years(Zambia) or 20-21 years onalawi), while spo-radic flowering has been widely and frequentlynoted. Both subsequent death of entire clumps(Chad, DR Congo) and regeneration of shootsfrom surviving rhizomes (Uganda) have beenrecorded. The fruit is dispersed as a largerpropagule (25-30 mm x 3-4 mm) includingseveral glumes; it is dislodged by animals andadheres to fur

Ecology Orytenonther0 o6yssinico is a low-land bamboo, occurring from sea-level up to2000 in altitude, but mainly at 300-1500 in. Itoccurs in savanna woodland subject to a cli-mate with an average annual rainfall of over800 min and 3-7 dry months (average rainfallless than 50 mm). It is absent from closed for-est and extends little into semi-and wooded

grassland and thicket. Prevailing average an-nualtemperatures are 20-27'C, with monthlyaverage daily maxima of 30-36'C and dailyminima of 7-17'C. Locally occasional frost mayoccur; if severe, this may scorch leavesOrytenonther0 o6yssinico grows on soils fromvarious types of parent rock, but over much ofits distributional range parent rock belongs tothe old crystalline basement complex. Soil rer-tinty is not a mayor influence. Orytenontheroo6yssinico is associated with impoverishedacrisols and ferralsols, moderately fertile Iuvi-sols, and younger relatively nutrient-rich cam-bisols and nitisols. The species is essentiallyabsent from arenosols that have poor moistureretention and gleysols having poor drainageKey site factors are good drainage combinedwith access to a reliable water supply. Charac-tenstic habitats are areas along rivers anddrainage lines, termite mounds and rockyslopes. Typical rocky slope microhabitats arewell-illuminated gullies with deep soil accumu-lated between boulders. Saline conditions areunfavourable

Propagation and planting Orytenontheroo6yssinico is usually raised from seed. Theweight of 1000 grainsis 70-110 g. They remainviable for 6-18 months if stored at ambient

temperatures under dry and pest-proof condi-tions. Storage for some months before use hasbeen advised. Germination percentages varyfrom 30% to 80% and germination periodsfrom11 days in warm moist conditions to 4 monthsin cool dry conditions. Seeds are sown in nurs-

eries when the mean daily temperature is 20'Cor higher. The nursery period may last 8-24months. Transfer to field sites is done in the

early part of the rainy season, suitable plantshaving two shoots, the larger one at least 30cm long.Where flowering is infrequent, rhizome divi-sion Is a realistic option for establishing newclumps and it is used traditionally to establishclumps in farmland. It has also been used bythe forestry service in Uganda. Early in therainy season, rhizome sections 12-30 cm longbearing healthy buds, or the lowest 45 cm of astem, are excavated and transplanted withoutdelay. Orytenonther0 o6yssinico can also bepropagated by stern cuttings.

Management Orytenonther0 o6yssinico israrely managed systematically, but weeding ofnewly established clumps, and removal ofoverhead shade, is recommended. Spacing ofnursery raised seedlings has been 3.8 in x 3.8in or 5 in x 5 in in experiments and in arboreta.In mixed plantations with hardwood trees thespacing is 6 in x 6 in. Clumps take up to 6years (from rhizome offsets) or 8 years (fromseedlings) to reach the stern harvesting stageFrom the third year, thinning (50% of shootsfrom previous years) appeared justified in aplanted stand in Kenya, because individualstems were smaller when stem numbers were

high. Clumps for sap ('wine') production aretypically established within areas cropped formaize, potatoes, pyrethrum or wheat. Theseclumps are thinned from the second year on-wards to prevent stem congestion, while prun-ing branches to around 2 in favours access, andloosening peripheral soil promotes rhizomeextension and unhindered shoot emergenceA pure stand of Orytenonther0 o6yssinico con-tains up to 750 clumps and 30,000 stems per ha

Diseases and pests Leafrust caused by thefungus Kuiei!ingio churno (synonym: Dusturellodiuino) has been recorded

Harvesting Harvesting of clumps starts whenthey contain 4-year-old stems; it continues atintervals of I-3 years, with 4-6-year-old sternsbeing cut. In a management system designed toimprove the productivity of natural standswith well-established clumps, rotational bar-vesting is practised during the first 4 yearsEach year shoots of all ages in one quadrantare cut; this treatment progresses round theclump year by year. Stems older than 6 yearsare used as fuel, those 4-6 years old as build-ing material, and those 2-3 years old may havevalue for weaving. After the initial phase, 4-

.

year-old sterns can be harvested annually or,for stems that are 4 or more years old, at 2-year or 3-year intervals. Apart from cullingdead or deformed stems, it is usual to leaveunharvested stems which are 3 years old oryounger, although clump congestion, or de-mand for supple material, may justify exploit-ing some of these.To collect wine in Tanzania, the tips of youngshoots are cut off, and the stem Is bruised twicea day during a week. The exudate from thebruises is collected and left to ferment for 2

days. The resulting wine ('ulanzi') contains 5-5.5% alcohol

Yield Yield estimates for natural vegetationwhere Orytenonther0 o6yssinico is abundantare 10-33 t dry stems per ha per year in Sene-gal. Experimental dry stern yields have beenreported equivalent to 8-11 t per ha per yearfrom DR Congo, and 14-28 t per ha per yearfrom Kenya. After a clump has reached an ageof 6 years such yields are sustainable underappropriate harvesting.

Genetic resources mequatorialdisjunctionand intervals east of Lake Chad and south ofEthiopia separate units of the range of Oryte, I-unther0 o6yssirtico. There are contrasting re-ports of annual flowering cycles in some placesand monocarpic clumps flowering at long in-tervals elsewhere, which could reflect geneticdifferences. This fact coupled with the frag-meritation and wide range of the species, sug-gest that there is appreciable genetic variation,prompting suggestions of subspecific taxa.There are no indications that Oxytenontheroabyssinico is currently threatened, but declinesin abundance have been reported, e. g. in Sene-gal and Uganda. Orytenonther0 o6yssinico hasbeen excluded, so far, from consideration by theFAO Panel of Experts on Forest Gene Re-sources although it is included in FAO'sREFORGEN information system, which sign-fies endangered status in Guinea. In responseto over-exploitation, bamboo forest reserveshave been gazetted in Uganda. The species hasbeen grown in a number of arboreta, but not asan explicit conservation initiative.

Prospects For savanna regions in Africa O^^-tenontheru o6yssinico has major attractions asa natural product amenable to sustainablemanagement. Straight, strong, light construc-tion materials can be generated in as little as 3years, and more items can be produced bysplitting stems with simple household toolsUndemanding propagation options are avail-able. Community bamboo stands established

and harvested following protocols optimizingproduct quality and quantity can be combinedwith land rehabilitation. Nevertheless, despitewide rural significance, Orytenonthero obys-sinico remained neglected in the forestry andagricultural sectors until the InternationalNetwork for Bamboo and Rattan (INBAR) re-cently included this species among 38 prioritybamboos of economic importance. ThroughINBAR there have now been appraisals ofbamboos in several African countries. Pros-

pects for cross-border management collabora-tion have resulted. Certain key areas of studyare relevant to continent-wide initiatives to

secure and improve Africa's savanna bambooresource, increasing its rural significance andvalue. One is clarification of geographic vana-tion in features of utility Interest, particularlystem size at maturity, a second one floweringcycles and their interaction with clump sur-vival, and a third is clarification of the conser-vation status. Fourthly, using knowledge of thedistribution to decide which geographic areasshould be represented, reciprocally planted,range-wide provenance trials monitoringgrowth and reproduction should be undertakenwith the aim of identifying superLor germplasmsources for various ecological conditions.

Major references Clayton, Harman & Wil-namson, 2002- b; Embaye, 2001; Binbaye ata1. , 2003; Fanshawe, 1972; Giffard, 1974;Inada, Kayambazinthu & Hall, 2003; ingomo& Kamiri, 1985; ingomo & Kamiri, 1987;Mgeni, 1983; Monteir0, 1949.

Other references Bayoumi, 1977; Burkill,1994; Bystriakova, Kapos & Lysenk0, 2004;Clayton, 1970; CTFT, 1962a; Doat, 1967; Fer-11n, 1970; Henke1, 1927; 1stas, Heremans &Raekelboom, 1956; 1stas & Raekelboom, 1962;Kerharo & Adam, 1974; Khristova at a1. , 2006b;Kobayashi, 1997; Launert, 1971; Le Hou6rou,1980; Rao, Rao & Williams, 1998; Rivi6re,1991; Shukla, Singh & Sanya1, 1988; Soder-strom & Ellis, 1987; Watson & Danwitz, 1992.

Sources of illustration Clayton, 1970Authors T. Triada & J. B. Hall

PARKiA 4/5

PARKiABicoLOR Achev.

Protologue Bull. SOC. Bot. France 55, meina 34 (1908).

Family Mimosaceae (Leguminosae - Mimo-soldeae)

Chromosome number2n=24

Vernacular names Faux Der6 (Fr)

.

4/6 TIMBERSl

Origin and geographic distribution Porkiobicolor occurs from Guinea and Sierra Leone

east to eastern DR Congo, and south throughGabon and Congo to Cabinda (Angola)

Uses The wood (trade names: eseng, essang,10) is used for planks, carpentry and canoes. Itis also suitable for light construction, interiortrim, joinery, furniture, cabinet work, ship-building, toys, novelties, implements, turnery,boxes, crates, matches, veneer, plywood, hard-board, particle board and pulpwoodThe fruit pulp is occasionally eaten and thefermented seeds serve as a condiment for sea-soning sauces and soups, in the same way asthose of African locust bean (Porkiu bigloboso(Jacq. ) R. Br. ex G. Don), although much lesscommonly so. The fruit is used as bait for fishand squirrels. The tree with its open, widelyspreading crown is used as a shade tree forcrops, and it is sometimes retained when cut-ting the forest for agricultural landSeveral parts of the tree are used in traditionalmedicine. A bark maceration is applied to treateye complaints, a bark decoction to treat tooth-ache, dried and powdered bark to enhancehealing of wounds and sores, and a vapourbath of the bark to treat rheumatism. Leaf

pulp is rubbed on smallpox and chicken poxThe roots serve to treat measles, infertility inwomen and sexually transmitted diseases.

Production and international trade A1-

though the wood of Porkio bicolor is often con-sidered to be of inferior quality, it is locallymarketed in C6te d'Ivoire and Cameroon, andprobably also elsewhere

Properties The heartwood is pale yellow,sometimes with darker irregular streaks; it isindistinctly demarcated from the wide sap-

wood. The grain is variable, from straight tointerlocked, texture coarse but even. The freshwood has an unpleasant odour.The wood is moderately light, with a density of460-630 kg/ina at 12% moisture content. It airdries rapidly, but distortion may occur. Therates of shrinkage are moderately high, fromgreen to oven dry 2.6-5.3% radial and7.5-9.6% tangential. The wood is moderatelystable to unstable in service

The wood is soft and moderately tough. At 12%moisture content, the modulus of rupture is35-126 N/mm2, modulus of elasticity9500-11,600 N/min2, compression parallel tograin 34-44 N/min2, shear 6.1-7.5 N/min2,cleavage 18-26 N/mm, Janka side hardness2470 N and Janka end hardness 3020 N

The wood saws moderately well and workssatisfactorily with both hand and machinetools. It can be planed to a smooth and lustroussurface, but the wood is difficult to polish andvarnish. The nailing and gluing properties aresatisfactory. It is not durable and liable to ter-mite, pinhole borer and marine borer attacksThe heartwood is moderately resistant totreatment with preservatives, the sapwood ispermeable.The wood yields about 57% sulphate pulp andis considered suitable for the paper industry.The seed oil has been analyzed and was foundto be non-toxic. It resembles the seed oil from

Porkio bigloboso. Arachidic acid is the mostabundant fatty acid (slightly over 40%), otheracids are behenic, stearic, palmitic, linoleicacids and the uncommon bicolargic acid(CmH, ,04-Preliminary phytochemical screening of theleaves demonstrated the presence of cardiacglycosides, tannins, alkaloids and steroids. Theroots contain gallic acid, 11chexanthone andIupeol. A root extract showed antibacterialactivity

Adulterations and substitutes The timber

of Porkio fineoideo WeIw. ex 011v. is similar tothat of Porkio bicolor and is sometimes tradedunder the same names

Description Medium-sized to fairly large,briefly deciduous tree up to 40 in tall; hole cy-11ndrical, usually straight but sometimestwisted or curved, up to 100(-150) cm in di-ameter, with thin and fairly spreading but-tresses up to 3(-6) in high; bark surface smoothto rough or scaly, shallowly pitted, grey-whiteto dark brown, inner bark granular or fibrous,brown or pinkish; crown widely spreading, urn-brella-shaped; young twigs densely reddish

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20 mm exserted; nectar-secreting flowers be-tween bisexual and staininodialflowers, calyxc. 5 mm long, stamens rudimentary, with nec-tar glands at base. Fruit a linear pod 25-45 cmx 1.5-3.5 cm, with stipe of 2-7 cm long, gla-brous, yellow to brownish red or wine red, be-coming black when old, with yellowish, mealypulp, indehiscent, up to 25-seeded. Seeds ellip-sold, 11-15 mm x 7-10 min, brown, with thin,membraneous seed coat. Seedling with epigealgermination; hypocoty1 5-7 cm long, epicotyl5-10 cm long; cotyledons thick and fleshy,rounded at apex; first leaves alternate, with2-3(-4) pairs of pinnae.

Other botanical information Porkio coin-

prises about 30 species and has a pantropicaldistribution. Only 3 species, all belonging tothe section Porkio, occur in continental Africa,and a fourth in Madagascar. The Porkio spe-cies from continental Africa seem to be closelyrelated, with Porkio Itlieoideo being the closestrelative of Porkio bicolor. The wood of Porkio

ingdogoscoriensis R. Vig. is used for construc-tion and planks. This poorly known speciesfrom moist forest in northern Madagascar re-sembles Porkio bicolor in leaves and innores-cences, but differs in its broad, flattened pods(5-9 cm broad). Porkio timoriono (DC. ) Merrfrom tropical Asia has occasionally beenplanted in East and southern Africa, but onlyas an ornamental and not as a timber tree

Anatomy Wood-anatointcal description (IAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;25: intervessel pits small (4-7 pin); 26: in-tervessel pits medium (7-10 pin); 29: vesturedpits; 30: vessel-ray pits with distinct borders;similar to intervessel pits in size and shapethroughout the ray cell; 42: mean tangentialdiameter of vessellumina 100-200 prn; 43:mean tangential diameter of vessellumina ^200 F1m; (45: vessels of two distinct diameterclasses, wood not ring-porous); 46: s 5 vesselsper square minimetre; (47: 5-20 vessels persquare minimetre); 58: gums and other depos-its in heartwood vessels. Tracheids and fibres61: fibres with simple to minutely borderedpits; 66: non-septate fibres present; 68: fibresvery thin-walled; 69: fibres thin- to thick-walled. Axial parenchyma: (76: axial paren-chyma diffuse); 80: axial parenchyma allform;81: axial parenchyma lozenge-allform; 83: axial

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PARKiA 4/7

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brown short-hairy. Leaves alternate, hipin-nately compound, up to 35(-45) cm long; stip-ules needle-shaped, c. 5 min long, caducous;petiole 5-10 cm long, swollen at base, abovethe base with an elliptical gland; rachis ribbed,reddish brown short-hairy, with 10-25 pairs ofpinnae; leaflets in (20-)28-50(-55) pairs perpinna, opposite, narrowly oblong, 5-10(-15)min x I-2.5 min, unequal at base, rounded toacute at apex, glabrous, only inIdrib prominentInflorescence a pendulous head arranged in araceme; head 5.5-8 cm x 3-4.5 cm, pinkish red,consisting of 2 parts, basal part initially cylin-drical, later depressed globose, apical part glo-hose; peduncle 5-25(-40) cm long. Flowersbisexual or sterile, regular, 5-merous, sessile;bisexual flowers in apical part of head, calyx7-11.5 min long with long tube, corolla lobesslightly longer, fused at base but free for morethan half their length, stamens 10, fused atbase, c. 7 mm exserted, ovary superior, shortlystiped, style slender, slightly shorter than sta-mens; staininodialflowers at base orbasal partof head, calyx c. 6 mm long, sterile stamens c

4/8 TIMBERSl

parenchyma confinent; (89: axial parenchymain marginal or in seemingly marginal bands);91: two cells per parenchyma strand; 92: four(3-4) cells per parenchyma strand; (93: eight(5-8) cells per parenchyma strand). Rays: 98larger rays commonly 4- to 10'seriate; 104: allray ceUs procumbent; 1/5: 4-12 rays per minMineral inclusions: 136: prismatic crystalspresent; 142: prismatic crystals in chamberedaxial parenchyma cells.(E. Ebanyenle, A. A. Oteng-Amoako & P. Baas)

Growth and development Seedlings canbecome I in tall in the first year. In C6ted'Ivoire the mean annual growth in diameter isabout 0.7 cm for trees with a diameter at

breast height of 10-70 cm in natural foreststands, and 1.7 cm per year in 14-year-oldplantations. The tree is deciduous after therainy season, and new leaves develop inbronze-red flushes, immediately followed bythe flowers still in the dry season. However, ithas been reported from south-western Camer-o0n that new leaves develop after fruiting atthe start of the rainy season. Individual treesflower for 6-8 weeks. At night, the sterile flow-ers near the base of the inflorescence produceabundant nectar, which attracts bats. Theflowers are probably most commonly pollinatedby these bats, but dormice and pottos have alsobeen recorded as feeding on the nectar. Fruitsdevelop in about 2 months. Monkeys, chimpan-zees and gorillas feed on the fruit pulp, andmay disperse the seeds. In C6te d'Ivoire fruit-ing occurs in February-March. Porkio bicolordoes not nodulate

Ecology Porkio bicolor occurs in a range offorest types, from wet evergreen forest to drysemi-deciduous forest. However, it is most

in mixed, moist, lowland,common semi-

evergreen rainforest, espedally in well-drainedplaces, but it can also be found along creeksand rivers and in swamps. In Cameroon it ismainly found in rainforest in the coastal zonetogether with numerous Coeso!pintoceoe spe-cies, but it extends along watercourses intomoist semideciduous forest. It occurs in regionswith mean annual rainfall of 2000-4000 min,and with a dry season of up to 3 months. It isfound up to 1200 in altitude. In some regions,e. g. in southern C6te d'TVoire, it is one of themost frequent large trees. Regeneration isfairly common in secondary forest

Propagation and planting The 1000-seedweightis 300-500 g. The seed loses its viahintyrapidly and should be sown quickly. Removal ofthe seed coat improves germination. Germina-

tion occurs 4-10 days after sowing. The seed-lings are sensitive to damping off. Young seed-lings are often found close to the mother treeand tolerate some shade, but saplings are lightdemanders. Porkio bicolor is classified as a

non-pioneer light demanderManagement In Gabon Porkio bicolor is

recorded as occurring in fairly low densities (onaverage 0.74 matha of total wood volume), andthis is also the case in south-western Camer-

o0n (0.72 ing/ha). In Cameroon the minimumfelling diameter is 60 cm.

Handling after harvest After felling thelogs should be removed from the forest rapidlyor be treated with a preservative immediately,because they are liable to blue stain attack

Genetic resources There are no indications

that Porkio bicolor is threatened by geneticerosion. It is not only widespread, but is alsocommon in several regions and has a wide eco-logical adaptation, including secondary forest

Prospects Untilrecently the wood of Porkiobicolor was considered of inferior quality. Nowthat the stands of many commercial timberspecies have become depleted in many regions,it is becoming more important, especially forlocal construction and joinery, but presumablyalso for export in the future. Tests on its eco-nomic potential for veneer and plywood produc-tion seem worthwhile. Much research is still

needed on growth rates, ecological require-merits and management of naturalstands on asustainable basis

Major references Bolza & Keating, 1972;Burki11, 1995; de Saint-Aubin, 1963; Hopkins,1983; Hopkins & White, 1984; SIepel, Poorter& Hawthorne, 2004; Takahashi, 1978; Viniers,1989; Voorhoeve, 1979; Wilks & TSSemb6, 2000

Other references Aiyelaagbe, Ajaiyeoba &Ekunday0, 1996; Ajaiyeoba, 2002; Ako6gninou,van der Burg & van der Maesen, 2006; Au-br6ville, 1959c; Bouquet, 1969; de in Mens-bruge, 1966; Dupuy, 1998; Durrieu de Madronat a1. , 1998a; Fotie at a1. , 2004; Gilbert & Bou-tique, 1952; Hawthorne, 1995; Hawthorne &Jongkind, 2006; InsideWood, undated; Luckow& Hopkins, 1995; Neuwinger, 2000; Normand& Paquis, 1976; Raponda-Walker & Sillans,1961; Sabatie, 1989; Sprent, 2001; Vivien &Faure, 1985; White & Ahemathy, 1997

Sources of illustration Voorhoeve, 1979;Wilks & Issemb6, 2000.

Authors AT. Tchinda

.

PARKIAFILICOIDEAWelw. ex 011v.

Protologue F1. trop. Mr. 2: 324 (1871)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Vernacular names ununde, inlopa, limenze,

innienzi, innyeusi(Sw).Origin and geographic distribution Porkio

flitcoideo is widespread from eastern C6ted'Ivoire east to southern Somalia and eastern

Kenya, and south to Angola, Zambia andnorthern Mozambique

Uses In East and southern Africa the woodis used for poles, mortars, water containers,stools and beehives. It is suitable for light con-struction, interior trim, furniture, cabinetwork, vehicle bodies, sporting goods, toys, nov-elties, handles, boxes, crates, matches, ply-wood, hardboard and particle board. It is alsoused asfirewood and for charcoal productionYoung pods, cut into pieces, and seeds are oc-casionally cooked and eaten, especially in perl-ods of food shortage. The fruit pulp Is alsoeaten, e. g. in Zambia where fruits ripen fromDecember to February, when rural food re-serves are at their lowest. The pods serve asfodder for livestock. The bark is used in tradi-tional medicine. A bark decoction is taken as a

galactagogue, and for treatment of malaria,rheumatism and toothache. The tree with its

open, widely spreading crown Is used as shadetree for crops, and it is sometimes planted asan ornamental. It provldes nectar for honeybees

Properties The heartwood is straw-colouredwith a pink or green sheen, turning reddishbrown on exposure, and often indistinctly de-marcated from the sapwood. The grain is inter-locked, texture coarse. Growth rings distinct.Theftesh woodhas an unpleasantodourThe wood has a density of 450-580 kg/ms at12% moisture content. The rates of shrinkagefrom green to 12% moisture content are 2.8%radial and 3.8% tangential. The wood air drLesrapidly, but with a tendency to twist and cup.Quarter-sawing before drying is recommendedThe wood is unstable in service

The wood is soft. At 12% moisture content, themodulus of rupture is 1/5 N/min2, modulus ofelasticity 10,200 N/min2, compression parallelto grain 48 N/min2, cleavage 16 N/min andChalais-Meudon hardness 2.9.

The wood is moderately easy to work withhand and machine tools, and a reduced cuttingangle is necessary to avoid picking-up duringplaning. The nailing and gluing properties are

satisfactory. The wood is not durable and isliable to termite, pinhole borer and marineborer attacks, but it is comparatively easy totreat with preservativesThe seed of Porkio Intocotdeoe has a high pro-teln content (32%) and low fat content (10%).The seed oil contains 54% unsaturated fattyacids; the main fatty acids are oleic acid (43%),stearic acid (17%) and linoleic acid (11%)

Botany Medium-sized, briefly deciduous treeup to 30(-40) in tall; hole cylindrical, usuallystraight, up to 120 cm in diameter, with thinand fairly spreading buttresses up to 2(-4) inhigh; bark surface smooth to slightly fissured,pale grey to yellowish brown, inner bark pink-ish red; crown widely spreading, umbrella-shaped; young branches short-hairy to gla-brous. Leaves alternate, hipinnately compound,up to 40 cm long; stipules needle-shaped, c. 5min long, caducous; petiole 5-12.5 cm long,swollen at base, with a deeply lobed gland;rachis short-hairy, with 4-12 pairs of pinnae;leaflets in 12-28(-32) pairs per pinna, oppositeto slightly alternate, oblong, 14-35 mm x 3.5-13 min, unequal at base, notched or rounded toacute at apex, glabrous or finely hairy, inIdriband I-2 other basal veins prominent. Inflores-cence a pendulous head arranged in a raceme;head 6-9 cm x 3.5-7.5 cm, bright red, DCca-SIonaUy white, consisting of 2 parts, basal partcylindrical to depressed globose, apical partglobose; peduncle 8-34(-40) cm long. Flowersbisexual or sterile, regular, 5-merous, sessile;bisexual flowers in aptcal part of head, calyx10-16 mm long with long tube, corolla lobesslightly longer, fused at base but free for morethan half their length, stamens 10, fused atbase, c. 3 mm exserted, ovary superior, shortlystiped, style slender, shorter than stamens;staininodialflowers at base of basal part ofhead, few, calyx c. 5 min long, sterile stamensslightly exserted; nectar-secreting flowers be-tween bisexual and staininodialflowers, calyx5-9 min long, stamens rudimentary, with nec-tar glands at base. Fruit a linear pod 25-45(-80) cm x 1.5-3.5 cm, with stipe of 2-10 cmlong, usually glabrous, dark green, becomingblack when old, with orange, mealy pulp, Inde-hiscent, up to 21-seeded. Seeds ellipsoid, 16-25mm x 10-13 min, brown, with thin, meinbra-neous seed coat. Seedling with epigeal germi-natlon

Porkio comprises about 30 species and has apantropical distribution. Only 3 species, anbelonging to the section Porkio, occur in conti-nentalAfrica, and a fourth in Madagascar. The

PARKIA 4/9

420 TIMBERSl

Porkio species from continental Africa seem tobe closely related, with Porkio bicolor A. Chev.being the closest relative of Porkio fineoideuThe name Porkio fineoideo has commonly beenmisapplied to Porkio big!o6080 (Jacq. ) R. Br. exG. Don, the well-known African locust bean,famous for its fermented seeds serving as acondiment; therefore, interpretation of the lit-erature should be done with care.

Porkio fineoideo trees are leafless for a briefperiod, often during the flowering period in thedry season. It has been reported that floweringtrees can be found throughout the year inKenya and Tanzania. At night, the sterile flow-ers near the base of the inflorescence produceabundant nectar, which attracts bats. Theflowers are probably most commonly pollinatedby these bats, but squirrels have also been re-corded as feeding on the nectar. Fruits developin 2-3 months. Monkeys, baboons, chimpan-zees, gorillas and elephants feed on the fruitpulp, and may disperse the seeds. Birds suchas parrots and bombills eatthe seeds

Ecology Porkio jiltcoideo occurs in wet ever-green forest to semi-deciduous forest. In Cen-tralAfrica it seems to occupy drier forest typesthan Porkio bicolor. In East and southern of-

rica it prefers swamp forest and riparian for-est, up to 1300 in altitude, but it may be foundin rainforest on well-drained localities. In West

Africa Porkio lineoideo often occurs scatteredin highland forest and in semi-evergreen rain-forest along the northern fringe of the Gumeo-Congolian region

Management Porkio nitcoideo can be pro-pagated by seed. The seeds have no period ofdormancy and deteriorate rapidly, so theyshould be sown quickly. Removal of the seed-coat, boiling for a short time or soaking in wa-ter overnight Improve germination. Freshseeds extracted from the fruit already startedto germinate without any contact with externalwater. In Kenya seeds found in fresh elephantand baboon faeces germinated soon after conec-tion. In tests in Malawi, however, seeds failedto germinate regardless of the pre-treatmentPods and seeds are easily attacked by insectsAfter felling, the logs should be removed fromthe forest rapidly because they are liable toblue stain attack.

Genetic resources and breeding FJrhiofineoideo is widespread, but in general of scat-tered distribution. There are no indications

that it is threatened by genetic erosionProspects Porkio nitroideo is not likely to

become of economic importance because It 00

curs in low densities and its timber is of rather

poor quality. Research on its possibilities as afruit tree appears to be desirable, particularlyin combination with other applications, e. g. asshade tree, forage, and fruit and seed vegeta-ble

Major references Bolza & Keating, 1972;Fouarge & G6rard, 1964; Hopkins, 1983; Hop-kms & White, 1984; Ruffo, Birnie & Tengnas,2002.

Other references Burki11, 1995; Engel,2000; Ibiyemi, 1987; Katende, Birnie &Tengiias, 1995; Mateke, Kamara & Chikasa,1995; Neuwinger, 2000; Tanzania Forest Divi-sion, 1967; Vivien & Faure, 1985; White &Ahemathy, 1997; Wimbush, 1957

Authors R. H. M. J. Lemmens

PERICOPSISANGOLENSiS (Baker) Meeuwen

Protologue Bull. Jard. Bot. Etat 32(2): 216(1962)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Chromosome number2n= 18

Synonyms Afrormosio origolensis (Baker)Harms (1913).

Vernacular names EastAfrican afrormosia

(En). Pau ferro, gainbo, inuanga (Po). Mu-vanga, inbanga (Sw).

Origin and geographic distribution Pert-copsis origolensis occurs from eastern DRCongo and Rwanda east to Tanzania, andsouth to Zambia, Angola, Zimbabwe and Mo-zambique.

Uses In Malawithe extremely durable woodwas highly esteemed for hoes and pestles, and

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for rims and hubs of wagon wheels. At presentthe wood is used for flooring and panelling, andit Is also suitable for heavy construction, rail-way sleepers, mine props, ship building, vehi-cle bodies, fence poles, vats, interior trim, join-ery, furniture, cabinet work, handles, ladders,agricultural implements, sporting goods, musi-cal instruments, toys, novelties, precisionequipment, carvings and turnery. In ZimbabwePertcopsis origolensis is one of the most impor-tant trees for poles used in local house con-struction. It is also used as firewood; it is diffi-cult to light, but produces great heat and littleash. It makes excellent charcoal.

In traditional medicine the roots, bark andleaves are commonly used. Root decoctions areused to stimulate the blood circulation, to treatdiarrhoea, bronchial and chest complaints,nausea, and eye problems. They are consideredtonic, abortifacient and aphrodisiac. Root POW-der Is applied externalIy to relieve pain, and totreat oedema and tumours. Bark decoctions or

macerations are taken to treat diarrhoea, sorethroat and toothache, and as eye bath. Leafsapis drunk as arithelmintic, whereas groundleaves are applied externalIy against headacheor vapour of a leaf decoction is inhaled for thispurpose

Production and international trade Pert-

copsis origolensis trees often occur too scatteredor their holes are too smallor poorly shaped forcommercial exploitation, except in Mozam-bique, where they are locally of some impor-tance for the timber market. The wood istraded in small amounts on the international

timber market, either under the name 'inu-wariga', or mixed with the wood of Pencopsiseloto (Harms) Meeuwen from West and Cen-tralAfrica as 'afrormosia'.

Properties The heartwood is greenish brown,turning dark brown to almost black upon dry-ing. It is distinctly demarcated from the whit-ish to yellowish grey, up to 2.5 cm wide sap-wood. The grain is interlocked, texture moder-ately fine. The wood is strikingly banded orshows a whorled figure. It is oily to the touchThe wood is heavy, with a density of 930-1030kg/ms at 12% moisture content. The wood airdries very slowly but with little degrade, exceptslight surface checking. The rates of shrinkageare low, from green to oven dry 2.0% radial and2.8% tangential. Once dry, the wood is moder-ately stable in serviceThe wood is very hard. At 12% moisture con-tent, the modulus of rupture is 80-106 N/min2,modulus of elasticity 12,600-13,100 N/mm2,

compression parallel to grain 64-73 N/min2,shear 13-16 N/mm2 and Janka side hardness

9070-12,230 N.The wood is difficult to saw and work because

of its high density. It is rather difficult to planeand a cutting angle of 20% is recommended,but it has a smooth finish. The wood holds

nails and screws well, but pre-boring is neces-sary. Gluing, staining and polishing do notcause problems. The wood turns well. Thebending properties are moderate. The wood isvery durable and resistant to fungi, termitesand allwood borers including marine borers. Itis also highly resistant to abrasive action andchemicals. In Malawi, where the wood of Pert-copsis origolensis is considered the most dura-ble native wood, remains of hoes made from thewood have been found that were at least 90

years oldAdulterations and substitutes The wood

of Pertcopsis origolensis resembles that of Pert-copsis e!oto, but is slightly heavier, darker andmore durable. It has some resemblance to the

wood of Bothioeo pittr4iugo Harms, which is astrong, durable wood used for similar purposes

Description Deciduous smallto medium-sizedtree up to 20(-27) in tall; hole branchless for up

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to 7.5 in, often curved or twisted, up to 100 cmin diameter; bark surface whitish grey tocreamy brown, initially smooth but later Ir-regularly fissured and flaking in thin piecesleaving red-brown patches, inner bark fibrous,yellowish, darkening rapidly on exposure;young twigs hairy but glabrescent. Leaves al-ternate, Imparipinnately compound with7-11(-13) leaflets; stipules spoon-shaped, 5-6min long, caducous; petiole and rachis together10-21 cm long; stipels thread-like, 2-5 minlong, caducous; petiolules 2-4 mm long;leafletsalternate, sometimes nearly opposite, ovate toelliptical, (2-)3.5-6.5G9.5) cm x (1.5-)2-3,565)cm, curieate to rounded at base, rounded tonotched at apex, hairy to glabrous below, pin-nately veined with 6-10 pairs of veins. Inno-rescence a panicle up to 15 cm long at the endof a shoot, hairy; bracts up to 1.5 mm long,caducous. Flowers bisexual, papilionaceous,whitish with purple veins to pink or purplish;pedice1 10-12 mm long, slender; calyx cam-panulate, 7-13 mm long, with lobes muchlonger than the tube, upper 2 lobes partlyfused, hairy; corolla with nearly circular stan-dard 13-15 mm in diameter, clawed, wing andkeel petals c. 18 mm long; stamens 10, free,8-14 mm long, glabrous; ovary superior, flat-tened, c. 9 min long, hairy, style slender, up-curved. Fruit an oblong-linear, flattened pod7-24 cm x 2-4 cm, shortly stiped, slightlywinged along margins, pale brown, smooth,glabrous or sometimes hairy, reticulatelyveined, indehiscent, I-4-seeded. Seeds disk-shaped, 12-15 mm in diameter, reddish

Other botanical information Pertcopsiscomprises 4 species, 3 of which occur in tropicalAfrica and I in tropical Asia. Pertcopsis origo-lensis is variable in the hairiness of the leaves

and fruits, and this has resulted in the distinc-tion of infraspecific taxaPertcopsis 10n/1070 (Benth. ) Meeuwen is asmall savanna tree occurring from Senegal toSudan that has been considered a subspecies ofPertcopsis origolensis. Its wood is similar andcan be used for the same purposes, but largersizes are not available. Pertcopsis 10n/lord ismore important as a medicinal plant

Growth and development In southernAfrica Pertcopsis origo!errsis trees flower fromSeptember to November, whereas fruits ina-ture from Aprilto May. The roots have nodulescontalning nitrogen-fixing bacteria.

Ecology Pertcopsis origo!errsis is IOCaUy coin-mon in deciduous, open or more or less closedwoodland and wooded grassland up to 1650 in

altitude. It is often found in intoinbo woodland,in association with Brachystegio, Combretumand Terming!to species. It has been reportedthat the occurrence of Pertcopsis origolensistrees in Malawiis an indication of fairly fertilesoil. The trees are fire resistant, but sensitiveto frost.

Propagation and planting In experimentsin Tanzania, seedlings showed poor survival,usually less than 60%, but the cause remainsunclear

Management In injoinbo woodland wherePertcopsis origolensis is common, a system ofcoppicing is often applied. For firewood produc-tion, a coppice rotation of 5 years Is possible,but for poles and certainly for larger sizes oftimber much longer rotation cycles are neededPertcopsis origolensis responds well to coppic-ing, although it often produces a large numberof small shoots. In some areas, e. g. in Zambia,the trees are considered too hard to cut, andoften left in the forest after cutting of otherspecies.

Harvesting The hardness of the wood is aserious drawback for felling using axes, andmore modern equipment, Including chainsaws,is preferred.

Genetic resources Pertcopsis origolensis iswidespread and locally common, and notthreatened. However, in several areas it hasbecome vulnerable because of overexploitation,

in Malawi where the boles are much col-e. g

Iected for poles and firewood, and the roots andbark for local medicine. In these areas largertrees are difficult to find, whereas inferior treespredominate in the remaining populations

Prospects Pencopsis origolensis produces ahighly valued timber. Its main constraint forlarger-scale commercial exploitation is itssmall and often poorly shaped hole. There isstintoo little information on growth rates andappropriate management schedules to developmodels for sustainable exploitation of poles andfirewood through coppicing. Harvesting thebark and roots for medicinal purposes is oftendestructive for the trees, and the developmentand promotion of harvesting techniques that donot kill the trees are needed. Considering itswide use in traditional medicine, the scarcity ofinformation on phytochemistry and pharma-cologicalproperties is astonishing

Major references Bolza & Keating, 1972;Bryce, 1967; Coates Palgrave, 1957; CoatesPalgrave, 1983; Gillett at a1. , 1971; Takahashi,1978; Toussaint at a1. , 1953; Williamson, 1955.

Other references Abbott & Lowore, 1999;

.

Dietrichs & SImatupang, 1974; Fitzgerald,Gunning & Donnelly, 1976; Gelfand at al. ,1985; Kokwar0, 1993; Neuwinger, 2000; vanWyk & van Wyk, 1997; Vyamana, Chainshama& Mugasha, 2007.

Sources of illustration Coates Palgrave,1957; Gillett at a1. , 1971

Authors A. U. Lumbile & 0. 0agile

PERICOPSISELATA (Harms) Meeuwen

Protologue Bull. Jard. Bot. Etat 32(2): 216(1962).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

SynonymsAfrormosio eloto Harms(1913)Vernacular names Afrormosia, African teak

(En). Afrormosia, assamela (Fr). Afrormosia,assamela (Po)

Origin and geographic distribution Pert-copsis elata occurs from Cote d'TVoire east tothe Central African Republic and DR Congo. Ithas been introduced in western Uganda.

Uses The wood (trade names: afrormosia,assamela, kokrodua) is highly valued on theinternational market, mainly for furniture andas decorative veneer, but also for interior andexterior joinery, stairs, flooring and boat build-ing. It is considered a substitute for teak. It isalso suitable for heavy and light construction,railway sleepers, vehicle bodies, Interlor trim,handles, ladders, agricultural implements,sporting goods, musical instruments, toys, nov-elties, boxes, crates, carvings, turnery anddraining boardsIn traditional medicine in Congo pulped bark isrubbed into scarifications as an anodyne.

Production and international trade Exploi-tation of Pertcopsis eloto for the internationaltrade started in 1948 in Ghana, soon followedby C6te d'Ivoire. The wood was an importantexport commodity from Ghana, but excessiveexploitation has drastically reduced the num-hers of exploitable trees. In 1966 Ghana ex-ported 8600 ina of logs and 19,600 in3 of sawnwood, in 2001 only 150 ina of logs. In the sameyear C6te d'Ivoire was the main exporter with68,000 ina of logs and 15,000 ina of sawn woodMore recently, Cameroon, Congo and DRCongo have become the main exporters, withonly smallvolumes exported from C6te d'Ivoireand Ghana. Cameroon exported 13,000 in3sawn wood in 2003, 7500m3 in 2004 and 6200ina in 2006. Congo exported 3000 ina of sawnPertcopsis eloto wood in 2003 at an averageprice of Us$ 371 per ms. and in 2004 the exportvolume was 5000 ms at an average price of Us$324 per ms. Pertcopsis eloto is among the fourmost Importanttimber Species in Congo

Properties The heartwood is yenowish brownto greenish brown with darker streaks, fairlydistinctly demarcated from the pale yellow-brown, up to 3 cm wide sapwood. The grain isstraight, sometimes interlocked, texture inod-erately fine. Quartersawn surfaces are roottled.The wood resembles that of teak, but lacks theoily textureThe wood is moderately heavy, with a densityof (620,700-800 kg/in, at 12% moisture con-tent. The wood air dries slowly and with littledegrade; it can be kiln dried under moderateschedules with good results. The rates ofshrinkage are moderate, from green to ovendry 3.0-3.7% radial and 5.8-7.0% tangential.Once dry, the wood is very stable in serviceAt 12% moisture content, the modulus of rup-ture is 93-155 N/min2, modulus of elasticity9450-13,300 N/min2, compression parallel tograin 63-71 N/min2, shear 9-16 N/min2, cleav-

17-24 N/mm and Janka side hardnessage6940-7115 N

The wood saws and works wellwith both hand

and machine tools. However, the blunting ef-rect is fairly high and stellite-tipped saw teethand tungsten-carbide-tipped cutting tools arerecommended. The wood has a smooth finish

and usually planes well, but sometimes with aslight picking up due to the presence of inter-locked grain; a cutting angle of 20' is advisedIt polishes satisfactorily, but the use of a filleris recommended. The wood is liable to splittingwhen nailed, but it holds nails and screws well;pre-boring is recommended. Gluing does not

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pose problems when it is done carefully to pre-vent staining. The bending properties are inod-erate. The wood must be well steamed for ve-

neer production. It turns satisfactorilyThe wood is very durable, but may occasionallybe attacked by marine borers. The heartwood isresistant to impregnation by preservatives, thesapwood fairly permeable. Wood dust maycause irritation to the nose, throat and eyes.Under moist conditions, the wood often stainsdark where it is in contact with iron.

Methanol extracts of the leaves showed moder-

ate in-vitro antlPIasmodial activity againstmultiresistant strains of Pigsmodium Idleipo-ruin. The wood contains stilbene derivatives

with antibiotic, antimalarial and blood-sugarreducing properties. They also play a role inthe colour development of the wood

Description Medium-sized to large tree upto 45(-50) in tall; hole branchless for up to 30(-35) in, straight and cylindrical, sometimestwisted, up to 130(-150) cm in diameter, oftenwith low and blunt buttresses; bark surfacecreamy or greyish, flaking in thin pieces Ieav-ing red-brown patches, inner bark yellowish,darkening to dirty orange; crown flat-topped,with massive, spreading branches; branchlets

drooping, glabrous. Leaves alternate, imparip-innateIy compound with (5-)7-9(-11) leaflets;stipules linear, c. 7 min long, caducous; petioleand rachis together 7-20 cm long, slightlygrooved on upper surface; stipels thread-like,2-5 mm long, persistent; petiolules 3-7 minlong; leaflets alternate, elliptical to ovate-elliptical, 3-8 cm x 1.5-3.5 cm, terminal onelargest, slightly curieate to rounded at base,shortly acuminate at apex, glabrous, pinnatelyveined with 5-11 pairs of fine veins. Inflores-cence a slender, drooping panicle up to 12 cmlong at the end of a shoot, sparsely hairy;bracts up to 4 min long. Flowers bisexual,papilionaceous, white to creamy or greenish;pedice1 5-9 min long, slender; calyx campanu-late, 6-12 mm long, with large triangularlobes, finely hairy; corolla with nearly circularstandard 11-13 mm in diameter, clawed, wingand keel petals c. 15 min long; stamens 10,free, 9-19 mm long, glabrous; ovary superior,flattened, 5-6 min long, hairy, style slender,UPCurved. Fruit an oblong-linear, flattened pod7-17 cm x 2.5-3 cm, shortly stiped, slightlywinged along margins, brown, smooth, reticu-lately veined, indehiscent, I-4-seeded. Seedsdisk-shaped, 12-15 mm in diameter, brown.Seedling with epigeal germination; hypocotylI-2 cm long, epicoty1 4-5 cm long; cotyledonsovate, c. 12 min long, slightly fleshy; first 2leaves opposite, simple, next leaves alternateand 3-foliolate.

Other botanical information Pertcopsiscomprises 4 species, 3 of which occur in tropicalAfrica and I in tropical Asia. The latter, Pert-copsis mooniono (Thwaites) Thwaites, washeavily exploited for its valuable timber inSouth-East Asia, but stands have been de-PIeted in many regions

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: a wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); (25: intervessel pits small (4-7 prn)); 26:intervessel pits medium (7-10 prn); 29: ves-tured pits; 30: vessel-ray pits with distinct bor-ders; similar to intervessel pits in size andshape throughout the ray cell; 41: mean tan-gential diameter of vessellumina 50-100 pin;42: mean tangential diameter of vessellumina100-200 pin; 47: 5-20 vessels per square inil-limetre; 58: gums and other deposits in heart-wood vessels. Tracheids and fibres: 61: fibres

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with simple to minutely bordered pits; 66: non-septate fibres present; 69: fibres thin- to thick-walled; 70: fibres very thick-walled. Axial pa-renchyma: (76: axial parenchyma diffuse); 78:axial parenchyma scanty paratrachea1; 79:axial parenchyma vasicentric; (80: axial paren-chyma allform); (81: axial parenchyma lozenge-anform); 83: axial parenchyma confluent; 84axial parenchyma unilateral paratrachea1; (89:axial parenchyma in marginal or in seeminglymarginal bands); 92: four (3-4) 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 min. Stoned structure: 1/8: all rays sto-Tied; 120: axial parenchyma and/or vessel ele-merits stoned. Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cells(P. Ng'andwe, H. Beeckman & P. E. Gasson)

Growth and development Initial growthmay be rapid in suitable conditions, with youngtrees attaining 8 in tallwith a hole diameter of9 cm after 7 years and 26 in tall in 16 yearsPlantation trials in C6te d'TVoire showed

growth rates up to 20 cm in bole diameter in 20years. Saplings tend to have a spreading,bushy habit. In Ghana trees flower inApril-May and fruit in August-November.Seedlings are usually found close to mothertrees, up to a distance of 35 in, indicating thatthe non-dehiscing pods are spread over onlyshort distances, although they may occasion-ally be spread over longer distances by strongwinds. The roots have nodules containing ni-trogen-fixing bacteria

Ecology Pertcopsis eloto is characteristic ofsemi-deciduous forest, especially in swampyand flat, disturbed localities. It occurs in re-gions with 1000-1500 min annual rainfall

Propagation and planting The 1000-seedweight is 200-450 g. Although seed productionis often abundant, lack of natural regenerationhas often been reported. The seeds of Pertcop-sis eloto germinate rapidly, in about 8 days,both in deep forest shade and in small gaps inthe forest, butthe seedlings only develop in thegaps and die in the deep shade. In fullsunlightthe germination rate is very low, only about5%. Localities where seedlings receive fullsunlight in the morning but which are shadedfrom direct midday sun have been recorded asideal for seedling growth. The seedlings areremarkably drought tolerant and can be foundon wet as well as dry forest soils. Sowing tests

showed that seedlings grow best when theseeds have been covered with 1.5 cm of soil. In

experiments, propagation by stem cuttingsshowed good results.

Management In a nursery experiment inGhana, the application of NPK fertilizershowed positive results on seedling growth, butthe potting medium (pure compost or compostmixed with up to 50% sand) had no influenceon seedling growth. In Ghana and DR CongoPertcopsis e!otu has been successfully used insmall-scale enrichment planting (line andgroup planting) as well as in taungya and pureplantations. For trees in natural forest, theminimum felling diameter is 50 cm in C6ted'TVoire, 60 cm in DR Congo, 100 cm in Camer-Don and 110 cmin Ghana.

Diseases and pests Larvae of the mothLamprosemu latentio!is have been recorded asa pest of seedlings and young trees. The effectsof defoliation by this insect can be serious andcan cause high rates of seedling mortality. Insome localities in Ghana, the fungus Beauueriobossiono was identified as a natural enemy ofLumprosemo loteritiolis

Harvesting In Ghana there is a minlmumfelling limit of 110 cm in diameter for Pertcop-sts e!oto, but even then logging is only allowedwith a permit. In Garnero0n the minimum di-ameter limit for Pertcopsis eloto ls 100 cm, butit has been recommended that this be reducedto 80 cm to relieve the pressure on lower di-ameter classes, which are cut indiscriminatelyand illegalIy due to the lack of trees with a bolediameter over 100 cm. In Congo the minimumexploitable diameter limitis set at 60 cm

Handling after harvest In several coun-tries (Ghana, Nigeria, Cameroon) the export oflogs has been banned, and logs are processedwithin the country of production. In Cameroonparquet flooring is an important product forexport

Genetic resources Pertcopsis eloto is in-cluded in the IDCN Red List of threatened

species as endangered, as a result of habitatloss and degradation and ongoing exploitationIt is listed in CITES Appendix H because Itmay become threatened with extinction unlesstrade is closely controlled. International trademay be authorized by the granting of an exportpermit in the country of production. In C6ted'TVoire, Ghana and Nigeria Pertcopsis eloto isseriously threatened due to forest fragmenta-tion and selective logging and in Garnero0n dueto illegal logging. The largest remaining standsare located in northern Congo and northern DR

PERTCOPsiS 425

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426 TIMBERSl

Congo, whereas there is no information on thestatus of Pertcopsis e!oto in the Central AfricanRepublic

Breeding Selection for plantation develop-merit has been undertaken in Ghana

Prospects Pertcopsis eloto produces one ofthe most highly valued timbers of tropical of-Tica. The large-scale selective exploitation hasresulted in serious reduction of populations,whereas the remaining larger populations arestill under much pressure. Although the criti-cal status of Pertcopsis eloto has been recog-nized by including it in International and na-tionallists of threatened species, protection isstill not adequate. There is still too little in-formation on growth rates, propagation andappropriate management techniques to developmodels for sustainable exploitation. More re-search is needed before measures can be taken

to allow harvesting of this species on a sris-tainable basis without threatening it with ex-tinction. Growth rates seem to be too slow to

establish economicalIy viable timber planta-tions, also because only the heartwood is highlyvalued, and the sale of material early in therotation cycle Is unattractive because thismainly consists of sapwood. In Ghana it hasbeen concluded that any future efforts to estab-lish plantations of Pertcopsis eloto must ad-dress the problem of defoliation by Loin-prosemo loterittolis. Sustainable exploitationfrom natural forest seems to be the only optionfor saving this valuable African timber for thefuture, but very long cutting cycles and thusvery low production levels may be necessary

Major references ATIBT, 1986; Bolza &Keating, 1972; Burki11, 1995; CTRAD ForestryDepartment, 2003; Farmer, 1972; G6rard at al. ,1998; Howland, 1979; Phongphaew, 2003;Siepel, Poorter & Hawthorne, 2004; Takahashi,1978.

Other references ACkah, 1997; AfricanRegional Workshop, 1998e; Agyili & Adam,1996; Ajaiyeoba at a1. , 2004; Appiah, 1994;Asare, 1994; Atuahene, 1996; Atuahene &Teyegaga, 1979; Burslem & Miller, 2001; de inMensbruge, 1966; Hawthorne, 1995; Haw-thorne & Jongkind, 2006; InsideWood, un-dated; Keay, 1989; Kyereh, Swame & Thornp-son, 1999; Neuwinger, 2000; Normand &Paquis, 1976; Sayer, Harcourt & Collins (Edi-tors), 1992; Tailfer, 1989; Toussaint at al. ,1953; Vivien & Faure, 1985.

Sources of illustration Tailfer, 1989;Toussaint at a1. , 1953

Authors L. C. N. inglaaere

PHYLLOXYLONPERRIERIDrake

Protologue Grandid. , Hist. phys. Madagascar30: 192 (1903)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution Phy!-lorylon perrieriis endemic to western Mada-gascar, where it occurs from near Analalavasouth to Morondava

Uses The wood, known as 'barahara', is tra-ditionally used for house construction, rumi-ture, agricultural implements and toolhandles,but it is also suitable for long-lasting poles andheavy-duty flooring. It is used as fuelwood andfor making charcoal. The bark serves as a fishpoison.

Properties The heartwood is pale reddishbrown and distinctly demarcated from the yel-lowish sapwood. The wood is very heavy, veryhard and elastic

Adulterations and substitutes The wood

is similar to that of other Phy!lorylon spp. andis known under the same Malagasy name. Itresembles that of Digitum unifoliolotumCapuron from eastern Madagascar, althoughthe wood of the latter species is less hard. Thewood of Humbertio ingdogoscoriensis Lain. isalso very heavy and used for similar purposes.

Description Deciduous shrub or smalltreeup to 12 in tall; hole up to 50 cm in diameter;bark pale brown to grey, flaking off in fibrouspieces; stems flattened, leathery, greyishgreen, much-branched, internodes leaf-like,linear, 10-40 cm x 2-7GIO) min, sharplypointed at apex, with buds and/or leaves alongthe margins, glabrous; older stems becomingrounded. Leaves alternate, simple and entire;

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but the other 3 have narrow andspecies,

rounded young stems.Anatomy Wood-anatointcal description OAWA

hardwood codes):Growth rings: a 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: in-tervessel pits alternate; 23: shape of alternatepits polygonal; 24: intervessel pits minute (S 4prn); 25: intervessel pits small (4-7 pin); 29vestured pits; 30: vessel-ray pits with distinctborders; similar to Intervessel pits in size andshape throughout the ray cell; 40: mean tan-gential diameter of vessellumina S 50 prn; 41mean tangential diameter of vessellumina 50-100 prn; 50:;;^ 100 vessels per square milllime-tre; 58: gums and other deposits in heartwoodvessels. Tracheids and fibres: 61: fibres with

simple to minutely bordered pits; 66: non-septate fibres present; 70: fibres very thick-walled. Axial parenchyma: 75: axial paren-chyma absent or extremely rare; 78: axial pa-renchyma scanty paratrachea1; 91: two cellsper parenchyma strand. Rays: 97: ray width I-3 cells; 104: all ray cells procumbent; 1/6:;a 12rays per mm. Stoned structure: 1/8: all raysstoned; 120: axial parenchyma and/or vesselelements stoned. Mineral Inclusions: 136

prismatic crystals present; 138: prismatlc crys-tals ln procumbent ray cells(L. N. Banak, H. Beeckman & P. E. Gasson)

Growth and development The growth isvery slow. Phy!!orylon perrieri usually flowersfrom September to November

Ecology Phylloxy!on perrieri occurs in de-ciduous forest and woodland, from sea-level upto 600 in altitude. It usually occurs on sandysoils, but can also be found on limestone

Harvesting Trees are difficult to fell withtraditional equipment because of the very hardwood

Genetic resources Phy!lorylon perrieriisclassified as an endangered species in theIUCN Red list. Although it is fairly widespreadin western Madagascar, it is uncommon be-cause of selective felling for its timber and fu-elwood, and because of severe fragmentation ofthe natural vegetation in which it occurs

Prospects The prospects for Phylloxylonperrieri and other Phyl!ory!on species as tim-her trees of commercial importance are verypoor because of the often small size of the bole,their uncommon occurrence and their slow

..

II

F.

Phy!!oxylon perrieri - I, ledh, twig, . 2, floweringtwig, . 3, twig with fruitRedrown ond adopted by Ishoh Syomsudin

stipules minute; petiole 2-6 mm long; bladeelliptical to ovate, 3-6(-7.5) cm x I-4(-5) cm,obtuse to rounded at base, acute to obtuse atapex, leathery, glabrous, pinnately veined;sometimes leaves absent. Inflorescence a few-flowered raceme produced from the buds on theleaf-like stem margins, 5-13 mm long; bractsnumerous on the axis, broadly ovate. Flowersbisexual, papilionaceous; pedice1 2-3.5 minlong; calyx shallowly cup-shaped, c. 2 min long,with short teeth, minutely brown hairy; corollapink to pale purple, with narrowly erripticalstandard c. 13 mm long, with white basal spot,wings 13-16 min long, keel 11-13 mm long;stamens 10, 9 united into a long sheath 6-7mm long and I free; ovary superior, narrowlyoblong to linear, style curved upward. Fruit acurved spindle-shaped pod 3.5-6.5 cm x 1.5-2cm, with beak at apex, leathery, eventuallydehiscent, I(-2)-seeded. Seed erripsoid, c. 16min x 11mm.

Other botanical information Phylloxyloncomprises 7 species, all endemic to Madagas-car. It belongs to the tribe Indigofereoe, inwhich it is considered to be basal because it

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PHYi, LowLON 427

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428 TIMBERSl

growth. Locally, they will remain important forthe production of durable posts and imple-merits. However, attention should be given toadequate protection

Major references du Puy at a1. , 2002; duPuy, Labat & Schrire, 1995.

Other references du Puy & Labat, 1998g;Gu6neau, Bedel & Thie1, 1970-1975; Inside-Wood, undated; Kerharo, GUIchard & Bouquet,1961

Sources of illustration du Puy, Labat &Schrire, 1995.

Authors R. H. M. J. Lemmens

PHYLLOXYLONXYLOPHYLLOIDES (Baker) DuPuy, Labat & Schrire

Protologue Kew Bull. 50(3): 489 (1995)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Synonyms Phy!104.10nensifolium Ban. (1896).Origin and geographic distribution Phyl-

lorylon xylophylloides is endemic to Madagas-car, where it mainly occurs on the upper slopesof the eastern margin of the Central Plateauxover nearly the whole length of the island.

Uses The wood, known as 'barahara' or 'ara-hara', is traditionally used for house construc-tion, furniture, agricultural implements andtool handles, but it is also suitable for long-lastlrig poles, heavy-duty flooring, turnery andnovelties. It is used as fuelwood and for makingcharcoal.

A decoction of the twigs is used in traditionalmedicine as a tonic, and to treat fever and skincomplaints. The leaves are credited with stimu-Iant, digestive and rebrifugal properties, theroots with depurative activity. The wood isused as fish poison, and a maceration of thewood is used against a poisonous water beetle('tsingala')

Properties The heartwood is dark reddishbrown and distinctly demarcated from the yel-lowish grey to brownish grey sapwood. Thegrain is straight or wavy and interlocked, tex-ture very fine.The wood is very heavy, with a density of about1200 kg/ing at 12% moisture content. The wooddries slowly, planks 25 min thick can be driedin 3-4 months under high plateau conditionswithout twisting, splitting or deformation.Shrinkage rates from green to oven dry arehigh: 6.3% radial and 11.2% tangential. Oncedried, the wood is very stable in service. Thewood is very hard and elastic. At 12% moisture

content, the modulus of rupture is about 260N/min2, modulus of elasticity 15,500 N/min2,compression parallel to grain 105 N/mm2 andChalais-Meudon side hardness 19.7

Sawing and working of the wood is difficult,and the use of stellite teeth and tungsten car-bide tools is necessary. The wood holds nailsand screws well, but pre-boring is necessaryGluing is difficult but varnishing and waxingproperties are good. The wood is very durableand extremely resistant to impregnation.

Botany Leaness shrub or smalltree up to15(-20) in tall; hole up to 35(-50) cm in diame-ter; bark pale brown, flaking off in fibrouspieces; sterns flattened, thinly leathery, glossygreen, much-branched, internodes leaf-like,elliptical to narrowly elliptical, 3-20 cm x0.5-3.5 cm, with buds along the margins, gla-brous except for the buds; older stems becoin-ing rounded. Inflorescence a few-flowered ra-ceme produced from the buds on the leaf-likestem margins, 3-8(-18) min long; bracts nu-merous on the axis, broadly ovate. Flowersbisexual, papilionaceous; pedice1 2-5 min long;calyx shallowly cup-shaped, c. 2 mm long, withshort teeth, minutely brown hairy; corolla pinkto pale purple, with elliptical standard c. 8 mmlong, with white or greenish basal spot, wingsand keelc. 10 min long; stamens 10, 9 unitedinto a long sheath c. 8 min long and I free;ovary superior, narrowly oblong to linear, stylecurved upward. Fruit a spindle-shaped pod2.5-4.5 cm x 1.5-2 cm, with beak at apex,leathery, greenish, glossy, eventually dehis-cent, I(-2)-seeded. Seed ovoid, with thin seedcoat. Seedling with epigeal germination; hypo-cotyl curved; cotyledons completely fused to ahomogeneous massPhylloxy!on xylophyl!oides shows a peculiargermination. A plumule is absent in the seedThe radicle penetrates into the soil while theseed remains at its position on top of the soil.In 3-4 months the seedling develops a spindle-shaped primary root 10-15 cm long and acurved hypocotyl. At this stage the cotyle-donous mass of the seed has been much re-

duced, and the seedling enters into a restingperiod untilthe onset of the next rainy seasonThen a leaf-like stem develops from a bud onthe hypocotyl. The growth is very slow. Phy!-lorylon xylophylloides usually flowers fromAugust to November and March to April.Phylloxylon comprises 7 species, all endemic toMadagascar. It belongs to the tribe Indigo-Iereoe, in which it is considered to be basalbecause it shows many primitive character-

.

states. The leaf-like young sterns are character-1stIC in 4 species, but the other 3 have narrowand rounded young stems.Phylloxylon orenicolo DU Puy, Labat & Schrirefrom north-eastern Madagascar is a shrub orsmalltree up to 4 in tall with hole up to 15 cmin diameter. It does not have leaf-like stemsand has obovate leaves. Its wood is heavy andextremely durable, and the stems are used forhouse posts, which may last for up to 100years. A leaf decoction is used in rituals.

Ecology Phyl!orylon xylophylloides occurs inhumid, evergreen forest, often along watercourses, usually at 600-1600 in altitude, butdescending to sea-level in the south-easternpart of Madagascar. It is found on latente andsandy soils

Management The seeds remain viable forup to 6 months. Trees are difficult to fell withtraditional equipment because of the very hardwood.

Genetic resources and breeding Phyl-lorylon xylophyl!oides is classified as unlner-able in the IUCN Red list. Although it is wide-spread in eastern Madagascar, its distributionarea is discontinuous and many larger popula-tions are threatened by forest clearing, e. g. fortitanium mines, and selective felling for itstimber. All other Phyl!orylon species are classi-fled as endangered or even criticalIy endan-gered.Phyl!orylon xylophylloides shows significantgenetic variation, particularly in the shape ofthe leaf-like stems and the length of the staini-nal sheath, which differ between northern,central and southern populations

Prospects The prospects for Phyl!orylon xy-lophylloides and other Phylloxylon species astimber trees of commercial importance are verypoor because of the often small size of the hole,their uncommon occurrence and their slow

growth. Locally, they will remain Important forthe production of durable posts and imple-merits. However, attention should be given toadequate protection

Major references Bolteau, Bolteau & A1-10rge-Bolteau, 1999; du Puy at a1. , 2002; duPuy, Labat & Schrire, 1995; Gu6neau, Bedel &Thie1, 1970-1975; Peltier, 1971

Other references Debray, Jacquemin &Razafindramba0, 1971; du Puy & Labat,1998h; Gu6neau, 1971; Kerharo, Guichard &Bouquet, 1961; Neuwinger, 2000

Authors R. H. M. J. Lemmens

PINUS CARIBAEAMorelet

Protologue Rev. Hort. C6te d'or I: 107(1851)

Family PinaceaeChromosome number2n=24

Vernacular names Caribbean pine, pitchpine, Caribbean pitch pine, Nicaragua pine,Cuban pine, Honduras pine, yellow pine (En)Pin mate, pin ianne, pin caratbe, pin desCaratbes, pin de Cuba (Fr). Msindano (Sw)

Origin and geographic distribution Pinusconboeo occurs naturally in eastern CentralAmerica, Cuba and the Bahama Islands. It isplanted throughoutthe tropics, including manycountries of tropical Africa, and in Australiaand New Zealand

Uses The wood (trade names: Caribbeanpitch pine, Honduran yellow pine) has a coin-paratively low density and exudes much resin,making it less suitable for e. g. joinery andflooring, and limiting its value as a timberNevertheless, it is used for a wide range ofpurposes, including construction, light flooring,joinery, inexpensive furniture, boxes, pallets,turnery, toys and, after treatment with pre-servatives, poles, posts, railway sleepers andmine props. Resin-soaked wood is popular forboat decking, because of its high durability.The wood is also suitable for interior trim, ve-neer, plywood, piles, vats, particle board andfibre board. It is used as fuelwood, although ittends to throw out sparks, and for the produc-tion of charcoal. The wood is also used for themanufacture of paper, but is not suitable fordissolving-grade pulp.When tapped, the tree yields a good quality ofo1eoresin which is distilled to give turpentine

PINUS 429

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Pinus cartboeo -planted

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430 TIMBERSl

and rosin. Turpentine is used in paint and ba-tik industries, and rosin is used in the produc-tion of paper, soap and glue. The o1eoresinproducts are often termed 'naval stores' be-cause of their historic use for ship caulkingPinus conboeo is planted in windbreaks and asan ornamental and shade tree. The mat ofnee-

dles on the ground is considered valuable forprotection against soil erosion. The seeds arelocally consumed

Production and international trade Pinus

curtboeo is fairly important for the productionof timber, but trade and export statistics areunavailable because the wood is often used

locally. The pulp is often mixed with that ofother species.

Properties The heartwood is yellowish toreddish brown and distinctly demarcated fromthe white or yellowish sapwood. The grain isstraight, texture medium to coarse. Growthrings are distinct. Numerous resin canals arepresent and clearly visible as straight brownstreaks on longitudinal surfaces.Wood properties show large differences be-tween sites and between trees. The wood is

moderately lightweight to fairly heavy, with adensity of 350-560(-820) kg/ing at 12% ino1s-ture content. The wood from slower-growingtrees from naturalstands has a higher densityand lower resin content than the wood from

faster-growing trees from plantations. Thewood air dries well, but end splits may occur.Boards of 30 min thick require about 6 weeksto air dry from green to 20% moisture contentConventional kiln drying is successful for both25 mm thick material(which takes 3-4 days todry to 12-14% moisture content) and 50 minthick material. High temperature drying hasproved successful for framing sizes. Boards of30 min thick of plantation-grown wood may bedried in a solar kiln to 12% moisture contentin

40 days. Rates of shrinkage from green to ovendry are 1.9-4.5(-6.3)% radial and 5.7-7.9%tangential.At 12% moisture content, the modulus of rup-ture is (50-)61-1/5 N/mm2, modulus of elastic-ity (2600-)6300-15,400 N/min2, compressionparallel to grain (22-)34-59 N/min2, shear 14N/mm2, cleavage 12 N/min radial and 13 N/mmtangential, Janka side hardness 3020-5520 Nand Janka end hardness 3740 N

The wood works and finishes well with hand

and machine tools, although resin may clogand gum up tool edges and surfaces. It takesnails and screws particularly well and gluessatisfactorily.

The wood is moderately durable, being suscep-tible to marine borer and termite attacks, butresistant to powder-post beetles and fungiResistance to insect attacks increases with

increasing resin content. The heartwood ismoderately resistant to impregnation withpreservatives; the sapwood is highly permeableand easily treated in open-tank or pressure-vacuum systems. The energy value of the woodis 20,300 kJ/kg.The wood fibres are on average 4.6 mm longand 41-52 pin wide. The wood fibres of 8-12-year-old trees from Tanzania were 2.9-3.0 mmlong, with a diameter of 40-47 prn and a cellwall thickness of 4.2-4.5 The chemicalF1mcomposition of the oven-dry wood was: bolocel-Iulose 64-65%, or-cellulose 40-42% and Iignin31%. The solubility in cold water was 1.8-2.6%,in hot water 2.1-3.3%, in alcohol-benzene 1.2-1.3% and in I% NaOH 11.6-12.8%. Pulpingwith the sulphate (kraft) process yielded 41-46% screened pulp, with a kappa number of23-48, satisfactory tensile and burstingstrengths, but relatively low tearing strength.The o1eoresin, which is found in the intercellu-Iar canals in especially the sapwood, is a paleyellow, clear and sticky mass, becoming brittleon evaporation. It is a hydrophobic substancesoluble in neutral, non-polar organic solventssuch as dry ethyl ether, hexane, and other pe-troleum solvents. On distillation, it producesgum rosin and gum turpentine in a ratio of 4-6:1. Gum rosin is a brittle solid, insoluble inwater, but soluble in many organic solvents. Itconsists primarily of a mixture of abietic andpimaric-type acids. Gum turpentine is a liquidmixture of mainly terpene hydrocarbons andterpenoids, including (+)-u-pinene, (-)-orpineneand (-)13-phellandrene. From the bark about10% tannin can be extracted; it can be dried toa reddish powder soluble in water. Hexaneextracts of the bark have shown strong anti-fungal and antibacterial activity, and may havepotential for control of fungi and bacteria inthe pulp and paper Industry

Description Evergreen, monoecious, medium-sized tree up to 30(-45) in tall, but in planta-tions usually much smaller; hole branchless forup to 21 in, up to 80(-135) cm in diameter,usually straight, cylindrical; bark surface red-dish brown to pale brownish grey, deeply fis-sured, lriner bark very resinous; crown thin,rounded to pyramidal, slightly spreading; twlgsorange brown, later turning grey-brownLeaves in bundles of (2-)3(-5) in whorls at theend of the shoots, needle-shaped, 15-25 cm

.

hondurensis is generally grown; It is the mostsuitable variety for tropical lowlands. In SouthAfrica it is recorded to have more crookedstems than the other 2 varieties. Var. 60-homensis is reported to have some tolerance toshoot moth attack. Pinus curtboeo var. hon-durensis forms natural hybrids with Pinuso0corpo Schiede ex Schltdl. Pinus curtbueo hasoften been mistaken for Pinus elliottii Engelmin the past

Anatomy Wood-anatoimcal description CAWAsoftwood codes)Growth rings: 40: growth ring boundaries dis-tinct; (42: transition from earlywood to late-wood abrupt);(43: transition from earlywood tolatewood gradual). Tracheids: (44: tracheidpitting in radial walls (predominantly) uniseri-ate (ea"Iywood only)); (45: tracheid pitting inradial walls coredominantly) z 2-senate (ear-Iywood only)); 46: tracheid pits in radial wallsoppositeIy arranged (earlywood only); 55: late-wood tracheids thick-walled (double wallthickness larger than radial lumen diameter);56: torus present (pits in earlywood tracheidsonly). Ray composition: 79: ray tracheids coin-monly present; 82: cell walls of ray tracheidsdentate; 83: cell walls of ray tracheids reticu-late; 85: end walls of ray parenchyma cellssmooth (unpitted); 87: horizontal walls of rayparenchyma cells smooth (unpitted). Cross-field pitting: 91: cross-field pits pinoid; 98: I-3pits per cross-field (earlywood only). Ray size:103: average ray height medium (5-15 cells);107: ray width exclusively uniseriate. Intercel-Iular canals: 109: axial intercellular (resin)canals present; 110: radial intercellular (resin)canals present; 1/7: epithelial cells thin-walled.

(P. Baas & I. Hemz)Growth and development Pinus conboeo

usually grows rapidly. In Nigeria 6-year-oldtrees were 7-8.5 in tall, and 11-year-old trees17 in. In trials in Rwanda 13-year-old treeswere 17 in tall, with a hole diameter of 18.7 cm.In Malawi 9-year-old trees (density 670trees/ha) were on average 19 in tall. In SouthAfrica (70 in altitude, mean annual tempera-ture 22'C, mean annual rainfall 965 mm) 34-year-old trees reached a height of 27 in and ahole diameter of 47 cm. Bole straightness gen-erally improves from the subtropics towardsthe tropics. Fairly often 'foxtails' occur, plantswithout branching and without growth rings inthe wood. Foxtailing is a reaction to off siteplantingIn southern Africa female flowering starts

Pinus cartboeo - I, tree hubit, . 2, bundle of ledues, .3, twig with mature fernole cone.Redrawn grid udopted by Achingd Sotzr, Nur-hornon

long, minutely toothed, stiff, dark or yellowlshgreen, slightly shiny. Male cone in dense clus-ters, 2-4 cm x 0.5 cm, red-brown. Mature fe-male cone solitary or in groups of 2-5, on apeduncle I-2 cm long, ovoid, 4-14 cm x 2.5-4cm, reddish brown, with reflexed or wLdespreading scales. Seeds narrowly ovoid, up to 6min x 3 mm, with a persistent membranouswing up to 20 mm long, black to inottled greyor pale brown. Seedling with hypogeal germi-nation.

Other botanical information Pinus is a

large genus comprising over 110 species, al-most allrestricted to the northern hemisphereMany Pinus species are cultivated outside theirnatural distribution area, in tropical, subtropi-cal and temperate regions. In the tropics 2 spe-cies are more important than all others: Pinuscartboeu in the lowland humid tropics andPinus potulo Schltdl. & Chain. in the coolerhighland tropics and subtropicsPinus conboeo is often divided into 3 varietles

60homensis (Griseb. ) W. H. GBarrett &Golfari, var. cartboeo and var. hondurensis(Sen6cl. ) W. H. GBarrett & Golfari. In planta-tions outside the native area of the species var

PINUS 431

3

2

var

432 TIMBERSl

when trees are 3-4 years old, but male flower-ing starts later. Pollination is by wind. Thetime between pollination and ripening of thefemale cones is 18-21 months. Cones are read-

11y shed from the branches, but sometimes per-SIStent for over one year. Seed dispersal is bywind, but sometimes birds, rodents and people,who gather the seeds for food, also dispersethem. Outside its native area Pinus curiboeorarely regenerates naturally

Ecology Pinus curtboeo is mostly grown upto 1000(-1500) in altitude, in areas with amean annual temperature of 20-27"C, a meanmaximum temperature of the warmest monthof 28-34'C, a mean minimum temperature ofthe coldest month of 8-23"C, an average an-nual "amfall of (650-)1000-3000(-4000) mm,and a dry season of up to 6 months. It is inod-erately drought resistant, but does not toleratefrost. The tree is moderately tolerant to wind,also to salt wind, and it may be planted nearthe coast. Pinus coriboeo grows on a wide van-ety of soils, but does best on well-drained, deep,fertile soils with pH 5-5.5. It tolerates season-ally waterlogged soils. Young trees are highlysusceptible to fire damage, but older trees aremoderately fire resistant. Pinus curtboeo isstrongly light-demanding.

Propagation and planting Pinus cartboeoIs easily propagated from seed. The 1000-seedweight is 12-33 g. To obtain seed, cones can becollected as soon as they begin to change fromgreen to brown on the tree; cones collected ear-lier may give seeds with short viahinty. Seedsare collected after sun-drying the cones in thesun or in open-sided sheds covered with plasticroofs. Seeds can be stored for up to 10 yearsunder dry (below 10% humidity), cold (0-10'C)and airtight conditions. Pre-treatment beforesowing is unnecessary, butsoaking in water for12-48 hours gives more uniform germinationGermination takes 8-21(-42) days, and up to80% germination is common. Mycorrhizae arenecessary for seedling growth, so it is recoin-mended to inoculate with spores or to add soilfrom near established trees. Seedlings aresuitable for planting out after 4-8 months,when they are 20-30 cm tall. The planting siteshould be thoroughly cleaned. Initial spacingsare 2-5 in x 2-5 in, depending on productionalms. with pulpwood plantations having theclosest spacing. For resin production the rec-Dinmended spacing is 4 in x 4 in. Direct sowingis uncommon. Vegetative propagation is possi-ble using stem cuttings, grafting, air-layeringor tissue culture

Management Weeding is recommended dur-ing the early years, also to decrease the risk offire. Pruning is recommended to reduce therisk of fire, improve access, improve tree formand reduce the size and frequency of knots. Inplantations for sawn wood, veneer and largeposts, trees are planted at an initial density of1100 trees/}Ia, pruned during the first years,and later thinned to a final density of 250-400trees/ha, with rotations of 15-25 years, Inpulpwood plantations there may be a singlethinning at 3-4 years to remove inalformedtrees or no thinning at all. Where both timberand pulpwood are produced, plantations can beheavily thinned when 10 years old to obtainpulpwood, with a more open plantation remain-ing for the production of timber

Diseases and pests Needle blightcaused byCercosporo pint-densi/10roe can seriously at-tack plantations. Damage by Armillorio meneohas been recorded in Tanzania, Malawi andMauritius. In nurseries damping off may occurPinus conbueu is resistant to pitch canker(Fusorium circinutttm), a diseaseserious

threatening pine plantations in South AfricaHarvesting To obtain the resin various

methods are practised. The first tapping orwounding of a living tree, at 30-40 cm abovethe ground, is about 1.3 cm wide and 30 cmlong and is followed by a series of tappingsuntilbreast height is reached. A sulphuric acidsolution (usually 40-60%) is applied jinmedi-ately after tapping. This process yields the'gum naval stores' which is the major source ofthe world's supply of rosin and turpentine'Wood naval stores' can be obtained by solventextraction of stumps of old trees; the resin ob-tained in this manner is less pure. Turpentineand wood resin can also be obtained as by-products from the kraft or sulphate pulping ofpines. Turpentine is removed from the chipdigester during the initialsteaming and is con-densed from the reliefgases. This method gives'kraft naval stores'

Yield Mean annual volume increments are

10-40 in3ftia. In Malawi 9-year-old trees (den-sity 670 treesA1a) yielded 179 msftia. Becausebranching is light, the yield of first-gradestructural timber is exceptionally high.

Handling after harvest Freshly felled logsexude copious amounts of resin. Although notnumerous, knots can cause degrade becausethey are large and tend to cause splitting ondryingFelled logs discolour easily and should be con-verted and dried rapidly to avoid blue stain

.

Seasoned boards should be stored under cover

Genetic resources Genetic linkage maps ofPinus curtboeo var. hondurensis have been

made using AFLP and microsatellite markers.Breeding Provenance trials have been con-

ducted in over 50 countries, including Gambia,Sierra Leone, C6te d'TVoire, Nigeria, Congo,Sudan, Kenya, Uganda, Tanzania, Malawi,Zambia, Zimbabwe, South Africa and Mada-

with var. hondurensis consistentlygascar,

showing the fastest growth. Clonal material ofabout 150 'plus trees' has been distributed bythe Oxford Forestry Institute, and clone bankswere established in Zimbabwe, Australia andBrazil. A hybrid of Pinus conboeu var. hon-durensis and Pinus enjottii has become impor-tant in plantations in Queensland (Australia)because of its superior growth and shape.

Prospects Pinuscoriboeoiscommonly plantedin the tropics because of its fast growth, abilityto grow on virtually all soil types, the detailedinformation available on Its SIIviculture, andthe suitability of the wood for a wide range ofapplications. It has become the most importantpine of the lowland humid tropics. However,the quality of the wood is often not high be-cause of the high resin content, and improve-merit of the wood quality should be a researchpriority.

Major references FAO, 1974; Katende,BITnie & Tengnas, 1995; Lamprecht, 1989;Nieto & Rodriguez, 2002; Suhardi at a1. , 1993;Takahashi, 1978; van Wyk, 2002a; Webb at al. ,1984; World Agroforestry Centre, undated.

Other references Bryce, 1967; Chauvet,1968; Evans, 2003; Fanon, 1984; Foot, 1967;Guti6rrez Gotera at a1. , 2004; HeInz, 2004;11vessalo-Pmffti, 1995; Lavers, 1969; Little,undated; AJIarais, 1997b; Mugunga. & van Wyk,2003; Palmer, Ganguli & Gibbs, 1984; Parant,Chichignoud & Curie, undated; Poynton, 1966;Shepherd at a1. , 2003; Sutter, 1990; vanVuuren, Banks & Stohr, 1978

Sources of illustration Fanon, 1984Authors A. A. Oteng-Amoako & M. Brink

Origin and geographic distribution Pinuserrtottii occurs naturally in the south-easternUnited States (Florida and adjacent coastalplains). It has been introduced into many partsof the southern hemisphere, including Africa,South America, Australia and New Zealand. InAfrica it is recorded from Burundi, Tanzania,Malawi, Zambia, Zimbabwe, Madagascar,Mauritius, Reunion, South Africa and Swazi-land. It also occurs as an escape.

Uses The lightweight and soft wood fromyounger trees is mainly used for pulping,whereas the heavier and harder wood of oldertrees is used as timber. The straightness of thehole makes it particularly suitable for poles,piles and solid-wood products, but treatmentwith preservatives is often necessary. Thewood is also suitable for construction, flooring,frames, joinery, interior trim, furniture, cabi-net work, ship and boat building, vehicle food-ies, toys, turnery, boxes, crates, veneer, ply-wood and particle board. It is used as fuelwood.01eoresin is tapped from the trees and distilledto obtain turpentine and rosin. Turpentine Isused in the paint industry, and rosin is used inthe production of paper, soap and glue.

Properties The heartwood is yellow to red-brown, and notclearly demarcated from the 5(-15) cm wide yellowish white sapwood. Thegrain is straight, sometimes spiral; texturemedium. Growth rings are distinct. The resincontent of the wood is high. The wood has adensity of 420-700 kg/m' at 12% moisture con-tent. It dries with little degrade, although somesurface checking and splitting down the centremay occur. The rates of shrinkage from greento oven dry are 4.2-5.5% radial and 7.9-8.5%tangential. At 12% moisture content, themodulus of rupture is 71-123 N/min2, modulusof elasticity 7800-14,700 N/mm2, compressionparallel to grain 37-59 N/mm2, compressionperpendicular to grain 6 N/min2, shear $9N/mm2 Janka side hardness 2930-4630 N andJanka end hardness 3800-5920 NThe wood is relatively difficult to work. Resinmay adhere to sawteeth and cutting edges, butthe use of long-pitched sawteeth reduces thisproblem. The wood holds nails and screws well,and it glues, finishes and paints satisfactorily.It is moderately durable to nori-durable, beingsusceptible to attacks by Ano61um borers, ina-rine borers and termites. The sapwood is notsusceptible to Lyetus borers. The heartwood isresistant to impregnation with preservatives,the sapwood is permeableThe wood fibre cells are 2.1-4.0 min long, with

PINUS 433

PINUS ELLIOTTll Engelm.

Protologue Trans. ACad. Sci. St. Louis 4(I):186-190, PI. I-3 (1880).

Family PinaceaeChromosome number2n=24

Vernacular names Slash pine, pitch pine,yellow pine (En). Pin d'Enjott, pin a aiguilleslongues (Fr)

.

434 TIMBERSl

a diameter of 39-54 prn and a cell wanthick-ness of 4.2-6.5 pin. The chemical compositionof the oven-dry wood is: holocellulose 57-73%,orcellulose 36-42% and Iignin 28-32%. Thesolubility in cold water is 1.4-3.3%, in hot wa-ter 1.9-4.7%, in alcohol-benzene 0.5-6.6% andin I% NaOH 8.5-15.8%. Pulping with the SUI-phate (kraft) process yields 40-51% screenedpulp, with a kappa number of 23-56, givingpaper with satisfactory tensile and burstingstrengths, but relatively low tearing strengthIn Zimbabwe the o1eoresin yields 12-13% tur-pentine, with as main components u-pinene(54.3%) and p-pinene (34.0%). The main coin-ponents of essential oil distilled from the nee-dles of Pinus enjottiifrom Mozambique were orpinene (43.0%), ^-pinene (27.1%) and orterpine01(9.6%).

Botany Evergreen, monoecious, medium-sizedtree up to 30(-40) in tall; hole slender, straight,up to 90(-100) cm in diameter; outer bark grey-Ish and furrowed in young trees, laterbrownish red with a plate-like pattern; crownovoid. Leaves crowded towards the ends of

branches, in bundles of 2-3, needle-shaped,17-30 cm long, stiff, dark green. Male cone indense clusters, up to 6 cm long. Mature femalecone usually solitary, sessile, ovoid-conical, 7-15 cm long, greyish or reddish brown, lustrous,with scales with a blunt greyish prickle. Seedsc. 6 min x 3 min, with wings c. 25 mm long,dark brown inottled with black. Seedling withepigealgermination.Initial growth of Pinus elliottiiis fast, but overthe whole rotation cycle growth is not as fast asthat of Pinus potulo Schltdl. & Chain. In Zaru-bia 30-year-old trees were on average 24 in tallIn Malawi 30-year-old trees in a thinned plan-tation with a density of 296 treesA1a were 30 intall, with a mean hole diameter of 37 cm. InMadagascar 40 year-old-trees were on average31 in tall with a bole diameter of 41 cm. In

South Africa 20-year-old trees (density 320treesA1a) were 23 in tall, with a hole diameterof 22 cm, and 33-year-old trees (density 320treesA1a) 33 in tall with a hole diameter of 42cm. Pinus enjottii is notable for the straight-ness of the hole. Self-pruning is common, re-SUIting in a relatively short crown. Pollinationis by wind. Female cones mature in 3 years,and trees start bearing seed when 7-8 yearsold

Pinus is a large genus comprising over 110species, almost all restricted to the northernhemisphere. Many Pinus species are cultivatedoutside their natural distribution area, In

tropical, subtropical and temperate regions. Inthe tropics 2 species are of outstanding impor-tance: Pinus curtboeo Morelet in the lowland

humid tropics and Pinus potulo in the coolerhighland tropics and subtropics. Pinus enjottiiis often confused with Pinus cartboeo, and the2 species hybridize

Ecology Pinus enjottiiis grown at (500,700-2500 in altitude, in areas with a mean annualtemperature of 15-24'C, a mean maximumtemperature of the warmest month of 23-32'C,a mean minimum temperature of the coldestmonth of 4-12'C, an average annual rainfall of650-2500 mm, and a dry season of I-4 monthsIt can be grown on a wide variety of soils, butdoes best on deep, well-draining, acid soilsPinus enjottiiis fairly tolerant to frost and saltwinds. Young trees are quite susceptible to fireinjury until they are 3-4.5 in tall, but then thebark becomes thick enough to insulate thecambium from high temperatures. It is a light-demanding species, competing wellwith weeds

Management Pinus enjottiiis usually pro-pagated by seed. The 1000-seed weight is 25-50 g. In Zimbabwe seeds are collected aftersun-drying the cones in open-sided sheds cov-ered with plastic roofs, sometimes supple-merited by kiln-drying at a maximum tempera-ture of 48'C. The seeds can be stored for yearsunder dry, cold and airtight conditions. Germi-nation normally takes 15-20 days, and thegermination rate of fresh seeds is usually 80-95%. The presence of mycorrhizae is highlybeneficial for survival and early growth, so it isrecommended to inoculate with spores or toadd soilfrom near established trees. Seedlingscan be planted out 4-8 months after germina-tion, when they are about 30 cm tall. Commonspacings are 2.5 in x 2.5 in. Vegetative propa-gation is possible using grafting, air-layering orcuttings, but branch cuttings do not root easily,especially those from older trees. In-vitro re-generation systems have been developed usingembryogenesis or organogenesis.Weed controlis necessary during the first 2years after sowing. There are no conclusiveexperiments of fertilizer use. Rotation cyclesdepend on production alms: the optimum rota-tion for pulpwood is about 25 years, whereasfor sawn timber longer rotations are applied,e. g. 45-55 years to obtain trees with a bolediameter of about 40 cm. Thinning is normallypractised. In plantations in Malawi with aninitial density of 1250 treesA1a the first thin-ning takes place when the mean bole diameterof the trees is 24 cm, so that the wood offelled

.

trees can be commercialised; this is usuallywhen the trees are about 13 years old. Thesecond thinning is carried out when the treesare 19 years old, and the third when they are25 years old, with a final density of 270treesA1a. For the production of sawn woodprunings are required at 5-10-year Intervals,starting at age 8-12, but even for pulp produc-tion pruning to a height up to 2 in Is recoin-mended when the trees are 6 in tall, to reducefire hazard. Pinus e!nottiiis reported to coppicewell.

Of all pines, Pinus e!Itottii is considered themost resistant to the Tinportant fungal pinepathogen Sphoeropsis soptneo (synonym: Dip-10dio pined) causing Sphaeropsis blight or die-back. The most serious disease is fusiform rustcaused by Cronortium lustforum. In MalawiPinus enjottiiis susceptible to honey fungus(Armi!loner sp. ), particularly below 1300 inaltitude. In nurseries damping off may occurPests include the grasshopper Mecostibuspintuoro in Zimbabwe. Pinus enjottiiis alsoattacked by the southern pine bark beetle(Dendroctonus frontolis).Due to its slower growth, wood yields of Pinuse!nottii are lower than those of Pinus potu!aHowever, Pinus e!!jotti can produce higherpulpwood yields because of its higher wooddensity. Mean annual volume increments inplantations are usually 10-20 ing/ha, but forMalawi annual increments of up to 36 mathahave been recorded. annual o1eoresin yields of3 kg/tree have been recorded from Zimbabwe.

Genetic resources and breeding Pinusel!jottii has been crossed with Pinus conboeq,Pinus potulu and other Pinus spp. A hybrid ofPinus el!jottii and Pinus conboeo var. hon-durensis has become important in plantationsin Queensland (Australia) because of its supe-nor growth and shape. Geneticlinkage maps ofPinus enjottii have been made using RAPD,APLP and microsatellite markers.

Prospects Pinus enjottiiis a useful plantationtree for the production of poles, piles and solid-wood products, and for paper making. It toIer-ates a wide range of climates and soils, and isrelatively resistant to the Tinportant pinepathogen Sphereropsis sopineo. Drawbacks areits relatively slow growth and the low durabil-ity of its wood.

Major references Barnett, 2002; Bolza &Keating, 1972; Lamprecht, 1989; Takahashi,1978; Theron at a1. , 1971

Other references Banks & Schoeman,1963; Bushy, 1982; Foot, 1967; Morris at al. ,

1997; Fagula & Baeckstr6m, 2006; Palmer &Gibbs, 1974; Palmer, Ganguli & Gibbs, 1984;Parant, Chichignoud & Curie, undated; Shep-herd at a1. , 2003; Tang, Newton & Charles,2006.

Authors M. Brink

PINUS KESIYA Royle ex Gordon

Protologue Gard. Mag. 16: 8 (1840).Family PinaceaeSynonyms Pinusinsu!ons End1. (1847), Pinus

hhosyo Hook. f. (1888)Vernacular names Khasya pine, henguet

pine, khasi pine (En). Pin a trois feuilles, pind'Indochine (F")

Origin and geographic distribution Pinushestyo is native to South-East Asia. It isplanted throughoutthe tropics and has becomea very important timber species, especially insouthern Africa, for instance in Zambia, whereit is the most widely planted pine.

Uses The wood of Pinus foesiyo (trade name:khasi pine) is used for construction, boxes,flooring, ceilings, panelling, Joinery, furniture,poles and mine props. It is also suitable forship and boat building, agricultural imple-merits, turnery, veneer, plywood and railwaysleepers. It is used for the manufacture of high-quality particle board, and its use as a pulp-wood is increasing. The wood is used as fuel-wood, for the production of charcoal and fortorches.

01eoresin of good quality Is tapped from thetrees. The o1eoresin is distilled to give turpen-tine and rosin, Turpentine is used in the paintindustry, and rosin in the production of paper,

PINUS 435

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Pinus foesiyo-planted

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soap and glue. Pinus hestyo is planted as anornamental.

Properties The heartwood is red-yellow tored-brown, darkening upon exposure, and usu-ally clearly demarcated from the 3.5-5 cm widepale yellow sapwood. The grain is straight,texture medium to fairly coarse and unevenGrowth rings are distinct. The wood containsnumerous resin canals, often visible as straightbrown scratches on longitudinal surfacesThe density of the wood is 400-750 kg/in3 at12% moisture content. The wood air dries well,without serious degrade when it is well piledand closely stacked. Kiln drying is easy, but itis advised to use a mild schedule, as fast dryingat high temperature may result in serioussplitting and excessive resin exudation. Therates of shrinkage from green to oven dry are2.3-7.0% radial and 5.5-11.5% tangential.Once dried, the wood is moderately stable inservice.

At 12% moisture content, the modulus of rup-ture Is 73-204 N/mm2, modulus of elasticity5700-20,700 N/mm2, compression parallel tograin 32-78 N/min2, shear 5-13 N/min2 cleav-age 10-19 N/mm, Janka side hardness 2220-4230 N, Janka end hardness 2120-3880 N andChalais-Meudon side hardness 1.0-6.2.The wood saws easily and can be worked to asmooth surface with all tools, but the highresin content may cause dulling of tool edgesThe nailing and gluing properties are good, andthe wood takes paints and varnishes well. It iseasy to cutinto smooth, tight veneer of uniformthickness at a cutting temperature of 50-70'C.During drying the veneer shows slight to inod-erate shrinkage and warping, and is usuallysplit free. To obtain an acceptable quality ofveneer it is often necessary to patch or fillim-perfections in the wood due to the presence ofknots and localized raised grain, and then tosand the surface.

The wood is only moderately durable. It is sus-ceptible to sapstain and attacks by pinholeborers, marine borers, wood wasps and ter-mites. The sapwood is not susceptible to Lyetusborers. The heartwood is moderately resistantto Impregnation with preservatives, the sap-wood is permeableThe wood is suitable for mechanical andchemical pulping. The wood fibres are 1.6-3.7min long, with a diameter of 44-62 prn and acell wall thickness of 4.3-7.2 pin. The chemicalcomposition of the oven-dry wood was: bolocel-Iulose 57-65%, u-cellulose 37-43% and Iignin28%. The solubility in hot water is 2.8%, in

alcohol-benzene 0.8-2.8% and in I% NaOH

10.9-16.1%. Pulping of material from Zambiawith the sulphate (kraft) process yielded 42-49% screened pulp, with a kappa number of28-48. The energy value of the wood is about23,160 kJ/kg

Description Evergreen, monoecious, largetree up to 45 in tall; bole branchless for up to20 in, straight, cylindrical, up to 140 cm in di-ameter; bark up to 4.5 cm thick, outer barkpinkish to reddish grey, reticulately and deeplyfissured; crown ovoid in young trees, flattenedor rounded in older ones; branches spreading,branchlets often with a waxy bloom. Leaves inbundles of (2-)3(-4), needle-shaped, (10-)12-21(-25) cm long, erect, flexible, soft, sharp-pointed, bright green. Male cone cylindrical tooblong, 3.5-5 cm x 0.5 cm, bright yellow or palebrown. Mature female cone up to 3 together,sessile or on a short stalk up to I cm long, pen-dulous, ovoid to ovoid-conical, (4-)5-8(-10) cmx 4-5 cm, shiny; with woody scales having atiny prickle. Seeds narrow, 1.5-2.5 cm long,short-winged. Seedling with hypogeal germina-tion

Other botanical information Pinus is a

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Pinus hestyo - I, tree hubit, 2, legb, twig, ' 3, bun-dle o11eoues, . 4, mature fernole coneSource. . PROSEA

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large genus comprising over 110 species, al-most all restricted to the northern hemisphereMany Pinus species are cultivated outside theirnatural distribution area, in tropical, subtropi-cal and temperate regions. In the tropics 2 spe-cies are of outstanding importance: Pinus curt-60eo Morelet in the lowland humid tropics andPinus potu!o Schltdl. & Chain. in the coolerhighland tropics and subtropicsThe taxonomy of Pinus hestyo is still open todebate. The inclusion of Pinus irisuloris Endl.into Pinus foesiyo has been disputed, because oftheir different field characteristics and prod-ucts, and some authors contend that Pinusfoesiyo has not been properly described

Anatomy Wood-anatonitcal description OAWAsoftwood codes)Growth rings: (40: growth ring boundaries dis-tinct); 41: growth ring boundaries indistinct orabsent; 43: transition from earlywood to late-wood gradual. Tracheids: 44: tracheid pittingin radial walls (predominantly) uniseriate (ear-Iywood only); 54: latewood tracheids thin-walled (double wanthickness less than radiallumen diameter); 56: torus present (pits inearlywood tracheids only). Ray composition: 79ray tracheids commonly present; 82: cell wallsof ray tracheids dentate; 85: end walls of rayparenchyma cells smooth (unpitted); 87: hori-zorital walls of ray parenchyma cells smooth(unpitted). Cross-field pitting: 90: cross-fieldpits window-like (fenestriform); (91: cross-fieldpits pinoid); 97: I-2 (large, window-like) pitsper cross-field (earlywood only); 98: I-3 pitsper cross-field (earlywood only). Ray size: 103:average ray height medium (5-15 ceUs); 107:ray width exclusively uniseriate. Intercellularcanals: 109: axial intercellular (resin) canalspresent; 110: radial intercellular (resin) canalspresent; (1/1: traumatic (resin) canals pre-sent); 1/7: epithelial cells thin-walledre. Baas & I. Hemz)

Growth and development Pinus foesiyogrows fairly fast. In Madagascar 38-year-oldtrees were 46 in tall. Pollination is by wind.Cones take about 23 months to mature. In

plantations trees start bearing seed when 5-7years old. Pinus hestyo generally bears seedsabundantly every year. Seed dispersal is bywind, but sometimes also by birds, rodents orpeople.

Ecology Pinus foesiyu is grown at (300-)600-1800(-3000) in altitude, in areas with a meanannual temperature of 14-23'C,maximum temperature of the warmest monthof 20-37'C, a mean minimum temperature of

the coldest month of 2-18'C, an average an-nual rainfall of 700-2200 mm, and a dry sea-son of I-7 months. Pinus foesiyo grows on arange of soil types, but prefers well-drained,neutral to acid soils. Once established the treeis fairly resistant to drought and frost. The treeis susceptible to fire damage during earlygrowth stages. Pinus hestyo is a light-demanding pioneer species colonizing areasdestroyed by fire or degraded by shifting culti-vation.

Propagation and planting Pinus foesiyo Isusually propagated by seed. The 1000-seedweight is 14-20 g. Seeds can be stored for sev-eral years under dry, cool and airtight condi-tions. Pre-treatment before sowing Is urineces-sary. The germination rate of fresh seed isabout 95%, usually in 8-20 days. Mycorrhizaeare necessary for seedling growth, so it Is rec-ommended to inoculate with spores or to addsoil from near established trees. Seedlings aresuitable for planting out after 4-7 months,when they are 20-30 cm tall. Normal spacingsare 1.5-3 in x 1.5-3 in for timber productionand 4 in x 4 in for o1eoresin production. In In-dustrialplantations in Zambia site preparatlonfor planting consists of removal of the existingvegetation, destumping, filling of holes, plough-ing to a depth of 25 cm and discing. Vegetativepropagation is possible by cuttings, grafting orair-layering. In-vitro regeneration methodshave also been developed.

Management Weeding is very importantforestablishment, and in industrial plantations inZambia it is done 8 times during the first 3years after planting out. Fertilizer may beadded according to specific site requirements.In the Mangoro valley in Madagascar, for In-stance, Zn is deficient. Plantations are usuallythinned. A schedule for Zambia is to thin to740 treesA1a at an age of 6 years, to 495treesA1a at an age of 9 years, to 300 treesA1a atan age of 12 years and to 185 treesA1a at an ageof 21 years. In Madagascar 12-year-old treeswere thinned from 1300 trees/ha to 300ftia, and

6 years later trees in thinned plots showed amuch higher diameter growth than those inunthinned plots, whereas densities of the woodand shrinkage rates were similar. Pruning isnecessary to produce high-grade timber, andstarts when the tree is 6-7 in tall. Dependingon production aims, rotations of 18-35 yearsare applied

Diseases and pests Pinus foesiyo Is suscep-tible to Dothistroma needle blight (Mycosyhoe-rello pint). In Malawi damage by Armillorio

PINUS 437

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meneo has been recorded. In nurseries damp-ing off may occur. Pests include the grasshop-per Mecostibuspiniuoro in Zimbabwe. In SouthAfrica Pinus hestyo is heavily attacked by thepine woolly aphid (Pineuspini)

Harvesting Pinus foesiyo takes 25-30 yearsbefore it can be felled for timber

Yield Mean annual increments are 10-30ms/ha. On good sites in Zambia trees reached amaximum annual increment of 40 matha at age18.

Handling after harvest The wood must beconverted or treated soon after felling, andopen-stacked under shade cover.

Genetic resources Substantial provenancevariation has been recorded, indicating poten-tial for selection and breeding. Provenancetrials have been recorded in Nigeria, Kenya,Malawi, Zambia, Zimbabwe, Madagascar andSouth Africa, and seed orchards have beenestablished in various African countries, in-cluding Malawiand Madagascar.

Breeding An important breeding objectiveis straightness of the hole. Hybridization ispossible with Pinus merhusii Jungh. & deVriese and Pinus o0corpo Schiede ex Schltdl. Aprotocol has been developed for the genetictransformation of embryogenic tissue, usingbiolistic transfer

Prospects Pinus hestyo is a fairly fast-grow-ing tree, and a useful source of timber andpulpwood. For tropical Africa it has best pros-pects at 800-1200 in altitude. Below 800 in itsyield is lower than that of Pinus curtboeo andPinus Docorpo, whereas above 1200 in Pinuspotu!o gives higher yields. As it rapidly colo-nizes degraded areas, Pinus heslyer may becomeInvasive when Introduced.

Major references Bolza & Keating, 1972;Chilufya & Tengnas, 1996; CTRAD ForestryDepartment, 2003; Gu6neau, 1963; Lamprecht,1989; Suhardi at a1. , 1993; Takahashi, 1978;van Wyk, 2002b; Webb at a1. , 1984; WorldAgroforestry Centre, undated

Other references Armitage & Buney (Edi-tors), 1990; Bouillet & Lefevre, 1996; Bouillet& Rakotova0, 1994; Chudnoff, 1980; Deb &Tandon, 2002; FAO, 1974; Hemz, 2004; 11ves-salo-Pmffli, 1995; Malabadi & Nataraja, 2007;Nandwani, Kumaria & Tandon, 2001; Palmer& Gibbs, 1969; Palmer & Gibbs, 1977; Parant,Chichignoud & Curie, undated; Parant, Chichig-noud & Rakotova0, 1985; Prasad, Prasad &Sharma, 2002; Rampanana at a1. , 1986; Salle-nave, 1955; Schmitt, Bouillet & Ratsly, 1995;Sutter, 1990; Willan, 1985

Sources of illustration Suhardi at al. ,1993.

Authors NyunaiNyemb

PINUS 00CARPA Schiede ex Schltdl

Protologue Linnaea 12: 491 (1838).Family PinaceaeVernacular names Ocote pine, Nicaraguan

pitch pine, o0carpa pine, egg-cone pine (En)Origin and geographic distribution The

natural distribution of Pinus o0co, :po is in Cen-tralAinerica, from Mexico to Nicaragua. It hasbeen introduced into tropical and subtropicalregions in Africa, Asia and South America,especially in Brazil. In Africa it is grown incountries along the western coast, from SierraLeone to Angola, in East Africa (Ethiopia,Kenya, Uganda, Tanzania), in southern Africaonalawi, Zambia, South Africa) and in Mada-gascar

Uses The wood (trade names: Caribbeanpitch pine, Nicaraguan pitch pine)is less proneto splitting and warping than that of mostother pines and therefore highly valued for theproduction of sawn timber. It is used for con-struction, boxes, poles, posts, handles, popsiclesticks, railway sleepers and plywood. It is alsosuitable for framing, flooring, Joinery, piles andparticle board. The wood is furthermore usedas pulpwood, fuelwood and for the productionof charcoal. 01eoresin, obtained from the bark,and firewood are the main products in CentralAmerica. Pinus o0cu, po has been planted as anornamental

Production and international trade Both

Pinus o0corpo and Pinus curtboeo Morelet aretraded under the name 'Caribbean pitch pine'In the 1970s the area planted worldwide withPinus o0corpo was estimated at about 23,400ha; in 2002 the total planted area was esti-mated at several hundred thousand ha

Properties The heartwood is yenowish brownto reddish brown, and distinctly demarcatedfrom the pale yellowish brown sapwood. Thegrain is straight, texture fairly fine and even.Growth rings are distinct. The density of thewood is 440-660 kg/ms at 12% moisture con-tent. The wood air dries well. The rates of

shrinkage from green to oven dry are 3.5-5.2%radial and 6.2-9.0% tangential. At 12% ino1s-ture content, the modulus of rupture is 90-123N/min2, modulus of elasticity 6600-15,500N/min2, compression parallel to grain 53N/mm2 and Janka side hardness 4050 N.

.

The wood saws and works easily with hand andmachine tools. It is only moderately durable. Itis highly resistant to white rot fungus, but onlymoderately resistant to brown rot. It does notweather well without being covered with paintor other coatings. The heartwood is resistant toimpregnation with preservatives, the sapwoodis permeable.Fibre cells in wood from Uganda were on aver-age 3 min long, with a diameter of 41 prn and acell wanthickness of 4 prn. The chemical coin-position of the oven-dry wood was: bolocellu-lose 65%, orcellulose 42% and Iignin 29%. Thesolubility in cold water was 1.5%, in hot water1.5%, in alcohol-benzene 1.1% and in I% NaOH10.8%. Pulping with the sulphate processyielded 42-48% pulp, with a kappa number of24-53. The pulps had relatively low tearingstrength; they were consldered suitable forgeneral purposes, but not for products wheretearing strength is a critical factor, such aspacking papersThe main constituents of o1eoresin from treesin Venezuela were u-pinene (40-50%), heptane(12-37%) and ^-pinene (5-14%). The o1e. restnand 2 terpenoids isolated from it (pimaric acidand longifolene) have shown trypanocidal ac-tivity against Tryponosomo cruz!, the causa-tive agent orchagas disease.

Botany Evergreen, monoecious, medium-sizedtree up to 30(-40) in tall; hole branchless for upto 15 in, up to 80(-125) cm in diameter, usuallystraight and cylindrical; bark 2-4 cm thick,bark surface grey to red brown, rough andscaly; crown wide, open; branches long, flexL-ble, curved upwards. Leaves clustered at theend of shoots, in bundles of (3-)5(-6), needle-shaped, 7-30 cm long, stiff, tips sharplypointed, bluish or pale green. Male cone cylin-drical, in dense clusters. Mature female coneoften in groups of 2-3, on a peduncle 3-4 cmlong, ovoid to ovoid-conical, 5-10 cm x 3-5 cm,pendulous, yellowish brown, with hard scaleswarty and pointed at apex. Seeds 4-7 mm long,dark brown, with a wing 10-12 mm long. Seed-ling with hypogealgermination.Growth of Pinus o0corpo is shrubby shortlyafter field establishment and it may take sev-eral years before a dominant stem develops.Once established, growth is rapid, with meanannual height increments up to 1.8 in duringthe first 10 years. Over the first 25 years anannual height growth of I in is possible, withan annual growth in hole diameter of 1.8-2.0

In southern Africa annual height Incre-merits of up to 85 cm and annual diameter in-

crements of 1.3-1.5 cm have been recordedPollination is by wind. The time from pollina-tion to mature cones is 18-21 months. Thecones remain on the tree for long times. Seedproduction is often poor near the equator.Seeds are dispersed by wind, but sometimesbirds, rodents or people disperse them. Pinuso0corpo resprouts easily from stumps afterfelling or damagePinus is a large genus comprising over 110

almost all restricted to the northernspecies,

hemisphere. Many Pinus species are cultivatedoutside their natural distribution area, intropical, subtropical and temperate regions. Inthe tropics 2 species are more important thananothers: Pinus cartboeo in the lowland humid

tropics and Pinus potu!a Schltdl. & Chain. inthe cooler highland tropics and subtropicsPinus o0corpo is a variable species, and 5 welldistinguished varieties can be recognized

Ecology Pinus o0corpo is grown at 250-2500in altitude, in areas with a mean annual tern-perature of 13-27'C, a mean maximum tern-perature of the warmest month of 20-34"C, amean minimum temperature of the coldestmonth of 7-20'C, an average annual rainfallof700-1500(-3000) mm, and a dry season of upto 6 months. Best growth is recorded underwarm-temperate to subtropical conditions, atabout 1500 in altitude. It is not tolerant offrost. Pinus o0corpo prefers light- to medium-textured, neutral to acid soils (optimum pH 5-6) which are well drained, but it tolerates shal-low soils and grows in a wide range of soiltypes. Trees are susceptible to fire when young,but become more resistant with age. Pinuso0corpo is light-demanding, but young treestolerate some shade. It rapidly colonizes ex-posed sites left bare by fire or erosion.

Management Pinus o0corpo is usually pro-pagated by seed. The 1000-seed weight is 8-24g. To obtain seeds, the cones can be air dried orkiln-dried, after which they open. Kiln tern-peratures of 40-44'C for 24 hours are recoin-mended, but temperatures of up to 50'C for12-18 hours can be applied without loss of vi-ability. The seeds can be stored for years underdry (6-9% moisture content) and cool(0-5'C)conditions. Germination usually takes 7-21

but indays. Pre-treatment Is unnecessary,commercial nurseries seeds may be soaked for24 hours to increase both rate and percentageof germination. Mycorrhizae are necessary forgood growth of seedlings, so it is recommendedto inoculate with spores or add soil from near

usuallyestablished trees. Seedlings are

PINUS 439

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planted out when they are 5-10 months oldand 20-30 cm tall. In Tanzania survival afterplanting out was much higher for 9-11-month-old seedlings than for 7-month-old ones. Anormal spacing is 2.7 in x 2.7 in. Direct sowingis only occasionally done. Vegetative propaga-tion by cuttings, grafting, air-layering or in-vitro techniques is possibleWeeds should be well controlled during estab-lishment of the tree. Trees are usually prunedand thinned several times, depending on pro-duction aims. Trees for pulpwood are pruned toabout 2 in height when 5-6 in tall. In planta-tions for sawn wood thinnings of 35-50% arecarried out every 5-6 years starting 8 yearsafter planting, and pruning up to 10 in highmay be applied in 3 rounds. In nurseries damp-ing off may occur. The fungus Cercosporo pint-densi/lorde causes a needle disease which cancause serious damage in plantations. In Zam-bia damage by Armi!!und meneu has been re-corded. Pinus o0corpo is highly resistant topitch canker (FMSorium circinotum), a seriousdisease threatening pine plantations in SouthAfrica. Pinus o0corpo is moderately resistantto termites, in general more so than Pinusconboeo. In Africa wild pigs are recorded tognaw at the roots and to uproottrees.Rotations of 23-30 years are normally appliedfor the production of timber. Mean annual in-crements in wood volume in plantations are10-40 ing/ha

Genetic resources and breeding Naturaland artificial hybrids with Pinus curtboeu havebeen recorded. Hybridization with Pinuspotulu is also possible. Provenance trials havebeen recorded in Nigeria, Kenya, Uganda, Tan-zania and southern Africa.

Prospects Pinus o0corpo Is a fast-growingpine species yielding wood of good quality,compared to that of other pines. It has beenplanted in many tropical African countries, butits actual importance and prospects in tropicalAfrica are difficult to assess

Major references Chudnoff, 1980; Chilufya& Tengnas, 1996; Dvorak, 2002a; Lamprecht,1989; van Wyk, 2002c

Other references Adegbehin, 2002; FAO,1974; Fanon, 1984; Palmer & Ganguli, 1985;Parant, Chichignoud & Curie, undated; Rubioat a1. , 2005; Schwarz, Beaty & Franc0, 1991;Suhardi at a1. , 1993; VelAsquez at a1. , 2000;Webb at a1. , 1984

Authors M. Brink

PINUS PATULA Schltdl. & Chain.

Protologue Linnaea a 354 (1831).Family PinaceaeChromosome number2n=24

Vernacular names Patula pine, Mexicanweeping pine, spreading-leaved pine, jelecotepine (En). Pin argent6, pin patula (Fr). Pinhopatula (Po). Msindano (Sw)

Origin and geographic distribution Pinuspotulo is native to Mexico. In 1907 it was in-troduced into South Africa, where it is pres-ently the most widely planted coniferous spe-cies and has become invasive in certain areasLater it was introduced into many other Mri-can countries, and it has become the most jin-portant pine in East and southern Africa. It isalso grown in Australia, New Zealand, Asiaand SouthAmerica

Uses The wood of young trees is mainly usedto manufacture boxes, and that of older treesfor light construction, light flooring, joinery,ceilings, panelling, shingles, furniture, cabinetwork, fence posts, poles, food containers, pal-lets, mine props, veneer and plywood. InMadagascar the wood is highly appreciated forglued laminated timber for carpentry and fur-niture, after the knots have been removed. It isalso suitable for hardboard, particle board andwood-wool. Pinuspotulo is an important sourceof pulpwood, e. g. for newsprint in South AfricaThe wood is excellent fuelwood and is also usedfor the production of charcoal. Pinus pornlo issuitable for reclamation of gullies, as the thickmat of fallen branchlets and needles retardssurface run-off, limiting the advance of soilerosion. It is also planted in windbreaks, as ashade tree and as an ornamental tree.

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Production and international trade World-wide, the area planted with Pinus potulo isapproximately I million ha, with plantations lnCentral, East and southern Africa accountingfor 95% of the planted area. Pinus potu!o isespecially important in Kenya, Tanzanla, Ma-lawi, Zimbabwe, Madagascar and South AfricaIn Kenya it accounts for about 25% of allforestplantations. In South Africa pines are grownon 54% of the total afforested area, with Pinus

potu!o being the most popular species, coveringan area of about 375,000 ha. Commercial plan-tations have also been established in ATgen-tina, Colombia, Ecuador, Peru and Brazil. Ex-ported products include round poles, pulp andpaper

Properties The heartwood is pinkish tocreamy white, and not clearly demarcated fromthe sapwood. The grain is straight, spiral or

texture fine. Growth rings are distinctwavy,

The wood contains little resin, although nu-merous resin canals are present and prominenton the tangentialsurface. It has a faint odourThe density of the wood is (330,430-650kg/ina at 12% moisture content. Air-dryingproperties are good, with only slight twisting ofsome boards. In Uganda 25 min thick boardstake 2-3 weeks to air dry to 20% inolsture con-tent. The rates of shrinkage from green to ovendry are 1.8-4.8(-9.5)% radial and 5.2-11.2%tangential. The dried wood is moderately sta-ble to stable in service

At 12% moisture content, the modulus of rup-ture is 47-154 N/mm2, modulus of elasticity5100-12,800 N/mm2, compression parallel tograin 25-57 N/mm2, shear 4-12 N/min2, cleav-age 9-21 N/mm, Janka side hardness 2350-2780 N, Janka end hardness 3360-3420 N andChalais-Meudon side hardness 1.5-3.8.The wood saws easily, but if fed too quickly arough surface will result. It planes easily, butboring, mortising and turning properties areless favourable. It takes and holds nails welland glues easily. The wood is not durable. It issusceptible to attacks by fungi, pinhole, long-horn, Anobium and marine borers, and ter-mites. The sapwood is not susceptible to Lyetusborers. Both heartwood and sapwood are easilyimpregnated with preservatives in open tankand pressure-vacuum systemsThe wood fibres are 2.0-4.9 mm long and 36-57 pin wide, with a cell wall thickness of 40-5.5 prn. The chemical composition of the oven-dry wood is: bolocellulose 59-73%, orcellulose40-44% and 11gnin 26-29%. The solubility incold water is 0.6-3.6%, in hot water 1.5-3.7%,

in alcohol-benzene 0.6-3.2% and in I% NaOH10.1-14.4%. Pulping with the sulphate (kraft)process yields 43-52% screened pulp, with akappa number of 21-61. Kraft pulps have inod-erate to good tensile and bursting strengthsand good tearing strength. The tearingstrength of the pulp was found to increase withthe age of the trees used for pulping, whereasbursting and tensile strengths decreased.

Description Evergreen, monoecious, medium-sized tree up to 30(-50) in tall, but in planta-tions often much smaller; hole branchless forup to 15 in, up to 120(-150) cm in diameter,usually straight and cylindrical; bark surfacegrey to dark brown, broken into longitudinal,irregular scales in the lower part of the hole,thin, papery and reddish brown higher up;

pyramidal; branches horizontal orturned upwards at their tips. Leaves in bun-dles of (2-)3-4(-5), needle-shaped, 12-30 cmlong, pendulous, bright to yellowish green.Male cone axiUary, small, yellow-brown. Ma-ture female cone in groups of 2-6, short-stalked, ovoid-conic, often curved, 4-12 cm x2.5-4 cm, oblique at base, pale glossy grey orbrown, persistent on the branches, with smooth

PINUS 441

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442 TIMBERSl

scales with a minute prickle, 40-80(-125)-seeded. Seeds triangular, 3-5 mm long, with awing 10-20 min long, grey inottled with blackSeedling with hypogealgermination

Other botanical information Pinus is alarge genus comprising over 110 species, al-most allrestricted to the northern hemisphereMany Pinus species are cultivated outside theirnatural distribution area, in tropical, subtropi-cal and temperate regions. In the tropics 2 spe-cies are more Important than all others: Pinuscartboeo Morelet in the lowland humid tropicsand Pinuspotulo in the cooler highland tropicsand subtropics.

Anatomy Wood-anatomical description OAWAsoftwood codes)Growth rings: (40: growth ring boundaries dis-tinct); 41: growth ring boundaries indistinct orabsent; 43: transition from earlywood to late-wood gradual. Tracheids: 44: tracheid pittingin radial walls (predominantly) uniseriate (ear-Iywood only); 54: latewood tracheids thin-walled (double wanthickness less than radiallumen diameter); 56: torus present (pits inearlywood tracheids only). Ray composition: 79ray tracheids commonly present; 82: cell wallsof ray tracheids dentate; 85: end walls of rayparenchyma cells smooth (unpitted); 87: hori-zorital walls of ray parenchyma cells smooth(unpitted). Cross-field pitting: 91: cross-fieldpits pinoid; 98: I-3 pits per cross-field (early-wood only). Ray size: 103: average ray heightmedium (5-15 cells); (104: average ray heighthigh (16-30 cells)); 107: ray width exclusivelyuniseriate. Intercellular canals: 109: axial in-tercellular (resin) canals present; 110: radialintercellular (resin) canals present; 1/7epithelialcells thin-walledre. Baas & I. Hemz)

Growth and development Pinus potulogrows very fast. Under favourable conditions itmay attain a height of 15 in after 8 years and35 in after 30 years, In southern Africa femaleflowering starts when trees are 2-3 years old,and male flowering I-2 years later. In south-ern Africa flowering of both male and femalecones is in August-October, with usually asecondary flush of only female cones in Janu-ary-May. In Kenya usually 2 flushes of bothmale and female flowering occur in April-Mayand in October-November, coinciding with therainy seasons, but female flowering throughoutthe year has also been recorded. Studies inZimbabwe showed that synchronization of pol-Ien shedding and female receptivity was goodat 1500 in altitude, whereas pollen shedding

occurred progressiveIy later at lower altitudesOutcrossing is predominant, and pollination ismainly by wind. Female cones mature in 22-30months. The production of viable seeds startswhen trees are 5 years old, and is prolific in 8-10-year-old trees. Seed dispersal is usually bywind, but sometimes also by birds, rodents orpeople

Ecology Pinuspotulo is grown at 1000-3300in altitude, in areas with a mean annual tern-perature of 9-23'C, a mean maximum tern-perature of the warmest month of 15-29'C, amean minimum temperature of the coldestmonth of 6-14"C, an average annual rainfallof(700-)1000-2200 min, and a dry season of upto 4 months. It grows best at higher altitudes:above 1000 in at 18-30' latitude, and above2000 in near the equator; several provenancestolerate severe frost. Cone production is best ata mean annual temperature of 13-16'C. Pinuspotulo has succeeded on a wide range of soils,but prefers well-drained, neutral to acid soilsIt is highly susceptible to fire. Pinus potulo isstrongly light-demanding. It is an aggressivepioneer species that grows readily in forestgaps created by fire. In Zimbabwe, South at-Tica and Swaziland it is now considered a sen-ous weed, invading forest margins, moist grass-land and road cuttings.

Propagation and planting Pinuspotulo ismainly propagated by seed. The 1000-seedweight is 6-11 g. Seeds are extracted by air orkiln drying of the cones, which open after 2-7days. In Zimbabwe cones are sun-dried inopen-sided sheds covered with plastic roofs,sometimes supplemented by kiln drying at amaximum temperature of 60'C. Seed can bestored for 5-10 years at 2-8'C and a moisturecontent of 6-10%. Germination starts 7-10days after sowing, and germination of fresh orwell-stored seed Is normally over 85% in Zim-babwe and South Africa. Pre-treatment is notnecessary, but germination may be improvedby soaking in cold water for I-8 days or in hy-drogen peroxide for I-4 days. The soil for sow-ing needs to be Inoculated with mycorrhizae byadding some soilcollected beneath mature pinetrees. Seedlings are planted out when they are4-12 months old and 10-30 cm tall. Wildlingsmay also be used as planting material. Normalspacings are 2.4-3 in x 2.4-3 in. Althoughnatural regeneration is often abundant, directsowing is usually not successful, probably be-cause the root system of seedlings is not deepenough to survive the first dry season. Pinuspotu!a should not be planted near crops be-

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cause of its shallow root systemVegetative propagation by grafting or air-layering is possible, and clonal orchards havebeen established using these methods. The useof cuttings for clonal propagation is limited byrapid initiation of ontogenetic aging in theplants, typified by the early onset of reproduc-tive maturity and resulting in variation In root-ing, growth habit, flowering and leaf inorphol-

Protocols for in-vitro propagation haveogy.

been developedNatural regeneration Is often prolific, e. g. inMalawi, Zimbabwe, South Africa and Mada-gascar, butts not common in East Africa. Seedsgerminate abundantly after a fire

Management Daring the firstyear after plant-ing out 2-3 weeding operations are required.The response of Pinus potu!o to fertilizers issite-specific. Pinuspotulo self-prunes poorly, sotrees are pruned when 4-6 year old to a heightup to 2.5 in, to reduce fire hazard and improveaccess ('low pruning'). In pulpwood plantatlonsno further pruning is done, although pruningup to a height of 6 in height has been recoin-mended to reduce the risk of fire. For the pro-duction of sawn timber, dead as well as livingbranches up to a height of 7(-12) in are re-moved to produce knot-free timber ('high prun-ing')Thinning depends on lnitial spacing, site qual-ity and end product. For the production of saw-logs in Zimbabwe and South Africa, the finalaim is a stand of over 400 trees/ha with a bolediameter of about 45 cm, which implies rota-tions of 25-35 years' In Zimbabwe plantationswith a density of 1100 treesA1a (spacing 3 in x3 in) may be thinned to 650 treesA1a after 6-8years, and to 400 trees/ha after 12-15 years' InMadagascar heavy thinning is recommended toarrive at a final density of 200-250 treesA1awhen the trees are 15 years old. For the pro-duction of pulpwood rotations of 15-25 yearsare normal, resulting in trees wlth a hole di-ameterofabout 30 cm

The density of the wood can be increased byplanting trees at lower rather than higher alti-tudes and allowing trees to grow longer beforeharvest. In Tanzania the density at 12% ino1s-ture content was found to increase from 380kg/in3 for 12-year-old trees to 510 kg/ina for 30-year-old trees

Diseases and pests When planted in hot,humid conditions, Pinus potulo Is susceptibleto infection by Sphoeropsis sopineo (synonym:Diplodio pined) after hail damage. In SouthAfrica Pinuspotulo is threatened by pitch can-

ker caused by the fungus FMSorium c, reLnotum,which was first recorded in the country in the1990s. In nurseries damping off may occur,mainly caused by Fusorium, Pythium andRhizoctonio spp. In Ethiopia Armingrid rootrot is common. Pests Include the grasshopperMecostibuspiniuoro in Zimbabwe. Pinuspotulois attacked by Ginoro cronortii (black aphid),Hy!OStes origustotus (Hylastes beetle) and sev-eral defoliating insects

Harvesting In Madagascar trees are felledwhen the hole diameter is 60 cm

Yield Mean annual increments are 10-40in31ha, in southern Africa 18-28 in31ha. In EastAfrica yields can be higher than in southernAfrica due to the shorter dry season. The totalyield (including thinnings) under favourableconditions maybe 630-700 msftia

Handling after harvest To avoidblue stain,log ends must be treated with a preservatlveimmediately after felling, and logs must beextracted and converted as soon as possibleAnti-stain dipping is highly recommended jin-mediateIy after sawing.

Genetic resources Various provenance col-Iections have been carried out in Mexico since1947, and provenance and progeny trials havebeen started in many countries, includingKenya, Malawi and Zimbabwe. Plantations InEast and southern Africa are believed to haveoriginated from a very restricted source ofgermplasm.Pinus potulo is included in the IUCN Red list,but it is classified in the lower rlsk category, inwhich it is considered to be of least concern.

Breeding Pinus potu!o responds readily togenetic improvement formcreased growth rate,stem form and branching. Advanced genetic

have been estab-improvement programmeslished in Zimbabwe and South Africa. Theseprogrammes have focused on the form and slzeof stem, branches and crown, and have re-suited in substantially increased timber yields.Hybrids of Pinus potu!o with Pinus o0corpoSchiede ex Schltdl. , Pinus rodiutu D. Don andother Pinus spp. have been developed. Treeimprovement programmes are also aimed atincreased productivity, increased vigourthrough modification of root systems and leafperformance, and tolerance to blotic and abioticstresses. Breeding for tree growth and formhas led to lower lncidence of reaction wood,wandering pith and knot associated defectsBiotechnological protocols have been developedfor somatic embryogenesis, cryopreservationand RAPD molecular markers. More recently

PINUS 443

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protocols for genetic engineering, including thegenetic transformation of embryogenic tissueusing biolistic and Agrobocterium-mediatedsystems, have been developed

Prospects Pinus potulo is and will remainan Important source of timber and pulpwood intropical Africa, especially in cooler highlandregions. On hot, dry sites at lower altitudesPinus hestyo Royle ex Gordon and Pinus o0-coreo are better able to withstand droughtstress, and on hot, humid sites Pinus curtboeoand Pinus enjottii Engelm. will do better. Themain problem associated with Pinus potulo isIts aggressiveness and weediness. Improve-merit of yield and quality of the wood is neededto meet growing timber demands. Research isneeded to improve timber quality by decreasingIignin content and increasing cellulose content;this would make it easier to process the woodinto pulp and paper products, and should jin-prove durability, hardness and stability.

Major references Bekele-Tesemma, 2007;Bolza & Keating, 1972; CTRAD Forestry De-partment, 2003; Dvorak at a1. , 2000; Maundu& Tengn6. s, 2005; Morris & Pallett, 2000;Nyoka, 2002; Poynton, 1984; Suhardi at al. ,1993; Takahashi, 1978.

Other references Chudnoff, 1980; Coutinhoat a1. , 2007; CTFT, 1959b; Dvorak, 2002b;Hemz, 2004; Lamprecht, 1989; Malabadi & vanStaden, 2005; Mitchell, Zwolinski & Jones,2004; Morris at a1. , 1997; Nigro at a1. , 2004;Nigro at a1. , 2008; Owen & van der Zel, 2000;Palmer & Gibbs, 1974; Palmer, Ganguli &Gibbs, 1984; Palmer at a1. , 1982; Parry, 1956;Sutter, 1990; Webb at a1. , 1984; World Agrofor-estry Centre, undated; Wormald, 1975.

Sources of illustration Fanon, 1984Authors S. A. Nigro

Africa it has been planted in Ghana, Nigeria,Sudan, Ethiopia, Kenya, Tanzania, Malawi,Zimbabwe, SouthAfrica andMadagascar

Uses The wood (trade names: radiata pine,Californian pine) is used for construction, join-ery, furniture, veneer, plywood, packing cases,poles, posts, shuttering, particle board andfibreboard. It is also suitable for flooring, inte-rior trim, toys, turnery, matches, railwaysleepers, hardboard and wood-wool. It is usedas fuelwood and widely used for paper making01eoresin is tapped from the trees and distilledto obtain turpentine and rosin. Turpentine isused for the production of pine oil, terpene res-ms. flavours and fragrances. Rosin is used inthe production of paper, inks, emulsifiers, syn-thetic resins, soap and glue. Pinus rodioto hasbeen planted for soil conservation, in wind-breaks and as an ornamental tree.

Production and international trade Pinusrodioto is the most widely planted pine in theworld, occupying about 4 million ha, of which90% is in the southern hemisphere. The maingrowing countries are Chile and New Zealand(about 1.5 million ha each), followed by AUStra-11a (more than 700,000 ha), Spain (about220,000 ha) and South Africa (about 55,000ha). In East Africa it was much planted untilitwas severely attacked by Dothistroma needleblight (Mycosyhoerellopini) in the 1960s, eaus-ing abandonment of large-scale plantings inmany parts of Africa in favour of more resis-tant species such as Pinus potulo Schltdl. &Chain. and Cupressus lusttonico Mill.

Properties The heartwood is pinkish brownand distinctly demarcated from the exception-ally wide creamy white sapwood, which consti-tutes the bulk of the commercial timber. Thegrain is often spiralled near the pith, but else-where usually straight, texture moderately fineand even. Growth rings are up to 3 cm wide.Numerous resin canals are present. Knots arePINUSRADiATAD. Don

Protologue Trans. Linn. SOC. London 17: 442(1836).

Family PinaceaeChromosome number2n=24

Vernacular names Radiata pine, Montereypine, irisignis pine (En). Pin de Monterey (Fr).Pinheiro irisigne (Po)

Origin and geographic distribution Pinusradioto occurs naturally in California (UnitedStates) and the Mexican islands Guadalupeand Cedros. It is widely planted elsewhere,especially in the southern hemisphere and ithas become naturalized in various countries. In

common.

The density of the wood is (330-)380-610kg/in3 at 12% moisture content. The wood airdries well. Wood to be used for joinery needskiln drying for 2-3 days to remove stressesafter air drying. Shrinkage from green to ovendry is about 3% radial and 5% tangential.The wood is fairly soft. At 12% moisture con-tent, the modulus of rupture is 58-99 N/min2modulus of elasticity 7900-15,400 N/min2compression parallel to grain 31-50 N/mm2shear 10-13 N/mm2, cleavage 16 N/min radialand 13 N/min tangential, Janka side hardness2270-5590 N and Janka end hardness 3510 N

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The wood saws easily, but splitting may occurin thick logs. It works well with hand and ina-chine tools, and careful planing gives a goodsurface. The wood takes nails well, but olderand denser wood may split. Holding propertiesfor screws are good. The wood glues, paints andvarnishes well. It is not suitable for steam

bendingThe wood, mainly consisting of sapwood, is notdurable. It is susceptible to attacks by fungiand termites, and occasional damage by Ano-binm, pinhole and longhorn borers is possible.The sapwood is not susceptible to Lyetus bor-ers. The sapwood is permeable to impregnationwith preservatives, the heartwood is moder-ately resistant to resistantThe wood can be pulped using mechanical,chemical and semi-chemical pulping processes.Wood fibre cells of 20-year-old trees grown inKenya were 2.8-3.1 min long, with a diameterof 34-37 prn and a cellwallthickness of 4.4-4.5prn. The chemical composition of the oven-drywood was: holocellulose 61-64%, orcellulose41-43% and Iignin 25-26%. The solubility Incold water was 2.3-3.2%, in hot water 2.4-3.5%, in alcohol-benzene 0.8-1.2% and in I%NaOH 12.0-12.9%. Pulping with the sulphate(kraft) process yielded 43-49% screened pulp,with a kappa number of 20-45. Wood fibre cellsof 25-year-old trees in Sudan were on average2.7 mm long, with a diameter of 39 pin and acell wall thickness of 4.2 prn. The chemicalcomposition of the oven-dry wood was: holocel-Iulose 71%, u-cellulose 45% and 11gnin 22%The solubility in hot water was 4.0%, in alco-hol-benzene 1.9% and in I% NaOH 15.0%

Pulping with the soda-arithraqulnone processyielded 48-52% pulp, with a kappa number of27-48 and good strength properties.The energy value of the wood is 20,470 kJ/kg.01eoresin from Kenyan trees contained 69%rosin and 25% turpentine.

Botany Evergreen, monoecious, large treeup to 55(-60) in tall; hole more or less straight,up to 150(-250) cm in diameter; outer barkdark brown, deeply grooved with age; crowndense; branches UPCurved. Leaves in bundles of(2-)3, needle-shaped, (3-)8-18(-20) cm long,soft, sharp-tipped, usually intense dark greenbut sometimes yellowish or bluish green. Malecone in dense clusters. Mature female cone in

groups of 3-6, 0void-conical, 5-21 cm x 2.5-12cm, oblique at base, shiny pale brown, persis-tent on the branches for many years, withscales having a blunt apex, c. 200-seeded.Growth can be exceptionally fast. After an inI-

tial phase of 2-5 years, plants have a growthrate of up to 2 in/year. In Madagascar 36-year-old trees had a height up to 37 in and a bolediameter up to 58 cm. Pollen production startswhen trees are 5-6 years old, and female flow-ering starts around the same age. Pollination Isby wind. Female cones take 2 years to mature.Mature cones may remain closed on the tree forup to 40 years'Pinus is a large genus comprising over 110

almost all restricted to the northernspecies,

hemisphere. Many Pinus species are cultivatedoutside their natural distribution area, in

tropical, subtroplcal and temperate regions. Inthe tropics 2 species are more important thanall others: Pinus conboeo Morelet in the low-land humid tropics and Pinus potulo in thecooler highland tropics and subtropics

Ecology Pinus radioto is grown at 1500-3000 in altitude, in areas with a mean annualtemperature of 8-18'C, a mean maximumtemperature of the warmest month of 13-30'C,a mean minimum temperature of the coldestmonth of -3-12'C, an average annual rainfallof 650-1600 mm, and a dry season of up to 6months. It requires more fertile soils thanother pines, and grows best on deep, well-drained, neutral to acid soils. Provenances varyin frosttolerance. Ground fires are tolerated byolder trees, but crown fires can killthe tree.Pinus rodioto is light-demanding but moretolerant of shade than most other pines;it mayeven form a vigorous understorey. It may be-

Management Propagation Is mainly by seed.The 1000-seed weight is 18-30 g. To open thecones, they are dried in the open or in kilnsKiln temperatures up to 55'C are toleratedSeeds can be stored for several years undercool, dry and airtight conditions. Germinationis rapid and uniform, pre-treatment Is urineC-essary. Mycorrhizae are necessary for seedlinggrowth, so it is recommended to inDCulate withspores or to add soil from near establishedtrees. Seedlings are ready for planting outwhen 4-8(-24) months old. Common spacingsare 1.5-3 in x 1.5-3 in. Vegetative propagationby shoot cuttings or grafts is also practised.The use of tissue culture and embryogenesis IstechnicalIy feasible.During establishment 2-3 weedings per yearare recommended. Herbicides and fertillzersare often applied in commercial plantations.Trees are usually pruned and thinned. Triten-SIve systems are common in New Zealand, withheavy thinning (down to 200-250 treesA1a) and

PINUS 445

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pruning (to 5 in height), and rotations of 25years or less, whereas in Australia light thin-rimg and low pruning are common, with rota-tions of up to 50 years. The number of knots inthe wood can be reduced by close planting andpruning

Pinus radioto is affected by Dothistroma nee-dle blight (Mycosphoerello pint), causing thedeath of needles and subsequently the wholetree. The disease has led to the abandonmentof PInus rodioto as a plantation species inmany parts of Africa, and the use of alternativespecies, such as Pinus potulo. Progress hasbeen made in breeding for resistance to thedisease. Pinus rodioto is one of the pines mostsusceptible to pitch canker caused by the fun-gus Fusorium circinotum, one of the most jin-portant pine diseases in the world and threat-ening plantations in South Africa. Symptomsare resin-soaked cankers on the hole and

branches, shoot die-back and death of youngand mature female cones. In nurseries itcauses damping-off, shoot and tip die-back anddeath of seedlings. mother important diseaseis Sphaeropsis canker caused by Sphoeropstssoptneo (synonym: Diplodio pined), which maylead to shoot die-back and death of the treeSphoeropsis sopineo is particularly common inAfrica. Pinus rodiuto is also susceptible to Ar-minorto root rot, common in Ethiopia. In SouthAfrica the pine emperor moth (lin brasio cythe-red) can be troublesome, and the pine whoollyaphid (Pineuspini)is causing concern.Annual volume Increments up to 40 in3ftia arepossible. Logs left on the ground after fellingwill discolour ifthey are not treatedTo obtain o1eoresin, trees over 23 cm in diame-ter are tapped by wounding the tree ('streak-ing') and attaching a cup to collectthe exudate.A rosin plant has been established in Kenya

Genetic resources and breeding Abun-dant genetic variation has allowed for highlysuccessful breeding programmes. Provenancetesting and breeding is done in South Africaand other major producing countries. Earlybreeding efforts focused on growth rate, treeform and disease resistance, but emphasis hasshifted towards wood properties. Molecularbiology is being explored, and protocols havebeen developed for genetic transformation ofembryogenic tissue using biolistic andAgroboc-tenum-mediated systems, and stably trans-formed plants have been regenerated

Prospects Its very fast growth, ease of es-tablishment, wide ecological adaptation andthe suitability of the wood for a wide range of

applications make it the utility softwood ofchoice wherever it can be grown satisfactorilyIn tropical Africa, however, its susceptibility todiseases is a major problem, which has pre-vented wide success

Major references Bekele-Tesemma, 2007;Bolza & Keating, 1972; Burdon, 2002; Lain-precht, 1989; Takahashi, 1978.

Other references Banks & Schoeman,1963; Coutinho at a1. , 2007; Khristova, Gabir &Tah, 1990; Njenga, 1995; Palmer at a1. , 1982;Rendle, 1970; Sutter, 1990; Walter at a1. , 2002;Webb at a1. , 1984; Willan, 1985

Authors M. Brink

PIPTADENIASTRUMAFRICANUM (Hook. f. )Brenan

Protologue Kew Bull. 1955(2): 179 (1955)Family Mimosaceae (Leguminosae - Mimo-

soldeae)Chromosome number2n=26

Synonyms Aptadeni0 o1>icono Hook. f. (1849).Vernacular names Dabema, danoma, African

greenheart(En). Dab6ma (Fr). Musence (Po)Origin and geographic distribution Pipto-

dentostrum of neonum occurs from Senegaleast to southern Sudan and Uganda, and southto DR Congo and northern Angola

Uses The wood (trade names: dabema, daho-ina) is used for construction, including marineconstruction and bridges, flooring, railwaysleepers, mine props, ship building, vehiclebodies, Interior trim, joinery, furniture, includ-ing garden furniture, cabinet work, sportinggoods, turnery, hardboard, particle board andpulpwood. It is used traditionally to make dug-

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out canoes. In Europe the wood is consideredan excellent replacement for oak (Quercusspp. ), and is sometimes called 'African oak'. Itis also used as fuelwood and for charcoal pro-duction.

Piptudeniostrum of neonum is commonly usedin traditional medicine, mostly the bark, some-times also roots and leaves. Bark decoctionsare used internally to treat cough, bronchitis,headache, mental disorders, haemorrhoids,genito-urinary infections, stomach-ache, dys-menorrhoea and male impotence, and as anantidote; externalIy, they are applied to treatfever, toothache, pneumonia, oedema, skincomplaints and rheumatism, to expelworms, todispelfleas, and as a purgative and abortifa-cient. A decoction of the bark also enters in acomplex treatment of leprosy. The bark is usedin arrow poison, and as ordeal poison and fishpoison; mixed with rice it is used to poisonmice. It is also used as a soap substitutePygmy people in Cameroon and DR Congo useboth root bark and stem bark as an ingredientof arrow poison. Root extracts or macerationsare applied against mental disorders, and asan abortifacient and aphrodisiac. Poundedleaves and leaf decoctions are applied as anenema to treat gonorrhoea and abdominalcomplaints. Leaves are used to poison miceThe tree is planted or left during forest clear-ing as a shade tree in coffee, cocoa and bananaplantations. The bark fibre has been used toweave mats. Edible caterpillars feed on theleaves, and the flowers are a source of nectarfor honey bees. In various countries Piptuden, -OStrL, in of neonum Is considered a magic tree

Production and international tradeAccord-

ing to ITTO, Ghana exported 4000 in31year ofsawn dabema wood in 2003 and 2004, at anaverage price of Us$ 3101m3. C6te d'TVoire ex-ported 10,000 ina of sawnwood in 2004, at anaverage price of Us$ 3971m3, and 4000 ing in2005, at Us$ 4391m3. Garnero0n exported 7000ms of logs in 2005, at an average price of Us$5551m3 and in 2006 the volume was 14,000 ina,at Us$ 3581m3. According to ATIBT ('Associa-tion technique Internationale des bois tropi-caux'), Cameroon exported 400 in3 of sawnwoodin 2003, 800 in3 in 2004, and 2000 ms In 2006,whereas log exports were 21,000 ina in 2006.Gabon exported 950 in31year of logs in 2003and 2004, and 15,000 ms in 2005.

Properties The heartwood is pale brown togolden brown, occasionally dark brown, withan attractive stripe pattern on quarter-sawnsurfaces, distinctly demarcated from the 5-15

cm thick, pale pink to greyish red sapwood.The grain is interlocked, texture coarse. Thewood has an unpleasant ammoniacsmellwhenfreshly cutThe wood is moderately heavy. At 12% ino1s-ture content, the density is (480,590-800(-900) kg/ms. The rates of shrinkage during dry-ing are moderate to high, from green to ovendry 2.5-5.2% radial and 7.0-10.5(-13.4)% tan-gential. The wood air dries rather slowly, withhigh risk of distortion and checking. In south-ern C6te d'TVoire boards 29 min thick in hori-zoritalroofed piles take 38 weeks to dry, and 50min boards 61 weeks. Air drying prior to kilndrying is recommended. After drying, the woodis moderately stable in service.At 12% moisture content, the modulus of rup.ture is 80-178 N/min2, modulus of elasticity9300-16,500 N/min2, compression parallel tograin 46-74 N/mm2, shear 7-18 N/mm2, cleav-age 14-32 N/min, Janka side hardness 6400-6860 N and Janka end hardness 8100 N.The wood is fairly easy to saw and work, butwith some blunting effect on cutting edges, sothe use of stellite-tipped sawteeth and tung-sten-carbide-tipped cutting tools is recoin-mended. A cutting angle of 10-15' is recoin-mended in planing and moulding operations toavoid picking-up of grain. The wood finisheswell, but the use of a filler is needed. It holdsscrews and nails well, but there is a slight ten-dency for splitting. The gluing properties aresatisfactory. The steam bending properties aremoderate. The wood is not particularly suitablefor veneer or plywood production; it should bewell steamed to make peeling possible. Thesawdust may irritate skin, throat and eyes.The heartwood is moderately durable. Itshowed moderate resistance to fungal, dry-wood borer and termite attacks. In tests inGhana, the wood showed no damage after be-ing subjected to Coptotermes formosonus ter-mites in exposure chambers, antermites beingdead within one week. It is recorded as durablein fresh water. The heartwood does not absorb

preservatives, but the sapwood is only moder-ately resistant. However, it is also reportedthat for use as railway sleepers Impregnatlonis needed. The degree of resistance to fungalattacks depends on the concentration of dihy-droflavonols, which is higher in the outer partof the heartwood than in the inner part, whichis thus less durable

Kraft pulping experiments showed that thewood has good prospects for paper production.In Ghana the charcoal madefrom the wood has

PIPTADENIASTRUM 447

.

448 TIMBERSl

been found to be suitable for preparation ofactivated, decolorizing carbon used in sugarrefining.The roots and stem bark contain saponins,

tannins, navonoids and Ieucoanthocyanes. Thestem bark is very toxic; this has been con-firmed by tests with rats. It showed moderateactivity against several pathogenic bacteria

Adulterations and substitutes The timberof Cylicodiscus gobunensis Harms is similaralthough slightly heavier, and is also traded asAfrican greenheart

Description Deciduous or evergreen large treeup to 50 in tall; hole straight and cylindrical,but sometimes sinuous, branchless for up to20(-30) in, up to 180(-300) cm in diameter atbase, with large, thin buttresses up to 5G8) inhigh, often extending into branched and sinu-ous surface plank roots; bark surface smooth,sometimes with ring marks and numeroussmalllenticels, in old trees sometimes slightlyfissured, yellowish brown to greyish brown orreddish brown, inner bark whitish to pale yel-low or brown, brittle; crown spreading, flat;young twigs densely brown short-hairy. Leavesalternate, hipinnately compound with 10-19

pairs of usually alternate pinnae; stipules 11n-ear, up to 9 mm long, early caducous; petiole0.5-3 cm long; leaflets opposite, 25-60 pairsper pinna, sessile, linear, 1.5-8.5 min x c. Imin, asymmetrical at base, obtuse at apex,hairy at margin. Inflorescence an axillary orterminal spike-like false raceme up to 11 cmlong, often many together at ends of twigs,hairy, densely flowered. Flowers bisexual,regular, 5-merous; pedicelc. 0.5 mm long; calyxwith c. 0.5 mm long tube, toothed; petals free,oblong to lanceolate, 2.5-3 mm long, whitish oryellowish; stamens 10, fused at base, c. 4 mmlong, arithers with gland at apex; ovary sripe-nor, ellipsoid, c. 2 mm long, with c. I mm longstipe, glabrous, style slender. Fruit a flattenedlinear pod 12.5-36 cm x 1.5-3 cm, shortly sti-ped at base, dark brown, transversely veined,dehiscent at one side, up to 9-seeded. Seedsoblong, flat, 3-9.5 cm long including the paperywing surrounding the seed, glossy brown, at-tached at the middle. Seedling with epigealgermination; hypocoty1 I-3 cm long, epicotyl c.3 cm long; cotyledons shell-shaped, c. I cm x2.5 cm, slightly fleshy; first leaves pinnatelycompound with many leaflets

Other botanical information Piptodenios-trum comprises a single species and seems tohave a rather isolated position within its fam.11y. It may be confused with Newtonio, whichdiffers in the presence of glands on the leaves,its hairy ovary and its seeds being attached atthe apex

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;26: intervessel pits medium (7-10 qrn); 27intervessel pits large (a 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 pin); 43mean tangential diameter of vessellumina z200 prn; 46: S 5 vessels per square minimetre;(47: 5-20 vessels per square minimetre); 58:gums and other deposits in heartwood vessels.Tracheids and fibres: 61: fibres with simple tominutely bordered pits; (65: septate fibres pre-sent); 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: (76:axial parenchyma diffuse); 79: axial paren-chyma vasicentric; 80: axial parenchyma an-

11 1<}*t f-

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Piptodeniostrz, in of neonum - I, base of 601e, . 2,flowering trotg, . 3, fruit, 4, seed.Redrown und udopted by AChmod Satin NILr-hornon

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form; 81: axial parenchyma lozenge-anform;83: axial parenchyma confluent; 91: two cellsper parenchyma strand; 92: four (3-4) cells perparenchyma strand; (93: eight (5-8) cells perparenchyma strand). Rays: 98: larger rayscommonly 4- to 10'seriate; 104: all ray cellsprocumbent; 1/5: 4-12 rays per min. Mineralinclusions: 136: prismatic crystals present; 142:prismatic crystals in chambered axial paren-chyma cells.(E. Uetimane, H. Beeckman & P. E. Gasson)

Growth and development Seedlings areoften common in the forest, even in dark shade.However, they grow very poorly in shade,where they may be only 20-35 cm tall when 3years old, whereas they may reach up to 150cm tall after 4 years in less shaded conditionsin the forest. Saplings and young trees demandlight, and are usually found in small gaps inthe forest. In Sierra Leone mean annual hole

diameter increments of 1.2 cm have been re-

corded for the first 20 years after planting. InC6te d'TVoire the mean annual diameter incre-

merit was 4.9 min. In Ghana mean annual di-

ameter increments were highest in the diame-ter classes of 30-49 cm and 50-69 cm, with 9.0mm and 11.1 mm respectively. In Nigeria atree reached 90 cm bole diameter in 71 years.Young trees have a rounded crown, which de-velops into 2 distinct layers, of which the lowerone disappears later and the upper one devel-ops horizontally and ultimately becomes frag-merited. Adult trees have huge, flat crownsthat spread in the upper canopy of the forest.The trees are often briefly deciduous, but fre-quently do not shed allleaves at the sametime. The leaflets fold up at sunset. In SierraLeone, Liberia and C6te d'Ivoire trees usuallyflower in May-August and fruits mature inDecember-March; in Nigeria flowering hasbeen observed in June-September and fruitingin October-March. In Ghana fruits are ripetowards the end of the dry season. In Ugandaflowering occurs mainly in July and August.The winged seeds are mainly dispersed bywind, but distribution by water and birds isalso possible. Piptodeniostrum o17iconumnodulates with rhizobia.

Ecology Piptoderriostrt, in of neonum occursin lowland evergreen and semi-deciduous for-est, up to 1200 in altitude. In Ghana it shows apreference for evergreen forest. In Gabon itoccurs scattered in primary forest, but may belocally more abundant in old secondary forest.Some preference for hillsides and moist butwell-drained levelground has been recorded for

Sierra Leone, but Piptodeniostrum @1nconumis also commonly found in riverme forest. InUganda it is a common and character1stlc spe-cies of the forest in the Lake Victoria belt

Propagation and planting Seedsfor plant-are collected from the forest floor. TheIng

1000-seed weight is about 180 g. Seeds losetheir viahinty quickly and cannot be stored formore than one month. They do not show dor-mancy, and germinate in I-3 weeks. In a ger-mination test, 96% of the seeds germinatedwithin 8 days. They should be sown in shadednursery beds. The growth of seedlings Is slowand they may stay in the nursery for morethan one year before planting. Wildlings aresometimes collected for planting.

Management In general Piptodeniustrum of-neonum occurs rather scattered in the forestIn Gabon the average bole volume is 0.6 mania,but in southern Garnero0n it is 1.6-4.8 in3ftiafor hole diameters over 60 cm. In some regionsPiptodeniostrum o17iconum occurs in highdensities, e. g. in some forests in Sierra Leone,where it may account for 10% of all trees ofover 60 cm hole diameter. In Liberia an aver-

age density of up to I hole over 60 cm diameterper ha has been recorded. In Uganda Piptode-ittostrum o17iconum has been planted success-fully in experimental plantations.

Harvesting The minimum felling diameteris 60 cm in C6te d'TVoire, Gamero0n and DRCongo, 80 cm in Liberia and the Central Mri-can Republic, and 90 cm in Ghana. The highbuttresses at the base of the bole necessltate

the construction of a platform before felling cantake place. The holes have a tendency to splitduring felling operations

Yield Trees with a hole diameter of 60, 90and 120 cm yield about 3.1, 7.2 and 13.1 in3 oftimber, respectively.

Handling after harvest Freshly harvestedlogs sink in water and thus cannot be trans-ported by river

Genetic resources Pintodeniostrum ofri-corium is widespread in different forest typesin West and Central Africa and is common in

many regions. Therefore, It does not seem to bethreatened at present. However, it seems to begaining in importance as a commercial timbertree, and the bark is commonly partially re-moved for applications in traditional medicineThis may make Piptodertiostrum of neonumliable to genetic erosion in the near future

Prospects There is much demand on the in-ternationaltimber market for Piptodeniustrt, inof neonum, although the applications of the

PIPTADENIASTRUM 449

.

450 TIMBERSl

wood are somewhat limited due to the highshrinkage rates. This demand offers possibili-ties for increased commercialization of the spe-cies, but research is needed on growth ratesunder different ecological conditions and thedevelopment of suitable management methodsfor forest in which it is a common constituentto guarantee sustainable production in thefuture.

Although Piptodeniostrum o17iconum has nu-meTous applications in traditional medicine,very little research has been done on its phyto-chemistry and pharmacological propertiesResearch is warranted to assess its possibilitiesfor drug development, but also in the light ofthe known toxic effects of the bark, which isthe most commonly used part of the tree inlocal medicine.

Major references ATIBT, 1986; Bolza &Keating, 1972; Burki11, 1995; CTFT, 1974a;Katende, Birnie & Tengnas, 1995; Neuwinger,1996; Phongphaew, 2003; Saville & Fox, 1967;Takahashi, 1978; Voorhoeve, 1979.

Other references Adjanohoun et al. (Edi-tors), 1988; CTRAD Forestry Department, 2003;de to Mensbruge, 1966; 060n & Schwartz,1988; de Saint-Aubin, 1963; Durrieu de Mad-ron, Nasi & 06tienne, 2000; Farmer, 1972;Hawthorne, 1995; Hawthorne & Jongkind,2006; InsideWood, undated; Keay, 1989;Latham, 2004; Lewis at a1. , 2005; Neuwinger,2000; Normand & Paquis, 1976; Noumi &Tchakonang, 2001; Onanga at a1. , 1999; SIepel,Poorter & Hawthorne, 2004; Sprent, 2001;Viniers, 1989; Vivien & Faure, 1985

Sources of illustration Brenan, 1959;Voorhoeve, 1979; Wilks & Issemb6, 2000

Authors R. B. Jiofack Tafokou

almost black, and distinctly demarcated fromthe white sapwood. The grain is fairly straight,texture fine to medium and even. The wood is

heavy, with a density of 810-900 kg/in3 at 12%moisture content, hard, and resistant to shock.The rates of shrinkage are high, and the woodmust be dried slowly. Once dry, it is stable inservice.

The wood works fairly easily, but polishing andvarnishing are difficult. It is durable, and re-sistant to termite attack, but the sapwood issusceptible to attacks by Lyctus borers. Theheartwood is extremely resistant to impregna-tion, the sapwood moderately resistant.

Botany Shrub or smallto medium-sized treeup to 18 in tall; bole usually straight, up to 60cm in diameter, without buttresses; twigsshort-hairy, soon becoming glabrous. Leavesalternate, imparipinnately compound with 3(-4) pairs of leaflets, up to 40 cm long; stipulescaducous; petiole and rachis grooved above,short hairy to glabrous; leaflets with needle-shaped, deciduous stipels and petiolules 2-5min long, oblong to elliptical or oblong-obovate,7-15 cm x 3-6 cm, obtuse to slightly curieate atbase, short-acuminate at apex, thinly leathery,appressed short-hairy below, pinnately veinedwith 7-10 pairs of lateral veins. Inflorescence aterminal or axillary panicle up to 40 cm long,densely hairy, many-flowered. Flowers bisex-ual, papilionaceous, with 2 bracteoles up to 2min long at base of calyx, caducous; pedice14-7min long; calyx 2-11pped with large hood-shaped upper lip up to 15 min long and un-equally 3-lobed lower lip up to 6 mm long; co-rolla whitish, with standard in bud covered bythe upper lip of the calyx, wings and keelc. 12min long; stamens 10, 9 united and I free;ovary superior, linear, with short stipe, denselyhairy, style curved. Fruit a linear-oblong pod7-10 cm x I-2 cm, flattened, with thickenedmargins, silky brownish hairy, 3-5-seededPlotysepulum comprises about 8 species and isconfined to tropical Africa.

Ecology Plotysepolum cheuolieri occurs indense primary or secondary forest up to 600 inaltitude, on dry to swampy soils

Genetic resources and breeding There isno reason to consider the little used Plotyse-polum cheuolieri as threatened by genetic ero-SIon, although it has a rather limited distribu-tion area

Prospects Plotysepolum cheuo!tents a goodsubstitute for hickory (Goryo spp. ) as a sourceof wood for sporting equipment. However, thehole is usually too small-sized to make it at-

FIATYSEPALUMCHEVALIERIHarms

Protologue Bull. SOC. Bot. France 54, Mein.a 15 (1907); Bot. Jabrb. Syst. 40: 37 (1907)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution PIO-tysepolum cheuolierioccurs in the Central Mri-can Republic, Congo and DR Congo

Uses The wood is suitable for construction,flooring, vehicle bodies, furniture, cabinetwork, handles, ladders, sporting goods, agricul-turalimplements, sleepers, poles, piles, mineprops, toys, novelties and turnery. In DRCongo a root decoction is taken to lose weight

Properties The heartwood is dark brown to

.

tractive for commercial exploitation.Major references Bolza & Keating, 1972;

Hauman at a1. , 1954b.Other references ILDIS, 2005Authors R. H. M. J. Lemmens

FIATYSEPALUMVIOLACEUMWelw. exBaker

Protologue Onv. , F1. trop. Mr. 2: 131 (1871)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Chromosome number it = 14

Synonyms Plotysepolum DonhoutteiDe Wild(1906).

Origin and geographic distribution PIO-tysepo!urn uioloceum occurs from southern Ni-gena east to DR Congo and south to northernAngola

Uses In DR Congo the wood is used to maketraps for game. In Nigeria the flowers, parchedand mixed with oil, are applied to sores.

Properties The heartwood is dark and hard.Botany Shrub or smalltree up to 15 in tall;

hole up to 20 cm in diameter; bark smooth,reddish brown; crown spreading; twigs gla-brous. Leaves alternate, imparipinnately coin-pound with (2-)3-4 pairs of leaflets; stipules 3-4 mm long, pointed, caducous; petiole 4-6 cmlong, rachis 4-11 cm long; leaflets with needle-shaped stipels 2-3 min long and petiolules 3-4mm long, obovate to elliptical, 9-13 cm x 2.5-4.5 cm, curieate at base, long-acuminate atapex, margin often slightly indented, thinlyleathery, glabrescent, pinnately veined withdistinct lateral veins. Inflorescence a terminal

or axillary panicle up to 30 cm long, denselyhairy, many-flowered. Flowers bisexual, papil-ionaceous, with 2 bracteoles up to I cm long atbase of calyx; pedice1 2-3 min long; calyx 2-lipped with large hood-shaped upper lip up to15 mm long and unequalIy 3-lobed lower lip upto 8 mm long; corolla pale purplish, with stan-dard in bud covered by the upper lip of thecalyx, wings and keelc. 15 mm long; stamens10, 9 united and I free; ovary superior, linear,with short stipe, densely hairy, style curved, c.6 mm long, glabrous. Fruit an ellipsoid or 11n-ear-oblong pod 9-11 cm x 2-2.5 cm, flattened,with thickened margins, silky hairy, 2-3-seeded. Seeds disk-shaped, c. 1.5 cm in diame-ter, blackPlotysepolum comprises about 8 species and isconfined to tropical Africa.

Ecology Plotysepolum uLoloceum occurs inprlmary and secondary forest, also in marshy

localities.

Genetic resources and breeding Plotyse-polum uto!Geeum is widespread in Central at-rica and probably not threatened by geneticerosion.

Prospects The boles of Plotysepolum uiolo-ceum trees are too small to be of importance incommercial timber logging operations

Major references Burki11, 1995; Hauman ata1. , 1954b

Other references Hepper, 1958Authors R. H. M. J. Lemmens

PLEIOCARPA 451

PLEIOCARPAPYCNANTHA (K. Schum. ) Stapf

Protologue Ofiv. , F1. trop. Afr. . 4(I): 99 (1902)Family ApocynaceaeSynonyms Pieiocorpu 110uescens Stapf(1902),

Pieiocorpo microntho Stapf(1902).Origin and geographic distribution Pieio-

euroo pyc, tontho is widespread from Senegaleast to Kenya, and south to Angola and Mo-zambique.

Uses The wood is used for local construction,combs, plane-blocks, implement handles andpestles, and also for carving, e. g. to make pipe-stems in Uganda. Ground roots mixed withseeds of Afromomum melegueto K. Schum. andpalm wine are taken as a laxative. In Benln aleaf maceration with lemon Julce is adminis-tered to patients suffering from jaundice, oe-dema, reduced urine excretion and infection byroundworms

Properties The wood is yellow to brown, hardand durable. Some indole alkaloids have beenisolated from PIeiocorpo pycnontho roots andbark, e. g. pycnanthine, PIeiocarpamlne, que-brachamine and macusine B.

Botany Shrub or smallto medium-sized treeup to 20(-30) in tall; hole up to 50 cm in diame-ter; bark smooth to fissured or reticulatelycracked, grey to reddish brown. Leaves OPPo-site or in whorls of 3-5, simple and entire; stip-ules absent; petiole 5-20 min long; blade nar-rowly elliptical to oblong, 4-22 cm x I-8 cm,base rounded to curieate, apex obtuse to acuteor acuminate, glabrous, pinnately veined with15-25 pairs of lateral veins. Inflorescence anaxillary cluster, I-2 cm x I-3 cm, 10-30(-400-flowered; bracts very small. Flowers bisexual,regular, 5-merous, very fragrant to bad-smelling; pedice1 I-3 mm long; sepals ovate orelliptical, I-2.5(-3) mm long, free or connate atbase, apex acute to rounded; corolla tube al-most cylindrical, 6-10 min x 0.5-2 min, with a

.

452 TIMBERSl

belt of hairs I-3 mm wide insidejust below theinsertion of the stamens, lobes ovate to almostorbicular, 1.5-4.5(-5) mm long, apex roundedto acute, spreading, recurved later, white toyellow-orange; stamens inserted just below thetop of the corolla tube, included, I-2 min long,arithers ovate, yellow; ovary superior, ovoid toglobose, consisting of 2 separate carpels unitedat base by a disk-like thickening, style 4-8 minlong, pistil head ellipsoid to ovoid, 0.2-I minlong. Fruit consisting of 2 globose to ellipsoidfollicles 13-23(-30) mm long, apex pointed torounded, yellow to orange, smooth to slightlyrough, 2-seeded. Seeds ellipsoid to oblong, 6.5-13.5 min long, brown.Pieiocu, per comprises about 5 species and isconfined to mainland tropical Africa. It is re-lated to Hunterto and Pierolimo

Pieiocorpo pycnontho can be found floweringand fruiting throughoutthe year

Ecology Pieiocorpo pyc, 10ntho occurs in theunderstorey of rainforest, gallery forest andmontane forest, up to 2300 in altitude. It canalso be found in disturbed forest

Genetic resources and breeding As Pieio-corpo pycnonthu is widespread and occurs invarious forest types, there is no reason to con-sider it threatened by genetic erosion

Prospects It is unlikely that Pieiocorpopyc-noritho will become an economicalIy importanttimber tree in the future because its size is

usually too small. However, its hard and dura-ble wood will remain useful for the construc-

tion of local houses and the production of jin-PIements, as long as natural stands of suffi-cient volume are available

Major references Burki11, 1985; CoatesPalgrave, 1983; 0min0, 1996

Other references Adjanohoun at a1. , 1989;Gorman at a1. , 2004; Neuwinger, 2000

Authors R. H. M. J. Lemmens

I.

"

Podocu, pus lotifo!jus - wild

from southern Sudan through eastern CentralAfrica and East Africa to Zimbabwe, Mozam-bique, Swaziland and eastern and southernSouth Africa, where it descends to sea-level. Itis also found in the highlands of south-easternNigeria and western Cameroon, and has beenrecorded from an iriselberg in EquatorialGuinea. It Is occasionally planted as ornamen-taltree within its natural area of distribution.

Uses The wood, often traded as 'podo', ishighly valued for furniture and ship building,e. g. for masts and planks, but it is also used forpoles, panelling, boxes, veneer and plywood. Itis popular for making butchers' blocks becauseit is fairly hard, without scent, and does notchip easily. It is suitable for construction, floor-ing, Jolnery, interior trim, vehicle bodies, rail-way sleepers, toys, novelties, agricultural jin-PIements, musical instruments, coffins, foodcontainers, vats, carving, pattern making,matches, turnery, hardboard and particleboard. It is also used as fuelwood, and it is con-sidered a high-quality pulpwoodPeople occasionally eat the fleshy receptaclesat the base of the seeds. In Kenya a bark infu-SIon is taken to treat stomach-ache. In Ugandaa leaf decoction is taken as vermifuge. In theMau region in Kenya the bark is used as wa-terproof cover for bee-hives. Podoco, pus loti-10nus is planted as ornamental and roadsidetree, having a characteristic appearance withits conical shape and bright green glossyleaves, and also as shade tree in coffee, cocoaand banana plantations. It is the national treeof South Africa

Production and international trade In thepast, the wood of Podocorpus lotifolius was in

PODOCARPUSLATIFOLIUS (Thunb. ) R. Br. exMirb.

L.

o

,^

Protologue Mein. Mus. natl. Hist. nat. , Paris13: 75 (1825).

Family PodocarpaceaeSynonyms Podocorpus milonjionus Rendle

(1894).Vernacular names Real yellowwood, East

African yellowwood, broad-leaved yellowwood,red-fruited podo (En). Pinho do muxito (FD)

Origin and geographic distribution Po-docorpus lotifolius occurs in mountain forest

..

.

high demand in southern Africa, especially forflooring, furniture and railway sleepers, and itwas harvested on a large scale. At present, theinternational trade in Podocorpus lotifoliustimber is very limited. In South Africa it hasbeen estimated that on average 750 in31year ofyellowwood roundwood (from Afrocorpus andPodocorpus spp. ) is harvested, of which Podo-corpus lotifo!jus probably constitutes the majorpart. The volumes sold by auction in SouthAfrica in 1997 and 1998 were 225 ing and 260

ms. respectively. There Is no information onproduction and trade from elsewhere, but inmany countries illegal logging of this valuabletimber species continues.

Properties The heartwood is pale yellowishbrown, and not demarcated from the sapwood.The grain is straight, occasionally spiralIy,texture fine and even. Reddish streaks of coin-

pression wood may be present. Resin is absent,and the wood has no distinctive odour

The wood is moderately lightweight, with adensity of 460-510(-610) kg/in3 at 12% ino1s-ture content. It usually air dries without prob-Iems, but distortion, end splitting and checkingmay occur; close stacking and heavy weightingare recommended. Kiln drying should be doneat lower temperatures. It has been recoin-mended to air dry for 4-5 weeks, followed by5-6 days of kiln drying to obtain optimal re-SUIts. The rates of shrinkage from green tooven dry are 2.0-3.9% radial and 3.7-6.1%tangential. Once dry, the wood is stable in ser-

resistant to acids. The wood of Podocarpuslotifolius from Sudan has a high Iignin contentand was found difficult to pulp.Several cytotoxic norditerpene dilactones havebeen isolated from the bark of Podocorpus lott-101ius, while the Iactone mumakilactone B hasbeen isolated from the wood, and heveaflavone,amentoflavone and derivatives from the leaves.

Adulterations and substitutes The wood

of Afrocorpus 101cotus (Thunb. ) GNPageclosely resembles that of Podocorpus lotifoliusand is used for similar purposes; it is alsoknown as yellowwood.

Description Evergreen, dioecious, medium-sized to fairly large tree up to 35(-40) in tall, athigh altitudes a shrub or small, stunted tree;hole branchless for up to 10(-20) in, usuallystraight and cylindrical, up to 150(-300) cm indiameter, sometimes with buttresses at base;bark surface greyish brown to dark grey,rather smooth, flaking in long strips, or longi-tudinally fissured, inner bark pale pink; twigsgrooved by decurrent leaf bases. Leaves ar-ranged spiralIy to nearly opposite, simple andentire; stipules absent; petiole indistinct, short;

PODOCARPUS 453

vice.

For wood originating from Sudan, at 12% ino1s-ture content, the modulus of rupture is 81N/mm2 modulus of elasticity 5100 N/mm2,compression parallel to grain 47 N/min2, shear16 N/min2 andJanka side hardness 6030 N

The wood is easy to saw and work with ina-chine and hand tools, having little dulling ef-fects on cutting edges. It can be planed to asmooth finish. The wood has a tendency to splitupon nailing, and pre-boring is recommended;it holds screws well. It requires support indrilling and mortising because of its brittle-ness. The gluing, painting, varnishing andstaining properties are moderately good. Steambending gives moderate results, but turningproperties are good. The peeling and slicingproperties are good; good-quality but brittleveneer can be produced from the wood. Thewood is not durable, being susceptible to bluestain, powder-post beetle, pinhole borer, long-horn beetle, termite and marine borer attacksIt is permeable to preservatives. The wood is

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Podocorpt, s lotifolius - I, tree hubit, . 2, led/; 3,twig with mole cones, . 4, trotg with seed.Redrown und udopted by Achingd Satin Nur-hornon

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454 TIMBERSl

blade linear-elliptical, (2-)3.5-10(-18) cm x5-12 min, curieate at base, acute to acuminateat apex, leathery, glabrous, glaucous green todark green, with a single main vein and sev-eral resin canals. Male cone axillary, solitary ormore rarely in groups of 2, nearly sessile, I-5cm x 3-5 mm, pinkish to reddish; scales many,arranged spiralIy, each bearing 2 pollen sacs.Female cone terminal on short, leafy or scalybranchlet, solitary, with I-2 fertile scales, atbase with swollen, fleshy receptacle 8-18 minlong, glaucous green turning pink to reddishpurple or blackish purple. Seeds drupe-like,globose to obovoid, 7-12 min long, glaucous togreyish green, seed coat slightly woody, en-closed in a leathery integument.

Other botanical information Podocarpuscomprises nearly 100 species and is widelyspread in the southern hemisphere, extendingnorthward to Central America, southern Chinaand southern Japan. In mainland Africa about3 species occur, and in Madagascar approxi-mately 5. Afrocu, :pus has been separated fromPodocu, :pus in 1988, mainly based on the ab-sence of a fleshy receptacle at the base of theseed and on chromosome number, but there isstill disagreement about this. However, ino-Iecular data support the recognition of Afrocor-pus as distinctfrom Podocarpus.Podocarpus madogoscuriensis Baker is en-demic to Madagascar, where it occurs in forestin the mountains at 800-2400 in altitude. It is

a smalltree up to 15(-30) in tall, with bole upto 30(-60) cm in diameter. The greyish yellowwood is used in local house construction, andfor flooring and joineryIn South Africa there are two other Podocarpusspp. , the wood of which is used for similar pur-poses as that of Podocorpus lotifolius: Podo-corpus e!origotus (futon) L'Her. ex Pers. en-demic to the Western Cape, and Podocorpushenhe!it Stapfex Dallim. & A. B. Jacks. endemicto Kwazulu-Natal and the Eastern Cape. Podo-corpus errsiculus MeIville, described from Tan-zania, may be a synonym of Podocarpus henke-lit

chyma diffuse (evenly scattered throughouttheentire growth increment)); 74: axial paren-chyma tangentially zoriate; 76: transverse endwalls smooth. Ray composition: (79: ray tra-cheids commonly present); 80: ray tracheidsabsent or very rare; 81: cell walls of ray tra-cheids smooth; 85: end walls of ray paren-chyma cells smooth (unpitted); 87: horizontalwalls of ray parenchyma cells smooth (unpit-ted). Cross-field pitting: (91: cross-field pitspinoid); (93: cross-field pits cupressoid); 98: I-3pits per cross-field (earlywood only). Ray size103: average ray height medium (5-15 cells);107: ray width exclusively uniseriatere. Baas & I. Hem")

Growth and development Growth of Po-docorpus lotifolius is slow under natural condi-tions; for trees with hole diameters between 10cm and 90 cm, mean diameter Increment is0.2-0.3 cm/year. In cultivation the growthrates are usually also quite low, but they de-pend on soil fertility and available water. For32-year-old trees in Rwanda a height growth of30-56 cm/year has been recorded, and a diame-ter growth of4-13 min/yearTrees 8-10 in tall may already fruit. In Kenyathe fruiting period is in March-April. Monkeys,10eries and birds such as bombills and turacos

feed on the fleshy receptacle at the base of theseed, and they usually discard the seed un-damaged, serving in this way as dispersers.However, most seed does not get far from themother tree. Bushpig eat receptacles and seedsafter falling. However, it has been reportedthat seed that has passed the digestive tract ofbushpig has a very low germination rate, 2% incomparison with 70% in control seeds. Beenests are often found in hollow Podocorpuslotifolius trees. At higher altitudes, above 2600in, the trees are usually stunted and rarelytaller than 10 in, and often covered with 11-chens and mosses; on exposed mountain sides,they may be shrubs that barely exceed 2 in inheight

Ecology In East Africa Podocorpus lotifoliusforest the mountains atInoccurs In

900-3200(-3500) in altitude. It prefers an an-nual rainfall of 1000-2000 mm and a mean

annual temperature of 9-19*C. It is oftendominant at higher altitudes in more humidlocalities, usually preferring sites with higherannual rainfall in comparison with Afrocorpusfoieutus. Podocorpus lotifolius may be doinI-nant in the climax vegetation of the lower andmiddle part of the montane zone, together with01eo spp. Locally it occurs in nearly pure

Anatomy Wood-anatointcal description (IAWAsoftwood codes):Growth rings: (40: growth ring boundaries dis-tinct); (41: growth ring boundaries indistinct orabsent); (43: transition from earlywood to late-wood gradual). Tracheids: 44: tracheid pittingin radial walls (predominantly) uniseriate (ear-Iywood only); (56: torus present (pits in early-wood tracheids only)). Axial parenchyma: 72axial parenchyma present; (73: axial paren-

.

stands, especially above 2600 in altitude, and itcan also be associated with bamboo. In south-

ern Africa it can be found from sea-level to

mountain forest up to 2300 in altitude. InSouth Africa it is characteristic of the so-called

Knysna forest, an evergreen subtropical type offorest of the southern Cape, comprising about60,000 ha. In Equatorial Guinea PodocorpL, slotifo!jus has been found in forest on rockyslopes on an iriselberg at 750 in altitude. Podo-corpus lotifo!jus is susceptible to drought. Ittolerates light frost, but young seedlings aresusceptible. It is characterized as a mediumshade-tolerant species. It performs best onwell-drained, deep, fertile soils. In cultivationit benefits from a layer of mulch on the groundStudies in South Africa showed that natural

regeneration of Podocorpus lotifolius is abun-dant, especially in small and medium-sizedgaps in the forest.

Propagation and planting The 1000-seedweight is 360-600 g. The fleshy receptacleshould be removed from the seed. Fresh seedsshould be cleaned and sown within 4 days in amixture of well-rotted compost and washedsand. They germinate in I-2 months, and thegermination rate is up to 80%. Cracking of theseed coat has been recommended to speed upgermination. Seeds can be stored in a cold storefor up to I year without losing much of theirgermination capacity, but moisture loss shouldbe avoided. Seeds stored in perforated polyeth-viene bags with damp sawdust showed a ger-mination rate of 72% after one year. Whentransplanting, care should be taken not todamage the taproot. The seedlings should bewell watered. It is recommended to plant inshade and not in the fulllight, but in a test inSouth Africa seedlings grown without shadeshowed the best growth. Wildlings are some-times also collected for planting.

Management Young trees up to pole sizecan be coppiced. In larger trees branches be-come drooping, and some pruning is often prac-tised. In natural forest in the southern Caperegion of South Africa, trees are selectivelyharvested under a felling cycle of 10 years, andselected according to externalIy visible criteriaof maturity

Diseases and pests A canker and die-backdisease, known as pink disease and caused byErythricium solmonico!or has been recorded inSouth Africa. It affects a wide range of treespecies. Seeds are commonly attacked by ln-sects, which may cause considerable losses

Handling after harvest Logs are suscepti-

ble to insect and fungal attacks and should beremoved from the forest and converted as soonas possible after felling, or treated with pre-servatives. Dipping in an anti-sapstain solutionimmediately after conversion and before stack-ing is needed to prevent discoloration by bluestain

Genetic resources Podocorpus lotifoltus iswidespread and in many regions within itsdistribution area quite common. However, Ithas been subject for many decades to large-scale felling and larger trees have become un-common. In Rwanda, for Instance, Podocarpuslotifo!jus has become quite rare as a result ofintensive exploitation. In Zimbabwe it occurs toosparsely to be of any commercial value. Podo-corpus lotifolius is legally protected in SouthAfrica. It is included in the IUCN Red 11st, al-though stillconsidered to be at lower risk.

Prospects Podocorpus lotifolius provides ahighly valued timber and as such deservesmore research attention. However, Its pros-pects as a plantation timber tree wlth coinmer-cial value, e. g. to substitute exotic pine planta-tions, seem to be limited because of its lowgrowth rates. The yield from natural forestmanaged on a sustainable basis Is very low.

Major references Bolza & Keating, 1972;Coates Palgrave, 1983; Geldenhuys, 1993b;Leistner, 1966; Maundu & Tengnds, 2005;Mbuya at a1. , 1994; Mtsweni, 2005; Palmer &Pitman, 1972-1974; Sommerlatte & Sommer-Iatte, 1990; Takahashi, 1978

Other references Barker, Muller & Mill,2004; Beentje, 1994; Burki11, 1997; CAB Inter-national, 2005; de Laubenfels, 1985; Fozdar,Khan & Shamsuddin, 1989; Gu6neau, Bedel &Thie}, 1970-1975; Hamill at a1. , 2000; Hemz,2004; Kabera, 1992; Katende, Birnie & TengnAs,1995; Lamprecht, 1989; Leistner, Smith &Glen, 1995; Neuwinger, 2000; Oilamb0, 1978;Palmer at a1. , 1986; Schaefer, 1990; Teal, 1984;Troupin, 1982; van Vuuren, Banks & Stohr,1978; van Wyk & Gencke, 2000.

Sources of illustration Coates Palgrave,1983; Leistner, 1966.

Authors J. M. Okeyo

PONGAMiOPSls 455

PONGAMIOPSIS PERVILLEANA (Baill. ) R. Vig

Protologue Notul. Syst. (Paris) 14(I): 74(1950)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution Poit-

.

456 TIMBERSl

gomiopsts perui!legno is endemic to Madagas-car, where it is widespread throughout thewestern part of the Island.

Uses The wood is used for house construc-tion and planks. The roots are used to makefish poison

Botany Deciduous shrub or smalltree up to14 in tall; hole up to 30 cm in diameter; barksmooth, shallowly longitudinalIy fissured, palegrey-brown; young twigs silky hairy, but soonglabrescent. Leaves alternate, imparipinnatelycompound with (9-)11-15(-19) leaflets; stip-ules and stipels usually absent; leaflets OPPo-site or alternate, elliptical to narrowly oblong-elliptical, I-4 cm x 0.5-1.5 cm, rounded or ob-tuse at base, rounded or notched at apex,thinly leathery, pubescent when young butoften glabrescent. Inflorescence an axillaryfalse raceme up to 7 cm long, 3-10-floweredwith flowers solitary or in pairs. Flowers bi-sexual, papilionaceous, sweet-scented; pedicelwith a pair of caducous bracteoles; calyx cup-shaped, 2.5-3.5 min long, 5-toothed with shortteeth, deep red, hairy; corolla pale purplish,standard circular, 8-10 min in diameter, withwhitish basal spot, wings and keel about aslong as standard; stamens 10, filaments fusedin lower half; ovary superior, I-celled, styleoften recurved or hooked. Fruit a broadly ellip-sold pod 1.5-4 cm long, inflated, with a shortbeak, leathery, thickened along the upper mar-gin, glabrescent, wrinkled, eventually splittinginto 2 valves, I-seeded. Seed ellipsoid, I-1.5 cmlong, flattened, brown to blackPortgomiopsis peru, !leano produces flowers onnew shoots with leaves stillimmature

Portgomiopsis comprises 3 species and is en-demic to Madagascar. It is related to Millettio,which differs in its flat, elongated pods andseeds with distinct anI

The wood of Poitgomiopsis Qinygdo!trio (Baill. )R. Vig. is used for construction and as firewood.This species occurs very locally in northernMadagascar and is listed as endangered in theIUCN red list of threatened species. It differsfrom Poitgomiopsis peruilleono in its largerleaflets, flowers and pods.

Ecology Poitgomiopsis peruilleono occurs indry scrubland or woodland, up to 1000 in alti-tude, often on sandy soils over limestone

Genetic resources and breeding Portgo-intopsis perui!leano is widespread and locallycommon in Madagascar and not yet endan-gered, although its habitat is declining rapidlyIn many regions

Prospects Poitgomiopsis perui!fourier will re-

main of local Importance for its wood as long asstands of some extentremain.

Major references du Puy at a1. , 2002Other references du Puy & Labat, 1998i;

du Puy & Labat, 1998j; Lewis at a1. , 2005Authors R. H. M. J. Lemmens

POUTERIAADOLFl-FRIEDERICll(Engl. )AMeeuse

Protologue Bothalia 7: 341 (1960)Family SapotaceaeSynonyms Sidero;:q. 10n odolfi-friedericii Engl.

(1913), Antngerio odolfi-friedericii(Engl. ) Robyns& G. CG. Gilbert (1947).

Vernacular names fullngeria (En).Origin and geographic distribution Poute-

rig odolfi-friedericii occurs from eastern DRCongo, southern Sudan and south-westernEthiopia south to eastern Zimbabwe

Uses In East Africa the wood (trade namesaningeria, aningre, inuria) is valued for furni-ture. It is also suitable for light construction,light flooring, interior trim, joinery, cabinetwork, boats, vehicle bodies, boxes and crates,veneer and plywood, and pulpwood. It is usedas firewood and for charcoal production. Thesweet fruit pulp is eaten raw. The seed oilisused for cooking. The tree is used as a shadetree in plantations. In Kenya a bark decoctionis drunk to treat stomach disorders. In Ethio-

pia the fruits are used as a traditional tae-nicidal drug, usually as a paste in barley POT-ridge

Production and international trade The

timber is traded on the international market,but statistics on production and export amounts

I.

Pouterio odo!It-friedericii- wild

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are not available. In 2003 in Ethiopia, the priceof primary grade Pouterio odo!It-friedericiisawn wood was Us$ 2751m3.

Properties The heartwood is pale greyishbrown with a pink tinge and not distinctly de-marcated from the sapwood. The grain isstraight or wavy, texture moderately fine tocoarse. The wood is lustrous and shows a inot-

tled figure when the grain is wavyThe wood is moderately light, with a density of450-545 kg/in3 at 12% moisture content. Dry-ing usually does not cause problems, but thereis a slight risk of distortion and checking, and atendency to blue stain, especiaUy in earlystages of air drying. Once dry, the wood ismoderately stable in service.The wood contains up to 0.3% silica and conse-quently the blunting effect on saws and cuttingtools may be high. Stentte-tipped sawteeth andtungsten-carbide tools are recommended. Thewood planes well and takes a good polish. Thenailing and screwing properties are good, andthe wood stains, paints and glues well. Goodveneer can be produced by slicing and rotarypeeling. The wood is not durable and liable toattack by fungi, termites, pinhole borers andmarine borers. It is permeable to preservativesTests in Ethiopia have shown that the expul-SIon time of the tapeworm Toenio saginoto byPouterto adolfi-friedericii fruits was about 30hours in humans. However, fruit extractsshowed moderate toxicity in mice when admin-1stered in the abdominal cavity. As a taenlcidaldrug the fruits are of comparatively poor valueconsidering their toxicity, potency and wormexpulsion time

Adulterations and substitutes The woodof Pouterto spp. from West Africa closely re-sembles that of Pouteriu odolfi-friedericii andis used for the same purposes. The wood ofsome Chrysophyllum spp. is often mixed intrade with Pouterio wood.

Description Very large tree up to 50 in tall;hole up to 150(-200) cm in diameter, branch-less for up to 27 in, straight and cylindrical,sometimes fluted, often with wide buttresses;bark surface greyish brown, shallowly fissured,inner bark pale pinkish brown, smelling un-pleasant, exuding latex; crown dense, rounded;young branches densely reddish brown hairyLeaves arranged spiralIy, simple and entire;stipules absent; petiole I-2 cm long, denselyhairy; blade elliptical to ovate-oblong, 4-22 cmx 2-8.5 cm, curieate at base, rounded to shortlyacuminate at apex, margins often mrolled,densely orange-brown pubescent on veins be-

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POUTERiA 457

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Pouterio udoni-friedericii- I, flowering branch, .2, flower, . 3, fruit, . 4, seedsRedrown grid oddpted byR. H. MJ. Lemmens

low, with glandular translucent dots that arevery obscure in full-grown leathery leaves,pinnately veined with 10-25 pairs of lateralveins. Flowers in fascicles in leaf axils, bisex-ual, regular, (4-)5-merous, fragrant; pedice1 5-10 mm long, densely hairy; sepals free, ovate toelliptical, up to 6 mm long, pubescent outside;corolla with up to 6.5 mm long tube androunded lobes up to 2 mm long, hairy at mar-gins, creamy white; stamens Inserted in upperpart of corolla tube, opposite corolla lobes;ovary superior, globose, long-hairy, 5-celled,style up to 6.5 mm long, stigma 5-lobed. Erult anarrowly ellipsoid berry up to 4 cm long, withc. I cm long beak, greenish, finely hairy, I-seeded. Seed ovoid to ellipsoid, up to 3 cm long,shiny brown, with very large scar. Seedlingwith hypogeal germination; cotyledons thickand fleshy.

Other botanical information Pouterio is

pantropical and comprises approximately 320species, about 200 of them in tropical America,120 in tropical Asia and only 6 in Africa. TheAfrican species were classified in the generaAningerio and Moldcontho, but both have beenincluded in Pouterio. Pouterio has been subdi-

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.

458 TIMBERSl

vided into 9 sections. The African species be-long to section Riotcoo, together with someAmerican species Including the well-knownfruit tree Pouterio coinpechiono (Kunth) Bae-hni(canistel or yellow sapote).The timber of Pouterio spp. is sometimes con-fused with that of Chrysophy!Ium spp. , but thelatter genus differs in the absence of translu-cent dots in the leaves and fruits containingseveral seeds. In the forest Pouterio udoni-friedericii often occurs together with Chryso-phyllum gorungosonum Engl. , of which theleaves have more closely appressed hairs be-low.

Pouterio udoni-friedericii is variable and 5subspecies have been distinguished, mainlybased on leaf characters. Most of them haverestricted distribution areas.

Anatomy Wood-anatomical description OAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; (23: shape of alternate pits polygo-na1); 25: intervessel pits small (4-7 prn); 26:intervessel pits medium (7-10 F1m); (30: vessel-ray pits with distinct borders; similar to in-tervesselpits in size and shape throughout theray cell); 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);33: vessel-ray pits of two distinct sizes or typesin the same ray cell; 41: mean tangential di-ameter of vessellumina 50-100 pin; 42: meantangential diameter of vessellumina 100-200F1m; 48: 20-40 vessels per square minimetreTracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled. Axialparenchyma: 77: axial parenchyma diffuse-in-aggregates; 86: axial parenchyma in narrowbands or lines up to three cells wide; (87: axialparenchyma reticulate); (89: axial parenchymain marginal or in seemingly marginal bands);92: four (3-4) cells per parenchyma strand; 93:eight (5-8) cells per parenchyma strand; (94:over eight cells per parenchyma strand). Rays97: ray width I-3 cells; (100: rays with multis-eriate portion(s) as wide as uniseriate por-tions); (106: body ray cells procumbent withone row of upright and/or square marginalcells); 107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-

ginal cells; (108: body ray cells procumbentwith over 4 rows of upright and/or square mar-ginal cells); 1/6: ;;: 12 rays per min. Mineralinclusions: (136: prismatic crystals present);(142: prismatic crystals in chambered axialparenchyma cells); 159: silica bodies present;160: silica bodies in ray cells.(N. P. Monel, P. Baas & A. A. Oteng-Amoako)

Growth and development The tree growsslowly. The roots are associated with arbuscu-Iar mycorrhizae. In Uganda fruits are ripe inApril, in Zimbabwe from January to April

Ecology Pouterio udoni-friedericii occurs inupland rainforest at (1000-)1200-2500 in alti-tude, where it is common in many regions. Athigher elevations it is often associated withPodocarpusspp

Propagation and planting Experiments inEthiopia revealed that regeneration in naturalforest was poor. Seeds lose viahinty veryquickly and should be sown directly after col-Iection. Sometimes wildlings are collected forplanting.

Management In Ethiopia and Uganda Pou-tend udoni-friedericii is planted in timberplantations and in degraded montane forest,but information on cultivation methods is not

available. The tree can be managed by coppic-ing and pollarding

Harvesting In south-western Ethiopia treesare still commonly felled with axes, followed bybucking into an average log length of 4 in, andcrawler bulldozer extraction

In Tanzania fruits are collected from the wild,from January to June

Yield In natural forest in south-western

Ethiopia the logging intensity is estimated at 2treesA1a with an average hole volume of 15.5ina per tree. However, the combined loggingand sawmilling recovery rate, or final yield ofproduct from a tree, was estimated at only14%. The low recovery rate is mainly due to thedifficulty of processing buttressed logs. Re-inoval of buttresses by sawing before fellingincreases timber yield considerably. The esti-mated recovery rate of delivered log volume atthe sawmill was 36%.

Handling after harvest Logs should beextracted from the forest as soon as possibleafter felling or directly treated with anti-sapstain preservatrves as they are very suscep-tible to blue stain. To collect the oil, seeds arepounded and boiled. The liquid is filtered andcooled, and subsequently the o111s skimmed off

Genetic resources In several regions FOM-tend odolfi-friedericii is under pressure be-

.

cause of large-scale deforestation of the moun-tains and selective logging, e. g. in south-western Ethiopia. This is especially alarmingsince the mountain forest in which PouterLo

udoni-friedericii occurs is floristICally the rich-est forest type in Ethiopia. In these forests, upto over 90% of logging comprises Pouterioudoni-friedericii. However, in other regions thespecies occurs locally abundantly, also in siteswhich are difficult to access for logging, andwhen the whole area of distribution is takeninto account it does not yet appear endangeredby genetic eroslon.

Prospects There is much scope for improvingyield and quality of Pouterio udoni-friedericiiveneer and sawn wood by utilizing optimalproduction techniques. In many areas, e. g. insouth-western Ethiopia, obsolete equipment Isstill used in sawmills and plywood factories,whereas felled trees are utihzed inefficiently. Ithas been demonstrated that the timber produc-tion from the same number of trees could be atleast doubled by using optimal techniques ofharvesting and processing. This could also con-tribute to the development of much neededecologicalIy sound sustaTnable managementtechniques for the remainlrig natural mountainforests in which Pouterio odolfi-friedericiiplays a prominent role

Major references Abebe & Holm, 2003;Bolza & Keating, 1972; Hemsley, 1968;Katende, Birnie & Tengnas, 1995; Ruffo, Birnie& Tengnas, 2002

Other references Beentje, 1994; Bekele-Tesemma, Birnie & Tengnas, 1993; CoatesPalgrave, 1983; Desta, 1995; Eggeling & Dale,1951; InsideWood, undated; Kokwar0, 1993;Kupicha, 1983; Lovett at a1. , 2006; Oxford For-estry Institute, 1997-2004; Sommerlatte &Sommerlatte, 1990; Tesfaye & Teketay, 2002;Wimbush, 1957

Sources of illustration Hemsley, 1968Authors R. H. M. J. Lemmens

south-western Ethiopia and eastern Kenya,and south to Mozambique.

Uses The wood is used locally for construction,implements, walking sticks and canoes, asfirewood and for charcoal production. Thesweet fruit pulp is eaten fresh. The leaves aresometimes used as fodder for livestock. Theyare also used as food wrappers. The leaves areapplied externalIy in traditional medicine inWest Africa to treat yaws, wounds and con-junctivitis, The pulped bark is administered inembrocations to treat rheumatism and as anenema to treat diarrhoea in children

Properties The heartwood is yenowish whiteand not clearly demarcated from the sapwoodThe specific gravity of the wood is 0.45. Thewood is fairly hard, durable and resistant totermites.

The leaves are well accepted as fodder bysheep, although they contain considerableamounts of tannin. The nutrient content per

100 g dry matter is: crude protein 13.3 g, crudefat 3.7 g, crude fibre 25.6 g and total ash 7.6 gPer 100 g dry matter the total phenol contentis 8.3 g of which tannin phenols 6.8 g and ex-tractable condensed tannins 6.6 g

Botany Shrub or smallto medium-sized deci-duous tree up to 25 in tall; hole up to 70(-100)cm in diameter, often twisted and fluted andwith small buttresses; bark surface greyishbrown, fissured and scaly, inner bark fibrous,whitish yellow to orange-yellow, exuding latex;crown dense; young branches densely reddishbrown hairy. Leaves arranged spiralIy, simpleand entire; stipules absent; petiole I-1.5 cmlong, hairy; blade elliptical to obovate, 12-25(-38) cm x 7-16(-20) cm, curieate to rounded atbase, rounded to slightly notched at apex,densely to slightly pubescent below, wlth glan-dular translucent dots, pinnately veined with15-20 pairs of lateral veins. Flowers in fasci-cles in the axils of current or fallen leaves, bi-sexual, regular, 5-meTous, slightly fragrant,sessile; sepals free, broadly elhptical, up to 5.5mm long, brownish pubescent outside; corollawith c. 4.5 mm long tube and rounded lobes c2.5 min long, hairy at margins, yellowish orgreenish white; stamens inserted in upper partof the corolla tube, opposite corolla lobes; ovarysuperior, globose, long-hairy, 5-celled, stylecylindrical, about as long as corolla, stigma 5-lobed. Fruit a globose to elhpsoid berry 1.5-2.5cm long, becoming red when ripe, with persis-tent style, finely hairy but glabrescent, I-seeded. Seed ellipsoid, 1.5-2 cm long, darkbrown, with rather narrow scar

POUTERiA 459

POUTERiAALNIFOLTA (Baker) Roberty

Protologue Bull. Inst. Franc. Mrique Noire15: 14/7 (1953)

Family SapotaceaeSynonyms Moldednth0 o1nifolio (Baker)

Pierre (1891), Moldedntho heude!otiono Pierre(1891).

Vernacular names Mguoguo (Sw).Origin and geographic distribution Poute-

rio Qinifolio is widespread from Senegal east to

460 TIMBERSl

Pouterto is pantropical and comprises approxi-mately 320 species, about 200 of them in tropi-calAmerica, 120 in tropical Asia and only 6 inAfrica. The African species were classified inthe genera Animgerio and Molocontho, butboth have been included in Pouterio. Pouteriohas been subdivided into 9 sections. The Mri-can species belong to section Riotcoo, togetherwith some American species including the well-known fruit tree Pouterio coinpechiono(Kunth) Baehni(canistel or yellow sapote). Inthe literature, Pouteri0 o1nifolio has beenmuch confused with other Pouterio spp. ,

mainly Poutsri0 o1tissimo (Achev. ) Baehniand Pouterio oningeri Baehni. It is character-ized by its sessile flowers and narrower seedscar. Pouterio Qinifolio usually flowers at thebeginning of the dry season

Ecology Pouterio Qinifo!to occurs in dry for-est and gallery forest, often in the transitionzone between savanna and forest, and also indisturbed forest. It is often found on rockysites. In south-western Burkina Faso Pouterio

dint/ono is most common on moister soils. InEast Africa it is often an understorey tree inlowland rainforest and deciduous forest. It isoften common, especially in West and CentralAfrica. Pouterio amifolio is a pioneer speciesthat tolerates fire. It regenerates abundantlyin regularly burned forest, where it is oftenmore common than in unhurried forest. In Be-

nin vegetation communities in which PouteriuQintfolio plays an important role are consid-ered an Indicator of sites with great potentialfor growing teak

Management Pouterto dintfolio can be ina-naged by coppicing

Genetic resources and breeding Pouterioamifolio is widespread and locally common,and not under threat of genetic erosion. How-ever, in several regions it Is considered rare,e. g. in Ethiopia, Kenya and Mozambique, andthere local populations may be threatenedVar. SOCleu"it (Lecomte) L. Gaut. , restricted toZanzibar (Tanzania), is classified as vulnerablein the 2006 IUCN Red list of threatened spe-cles.

Prospects Pouteriu Qinifolio is of limitedvalue as a commercial timber tree because ofits often poorly shaped and small-sized holeHowever, It will remain of local importance indrier regions. It deserves more attention as amultipurpose tree, providing timber as well asedible fruits, forage and traditional medicine.

Major references ATbonnier, 2000; Au-br6ville, 1964; Burki11, 2000; Hemsley, 1968;

Lovett at a1. , 2006Other references Aschfalk at a1. , 2000;

Beentje, 1994; Ganglo, Leioly & Pipar, 1999;Hawthorne, 1995; Heme, 1963; Irvine, 1961;Kupicha, 1983; Lovett & Clarke, 1998e; Neu-winger, 2000; Pennington, 1991.

Authors R. H. M. J. Lemmens

POUTERiAALTiSSiMA (Achev. ) Baehni

Protologue Candonea 9: 292 (1942).Family SapotaceaeSynonyms Sidero"ylon o1tissimt, in (Achev. )

Hutch. & Dalzie1 (1931), Antngerio giltssimo(Achev. ) Aubr6v. & Penegr. (1935)

Vernacular names mingeria (En). Aningr6blanc, anlegr6 blanc (Fr)

Origin and geographic distribution Poute-rid o1tissimo occurs from Guinea east to south-western Ethiopia, western Kenya and north-western Tanzania, and south to northern Zam-bia.

Uses The wood (trade names: aningr6,antsgr6, anIgr6, asanfena, asanfona, OSan, inu-kangu) is especially recommended for high-quahty sliced and peeled veneer. In West at-rica it is also used for light carpentry, interiorjoinery, high-class furniture and moulding. InEast Africa it is considered suitable for the

same purposes and additionally for light con-struction, vehicle bodies, musical instruments,boxes and crates, railway sleepers, toys andnovelties, turnery, and pulpwood for paperproduction. It is also used as firewood and forthe production of charcoal. In Ethiopia andUganda Poutsriu o1tissimo is used as a shadetree in coffee, banana and cocoa plantations,

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and it is considered useful as a bee plant. It issometimes planted as roadside tree.

Production and international trade Poute-rt0 o1tisstmo timber is exported as sawn woodand veneer from C6te d'TVoire and Ghana in

mixed consignments with Pouterio onInger!Baehni(synonym: Awingend robusto (Achev. )Aubr6v. & Penegr. ); however, the latter consti-tutes the larger part of the export.In 2001the export of'anlegr6' veneer from C6ted'Ivoire was 15,000 ms with an average price ofUs$ 7691m3. From Ghana 19,000 ina'asanfena'veneer was exported in 2002 (average price ofUs$ 9231m, ), 14,000 in, in 2003 (average priceof Us$ 1243/ina), and 13,000 in, in 2004 (aver-age price of Us$ 1164/ms). Cameroon exported1000 in3 'anIgr6' veneer in 2003 with an aver-age price of Us$ 1864/in3The Central African Republic exported 21,000ms'aningr6'logs in 2003, with an average priceof Us$ 8011m3. Congo exported 6000 ing 'anin-gr6' logs in 2003 with an average price of Us$17/1m3 and 4000 main 2004 with an averageprice of Us$ 1561m3. The statistics for the Cen-tral African Republic and Congo may referentirely to Poutsri0 o1tissimu because Pouteriooningeri has not been recorded for these coun-tries.

In Uganda Pouteri0 o1tissimo belongs to thecommonly harvested timber species, but fig-ures on production and trade are not available

Properties The heartwood is creamy whiteto reddish brown and indistinctly demarcatedfrom the 3-6 cm wide sapwood. The grain isstraight, sometimes interlocked, texture fine tomoderately coarseThe wood is moderately light, with a density of500-580 kg/in3 at 12% moisture content. Dry-ing usuaUy does not cause problems, but thereis a slight risk of distortion and checking and atendency to blue stain, especiaUy in earlystages of air drying. The shrinkage rates aremoderate. Once dry, the wood Is moderatelystable in service.

At 12% moisture content, the modulus of rup-ture is 90 N/min2, compression parallelto grain48 N/min2 Janka side hardness 5560 N andJanka end hardness 3780 N.The wood contains about 0.3% silica and con-

sequently the blunting effect on saws and cut-ting tools is high. Stellite-tipped sawteeth andtungsten-carbide tools are recommended. Itsometimes finishes poorly after planing or saw-ing. The nailing, screwing and slicing proper-ties are good, and the wood stains, paints andglues well

The wood is not durable and liable to attacks

by fungi, termites, dry-wood borers and marineborers. It is fairly permeable to preservatives

Adulterations and substitutes The woodof some Chrysophyl!urn spp. (e. g. Chrysophy!-Ium gigonteum Achev. ) closely resembles thatof Pouterio and is sometimes traded under thesame name, e. g. 'anfogr6', in C6te d'TVoire.

Description Large tree up to 45(-50) in tall;hole up to 200(-250) cm in diameter, branch-less for up to 30 in, straight and cylindrical,sometimes fluted, with triangular, oftenwinged buttresses up to 3 in high; bark surfacecreamy grey, irregularly fissured, inner barkfibrous, reddish brown, exuding latex; crownspreading; young branches finely hairy. Leavesarranged spiralIy, SLmple and entire; stipulesabsent; petiole up to 1.5 cm long; blade ellipti-cal to ovate-oblong, 4-16 cm x 2.5-7 cm,slightly curieate to rounded at base, rounded toshortly acuminate at apex, glabrous exceptlower side of inIdrib, with glandular translu-cent dots, pinnately veined with 11-23 pairs oflateral veins. Flowers in fascicles in leaf axils,bisexual, regular, 5-merous, fragrant; pedicel3-6 min long; sepals free, elliptical to ovate-oblong, 3.5-5.5 mm long, pubescent outside;

POUTERIA 461

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462 TIMBERSl

corolla with up to 3.5 mm long tube androunded lobes up to 2 mm long, hairy at mar-gins, creamy white; stamens inserted in upperhalf of the corolla tube, opposite corolla lobes;ovary superior, globose, long-hairy, 5-celled,style up to 4 min long, stigma 5-lobed. Fruit anobovoid to globose berry up to 2 cm long, be-coming red when ripe, finely hairy but glabres-cent, I-seeded. Seed obovoid, up to 1.5 cm long,shiny brown, with very large scar. Seedlingwith hypogealgermination.

Other botanical information Pouterio is

pantropical and comprises approximately 320species, about 200 of them in tropical America,120 in tropical Asia and only 6 in Africa. TheAfrican species were classified in the generaAn!rigerio and Moldcontho, but both have beenincluded in Pouterto. Pouterio has been subdi-vided into 9 sections. The African species be-long to section Riotcoo, together with someAmerican species including the well-knownfruit tree Pouterio coinpechiono (Kunth) Baeh-ni(canistel or yellow sapote)The timber of Pouterio spp. is sometimes con-fused with that of Chrysophy!!urn spp. , but thelatter genus differs in the absence of translu-cent dots in the leaves and fruits containingseveral seeds. Pouterio oningeri differs fromPouteri0 o1tissimu in its leaves being denselyhairy below. However, the two species are oftenconfused and mixed in the timber trade.Pouterio pseudorocemoso (J. H. Hemsl. ) L. Gaut.(synonym: Antngerio pseudorocemoso J. H. Hemsl. ;Swahili name: inkuti) is a tree up to 40 in tall,endemic to Tanzania and occurring in moistlowland and submontane forest. It is used fortimber, fuelwood, making charcoal and as ashade tree. It is classified as vulnerable in the2006 IUCN Red list of threatened species

Anatomy Wood-anatomical description CAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: (10: vessels in radial multi-PIes of 4 or more common); 13: simple perfora-tion plates; 22: intervessel pits alternate; (23shape of alternate pits polygonal); 25: interves-sel pits small (4-7 prn); 26: intervessel pitsmedium (7-10 prn); (30: vessel-ray pits withdistinct borders; similar to intervessel pits insize and shape throughout the ray cell); (31vessel-ray pits with much reduced borders toapparently simple: pits rounded or angular);32: vessel-ray pits with much reduced bordersto apparently simple: pits horizontal(scalari-form, gash-like) to vertical (palisade); 33: ves-

sel-ray pits of two distinct sizes or types in thesame ray cell; (35: vessel-ray pits restricted tomarginal rows); 41: mean tangential diameterof vessellumina 50-100 pin; 42: mean tangen-tial diameter of vessellumina 100-200 F1m; 475-20 vessels per square minimetre. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;69: fibres thin- to thick-walled. Axial paren-chyma: (77: axial parenchyma diffuse-in-aggregates); 86: axial parenchyma in narrowbands or lines up to three cells wide; (87: axialparenchyma reticulate); (89: axial parenchymain marginal or in seemingly marginal bands);92: four (3-4) cells per parenchyma strand; 93:eight (5-8) cells per parenchyma strand. Rays:97: ray width I-3 cells; (100: rays with multis-eriate portion(s) as wide as uniseriate por-tions); (106: body ray cells procumbent withone row of upright and/or square marginalcells); 107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells; 1/5: 4-12 rays per min. MineralInclusions: 159: silica bodies present; 160: silicabodies in ray cells; 161: silica bodies in axialparenchyma cells(N. P. Monel, P. Baas & A. A. Oteng-Amoako)

Growth and development In natural for-est in Gabon Poutsri0 o1tissimo trees showed a

mean annual bole diameter Increment of 3.3

mm. In the Central African Republic the an-nualbole diameter increment was 3.2 min in anon-perturbed forest, 4 mm after exploitationof the forest and 6 min after exploitation andchemical thinning. In Mbaiki forest (CentralAfrican Republic) there are 4.3 Pouteri0 o1tis-simo trees per ha, corresponding to a basalarea of 0.2 in2ftia and a timber volume of 2

mama. In C6te d'Ivoire and Ghana Pouterio

o1tissimo flowers in December-March. Mosttrees with holes over 50 cm in diameter are

capable of producing fruits. In West Africa thefruits are ripe around the start of the rainyseason. They are eaten by birds and chimpan-zees, which may disperse the seeds.

Ecology In West Africa, where the areas ofdistribution of Pouteri0 o1tissimo and Pouteriooningeri overlap in the semi-deciduous forestzone, the former generally occurs in drier foresttypes than the latter. In Ghana Pouteri0 o1tis-simo occurs in the driest types of semi-deciduous forest and is locally fairly common.In C6te d'TVoire it is found near the northernborder of the semi-deciduous forest zone and ismost common at higher altitudes. In Uganda itis common, locally even dominant, in rainforest

.

at 1000-1700 in altitude.

Propagation and planting Seedhngs areclassified as non-pioneer light demanders. A1-though they may be abundant around parenttrees, further development depends on thepresence of gaps in the forest canopy. However,research in Uganda showed that large-scalelogging operations in the forest negatively in-fluence regeneration. The 1000-seed weight isabout I kg. Seeds lose viahinty very quicklyand should be sown directly after collection

Management Testson regeneration in Kenyashowed that artificial regeneration of Pouterioo1tissimo in buffer plantations around thenatural timber production forest may beneeded to maintain the species in sufficientnumbers after logging. In cultivation, the treecan be managed by coppicing or pollarding.

Handling after harvest Logs should be ex-tracted from the forest as soon as possible afterfelling or directly treated with preservatives asthey are liable to blue stain. Assessments ofsliced veneer recovery of Pouteri0 o1tissimologs in Ghana showed 30% recovery, whichmeans that the veneer mills generate a lot ofwaste.

Genetic resources Although Pouteri0 o1tis-sting is more widespread than Pouterio onin-gen, it is less common in West Mrlca, e. g. inGhana. Moreover, it is heavily exploited inGhana as well as C6te d'TVoire. In severalcountries in East Africa it is rare or occurs only

locally, e. g. in Kenya, Tanzama and Zambia. Itis classified as a lower risk species ln the 2006IUCN Red list of threatened species, but it isexpected to become threatened within a periodof 5 years without conservation programmesIn Ghana it is considered a still common spe-cies, but under pressure from exploitation, andtherefore in need of careful control.

Prospects In recent years PouterL0 o1tissi-ino has become an important timber tree, par-ticularly for veneer production, in some WestAfrican countries (06te d'TVoire, Ghana), whileit has already been of importance in Ugandaand locally in western Kenya (Kakamega For-est) for a longer time. It will only maintain itsimportance if its exploitation Is carefully con-trolled, as it is already overexploited in many

Research should therefore focus onregions.

methods of sustainable exploitation. Studies inGhana showed that there is still scope for tin-proving yield and quality of Pouteri0 o1tissimoveneer and sawn wood by utilizing optimalproduction technlques

Major references Aubr6ville, 1964; Bolza &

Keating, 1972; Burki11, 2000; Durrieu de Mad-ron, Nasi & 06tienne, 2000; Hawthorne, 1995;Hemsley, 1968; Katende, Birnie & Tengnas,1995; Normand, 1970; Takahashi, 1978; Wim-bush, 1957.

Other references Adjei-Sakyi, 2000; Mri-can Regional Workshop, 1998f; Aubr6ville,1959d; Aubr6ville, 1962b; Beentje, 1994;Chudnoff, 1980; CTRAD Forestry Department,2003; CTFT, 1970; Durand, 1983a; ForestProducts Research Laboratory, 1955; G6rard ata1. , 1998; InsideWood, undated; ITT0, 2006;Kalema, 1994; Nama & Kiyiapi, 2001;Kupicha, 1983; Lovett & Clarke, 1998f; Lovettat a1. , 2006; Pennington, 1991; Sommerlatte &Sommerlatte, 1990; Taylor, 1960

Sources of illustration Kupicha, 1983;Normand, 1970

Authors R. H. M. J. Lemmens

POUTERIA 463

POUTERLAANiNGERIBaehni

Protologue Candonea 9: 289 (1942).Family SapotaceaeChromosome number2n=28Synonyms Antngerio robusto (Achev. ) Au-

br6v. & Fellegr. (1935)Vernacular names fujingeria (En). Aningr6

blanc, antsgr6 blanc (Fr)Origin and geographic distribution Poute-

rig oningeri occurs in the forest zone fromGuinea Bissau east to Gamero0n.

Uses The wood (trade names: animgr6, am6gr6,anigr6, asanfena, asanfona) is especially rec-ommended for high-quality sliced veneer. It isalso used for light carpentry, Triterior 101nery,high-class furniture and moulding. The fruit

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464 TIMBERSl

pulp Is edible.Production and international trade Coin-

mercial exploitation of aningr6 wood beganaround 1965. Pouterio oningeriis exported assawn wood and veneer from C6te d'Ivoire andGhana in considerable quantities in mixed con-signments with Pouteri0 o1tissimo (Achev. )Baehni. In 2001 the export of 'anlegr6' veneerfrom C6te d'Ivoire was 15,000 in3 with an aver-age price of Us$ 7691m3. From Ghana 19,000in3 'asanfena' veneer was exported in 2002 (av-erage price of Us$ 9231m3), 14,000 ms in 2003(average price of Us$ 1243/in3), and 13,000 msin 2004 (average price of Us$ 1164/ms). Cam-eroon exported 1000 ms 'antgr6' veneer in 2003with an average price of Us$ 1864/in3.

Properties The heartwood is creamy whiteto pale pinkish brown and indistinctly demar-cated from the 3-6 cm wide sapwood. The grainIs straight, sometimes slightly interlocked,texture fine. The wood is lustrous

The wood is moderately light, with a density of540-590 kg/in3 at 12% moisture content. Dry-ing usually does not cause problems, but thereis a slight risk of distortion and checking and atendency to blue stain, especially in earlystages of air drying. The shrinkage rates aremoderate, from green to oven dry 3.9-4.1%radial and 6.7-7.6% tangential. Once dry, thewood is moderately stable in service.At 12% moisture content, the modulus of rup-ture is 93-130 N/mm2, modulus of elasticity11,100 N/min2, compression parallel to grain52-57 N/min2, shear 6.8-9.5 N/mm2 cleavage16 N/mm and Chalais-Meudon hardness 2.6The wood contains about 0.3% silica and con-sequently the blunting effect on saws and cut-ting tools is high. Stellite-tipped sawteeth andtungsten-carbide tools are recommended. Thenailing and screwing properties are good, andthe wood stains, paints and glues well. Theslicing properties are good, and excellent ve-neer can be produced. The wood is not durableand liable to attacks by fungi, termites anddry-wood borers. It is fairly permeable to pre-servatives

Adulterations and substitutes The wood

of some Chrysophyllum spp. (e. g. Chrysophyl-Ium gigantet, in Achev. ) closely resembles thatof Pouterio and is sometimes traded under thesame name, e. g. 'anlegr6', in C6te d'TVoire

Description Large tree up to 40 in tall; boleup to 150 cm in diameter, branchless for up to25 in, straight and cylindrical, with triangular,often winged and branched buttresses up to 3in high; bark surface grey with whitish patches,

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fissured, inner bark fibrous, reddish brown,exuding latex; crown dense, dark green; youngbranches densely hairy. Leaves arranged spi-rally, simple and entire; stipules absent; peti-o1e c. I cm long, silky hairy; blade elliptical toovate-oblong, 8-15 cm x 4-6 cm, slightlycurieate to rounded at base, rounded to veryshortly acuminate at apex, densely reddishbrown pubescent below, with glandular trans-Iucent dots, pinnately veined with (10-)15-20pairs of lateral veins. Flowers in fascicles inthe axils of fallen leaves, bisexual, regular, 5-meTous; pedicel c. 3 min long; sepals free,ovate, c. 4 min long, pubescent outside; corollawith c. 3 mm long tube and rounded lobes c. 1.5min long, hairy at margins, creamy white; sta-mens inserted halfway up the corolla tube,opposite corolla lobes; ovary superior, globose,long-hairy, 5-celled, style c. 2 mm long, stigma5-10bed. Fruit a globose berry 1.5-2 cm long,becoming red when ripe, finely hairy, I-seededSeed ellipsoid, c. 1.5 cm x I cm, dark brown,with very large scar. Seedling with hypogealgermination; hypocoty1 0.5-I cm long, epicotyl

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7-9 cm long, hairy; cotyledons thick and fleshy;first two leaves opposite, subsequent ones al-ternate

Other botanical information Pouterio is

pantropical and comprises approximately 320species, about 200 of them in tropical America,120 in tropical Asia and only 6 in Africa. TheAfrican species were classified in the generaAntngerio and Moldcontho, but both have beenincluded in Pouterio. Pouterio has been subdi-

vided into 9 sections. The African species be-long to section Riuicoo, together with someAmerican species including the well-knownfruit tree Pouterio coinpechiono (Kunth) Baeh-in (canistelor yellow sapote).The timber of Pouterio spp. is sometimes con-fused with that of Chrysophy!Ium spp. , but thelatter genus differs in the absence of translu-cent dots in the leaves and fruits containingseveral seeds. Pouteri0 o1tissimo differs from

Pouterio oningeri in its leaves being glabrousbelow. However, the two species are often con-fused and their wood mixed in the timbertrade.

Pouterio superbo ryermoesen) L. Gaut. (syno-nyms: Molocontho superbo Vermoesen, Antn-geno superbo ryermoesen) A. Chev. ), describedfrom south-western DR Congo (Bas-Congo),closely resembles Pouterio oningeri. It has astraight hole branchless for up to 25 in and upto 100 cm in diameter, with large buttresses,and the wood is similar to that of Pouterio on-

ingeri and probably also traded internationallyin smallamounts

Anatomy Wood-anatomical description CIAWAhardwood codes):Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23?: shape of alternate pits polygo-na1; 26: intervessel pits medium (7-10 prn); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; 31: vessel-ray pits with much re-duced borders to apparently simple: pitsrounded or angular; 32: vessel-ray pits withmuch reduced borders to apparently simplepits horizontal(scalariform, gash-like) to verti-cal (palisade); (33: vessel-ray pits of two dis-tinct sizes or types in the same ray cell); 41:mean tangential diameter of vessellumina 50-100 pin; 47: 5-20 vessels per square minimetreTracheids and fibres: 61: fibres with simple tominutely bordered pits; 66: non-septate fibrespresent; 69: fibres thin- to thick-walled. Axial

parenchyma: (77: axial parenchyma diffuse-in-aggregates); 86: axial parenchyma in narrowbands or lines up to three cells wide; (87: axialparenchyma reticulate); 92: four (3-4) cells perparenchyma strand; 93: eight (5-8) cells perparenchyma strand. Rays: 97: ray width I-3cells; 107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells; (108: body ray cells procumbentwith over 4 rows of upright and/or square mar-ginal cells); 1/6:;a 12 rays per mm. Mineralinclusions: (159: silica bodies present); (160:silica bodies in ray cells); (161: silica bodies inaxial parenchyma cells)re. Mugabi, A. A. Oteng-Amoako & P. Baas)

Growth and development Germinationtakes 2-3 weeks, and initial growth is slow. InMopri forest (C6te d'Ivoire) during a period of14 years (1978-1992), the annual growth indiameter of trees in diameter class 10-70 cm

was 2.8 min in non-disturbed forest and 4.4after chemical thinning. The annual

growth in diameter was I-2 mm faster fortrees in diameter class 40-70 cm than for thin-

ner ones. In C6te d'TVoire Pouterio oningeriflowers in February-April and ripe fruits canbe found in April-May. In Ghana the tree has adeciduous period in February-March, withflowering taking place in the same period;fruits are ripe in March-April. The fruits areeaten by birds and chimpanzees, which maydisperse the seeds. Pouterio oningeri regener-ates abundantly.

Ecology In Ghana Pouterto oningeri occursin semi-deciduous forest and is locally commonIn C6te d'TVoire and Garnero0n it is reportedIymost abundant in the transition zone between

semi-deciduous forest and humid evergreenforest. Pouterio oningeriis susceptible to fire;after a fire in a forest in C6te d'Ivoire 50% ofthe trees died within 6 years

Propagation and planting Seedlings areclassified as non-pioneer light demanders. A1-though they may be very abundant aroundparent trees, further development depends onthe presence of gaps in the forest canopy. The1000-seed weightis 700-800 g.

Management Burning negatively affects re-generation of Pouterio oninger!; in Ghana alltree sizes were more abundant in unburnedthan in burned forest. Although seedlings pre-for some openings in the forest canopy forproper development, larger-scale logging op-erations in the forest negatively affect regen-eration

Handling after harvest Logs should be ex-

POUTERiA 465

min

.

466 TIMBERSl

tracted from the forest as soon as possible afterfelling or directly treated with preservatives asthey are liable to blue stain. Studies in Ghanashowed that the strength properties ofbranchwood of Pouterio oningeri comparedfavourably with those of holewood and thatlarger branches (15-30 cm in diameter), whichare usually regarded as waste, can be sawn forstructural material. Assessments of sliced ve-

neer recovery of Pouterio oningeri logs inGhana showed 30-34% recovery, which meansthat the veneer mills generate a lot of waste.Occupational asthma and contact urticariacaused by wood dust of Pouterio oningeri havebeen recorded in workers.

Genetic resources Although Pouterio onin-gents less widespread than Pouteri0 o1tissimo,it is locally common in West Africa, e. g. in C6ted'Ivoire and in Ghana, where it is more coin-mon than the latter species. However, it isheavily exploited in both countries, and moni-tonng of the populationsis recommended

Prospects In recent years Pouterio oningerihas become an important timber tree, particu-Iarly for veneer production, in some West Am-can countries (C6te d'Ivoire, Ghana). Severalcharacteristics (favourable wood properties,good regeneration) indicate that it may upholdits Importance if it is not too heavily exploited.Research should therefore focus on methods of

sustainable exploitation. Studies in Ghanashowed that there is stillscope to improve yieldand quality of Pouterio owingeri veneer andsawn wood by utilizing branchwood and opti-inalproduction techniques.

Major references Aubr6ville, 1964; Ber-tault, 1982; Durrieu de Madron at a1. , 1998b;Hawthorne, 1995; Normand, 1970; Okai, Frim-pong Mensah & Yeboah, 2004; Takahashi,1978; Voorhoeve, 1979.

Other references Adjei-Sakyi, 2000; Agya-pong, 2000; Aubr6ville, 1959d; Bertault at al. ,1999; Chudnoff, 1980; CTRAD Forestry De-partment, 2003; de in Mensbruge, 1966; Du-rand, 1983a; Oarces Sotillos at a1. , 1995;G6rard at a1. , 1998; InsideWood, undated; Ir-vine, 1961; ITT0, 2006; Of on & Appiah, 1998;Pennington, 1991; Taylor, 1960; Vermoesen,1923; Yeboah, 2000.

Sources of illustration Aubr6ville, 1964;Normand, 1970

Authors R. H. M. J. Lemmens

PREMNAANGOLENSIS Gtirke

Protologue Bot. Jabrb. Syst. 18: 165 (1893)Family Verbenaceae (APG: Lamiaceae)Synonyms Premno zenkert Gtirke (1903).Vernacular names Musalengue (Po).Originand geographic distribution Premno

origolensis is widely distributed in tropical of-rica, from Senegal to Ethiopia, Kenya, Tanza-nia and Angola.

Uses The wood of Premno origolensis (tradename: inuhorro) is suitable for construction,flooring, mine props, ship and boat building,furniture and cabinet work, interior trim, toysand novelties, agricultural implements, boxesand crates, carvings, turnery, draining boards,food containers, veneer and plywood. In Kenyait is used for carving and beehives, and it hasbeen used in construction work in mines. In

Tanzania it is used for animal traps, tool han-dles, and as firewood.In 06te d'TVoire the bark of Premno origo!errsisis used in enemas and baths to treat fever in

children. In Sao Tome at Principe the bark is aremedy for malaria and fevers. In Gabon thebark is used in enemas and in fumigationsagainst madness. In DR Congo juice squeezedfrom the crushed bark is instilled in the nos-

trils to treat epilepsy. In Tanzania a prepara-tion of the bark is taken to treat stomach pain,and sap exuded from the bark is taken againstdysentery.In Gabon Premno origolensis is taboo for somepeople, and its use in the kitchen is not al-lowed. The bad smell of leaves and twigsthrown on a fire is believed to keep away badspirits

Properties The heartwood of Premno origo-lensis is pale yellow-brown, the sapwood is onlyslightly paler. The grain is usually straight,but sometimes interlocked, texture mediumand even. The smell has been described bysome as sweet, by others as unpleasant. Thedensity of the wood is 730-800 kg/ina at 12%moisture content. Logs are usually hollow orhave rotten cores. The wood seasons well, withlittle distortion, but some checking may occur.The wood works and planes well. It is recordedto be suitable for peeling and slicing, holdsnails well, glues satisfactorily, and paints, pal-ishes and varnishes well. The steam-bendingproperties are moderate to poor. The wood isdurable, due to the presence of oil. The sap-wood is not susceptible to Lyetus borer attack

Botany Smallto medium-sized tree up to 21(-33) in tall, less often shrubby; hole up to 120 cm

.

in diameter, often crooked, sometimes fluted,usually hollow; outer bark pale grey or reddishgrey, finely furrowed; crown spreading, withmore or less horizontal branches; branchletssparsely pubescent and glandular, becomingglabrous. Leaves mostly in whorls of 4, lessoften opposite, simple and entire; petiole 3-10cm long; blade ovate, oblong or elliptical, 4-21cm x 3-13(-17) cm, base curieate or rounded toslightly cordate, apex acuminate, glabrousabove, pubescent beneath on main veins, glan-dular punctate. Inflorescence a large thyrsoidpanicle, terminal or axillary, 5-30 cm long, themain branches in whorls of 2-5 and 3-9 cm

long; bracts linear to lanceolate, up to I cmlong. Flowers bisexual, zygomorphic; pedicelupto I min long; calyx 1.5-2 mm long, truncate orobscurely 2-lipped, pubescent or glabrous, per-SIStent; corolla 4-10bed, white, glabrous outside,tube 2-2.5(-3) min long, lobes c. I min long;stamens 4, inserted in corona tube, 2 longerand 2 shorter; ovary superior, 2-celled orfalsely 4-celled, style subulate, shortly 2-lobedat apex. Fruit a globose drupe 4-6 min in di-ameter, apex acute, green turning purple; en-docarp bony, few-seeded. Seeds oblongPremno comprises about 225 species, distrib-uted mainly in the Old World tropics and sub-tropics. Various other Premno spp. are localsources of timber in tropical Africa. The woodof Premno hi!debrondtii Gtirke, distributed inKenya and Tanzania, is hard and used forbuilding poles, tool handles and as firewood.The foliage is browsed by goats, and a decoc-tion of the root is used as a medicine for stom-achache. The wood of Premno mooiensis (Pear-son) Pieper, distributed in Mozambique andSouth Africa, is durable in the ground and hasbeen used for fencing poles. The wood ofPremno sch!iebenii Werderm. , distributed inTanzania and Mozambique, is tough and hard,and is used for building poles, toolhandles andas firewood; it is unclear, however, whetherthis should be considered a separate specles orbe included in Premno chrysoclodo (Boj. )Gtirke. Premno schliebeniiis classified as vul-nerable on the IUCN red list of threatened

specles.

In Kenya Premno origolensis flowers inApril-May.

Ecology Premno origo!errsis occurs up to 2100in altitude, in forest, bushland and grasslandIn forest it occurs mainly in margins and clear-IngS.

Genetic resources and breeding In viewof its wide distribution Prem, IQ origo!errsis is

notthreatened with genetic erosionProspects Premnu origolensis is a useful local

source of wood for various purposes, Includingconstruction. Commercially the wood has littlepotential, because logs are often hollow; so Itsimportance is unlikely to increase

Major references Bolza & Keating, 1972;Burki11, 2000; Fernandes, 2005; Lovett at al. ,2006; Verdcourt, 1992.

Other references Beentje, 1994; Chifun-dera, 2001; Coates Palgrave, 1983; Do C6u deMadureira at a1. , 2002; Eggeling & Dale, 1951;Huber, Hepper & MeIkle, 1963; Lovett &Clarke, 1998g; OSolo at a1. , 1996; Raponda-Walker & Sinans, 1961; Wimbush, 1957

Authors M. Brink

PREMNA 467

PREMNAMAxiMAT. GE. Fr.

Protologue Notizbl. Bot. Gart. Berhn-Dahlema 700 (1924)

Family Verbenaceae (APG: Lamiaceae)Origin and geographic distribution Premno

maximo is endemic to Kenya, where it is con-fined to a few smalllocalities near Meru andMarsabit

Uses The wood of Premno morimo Is suit-able for construction, poles and pites, lightflooring, mine props, ship and boat building,furniture and cabinet work, joinery, toys andnovelties, agricultural implements, boxes andcrates, vats, turnery, draining boards, foodcontainers, veneer and plywood. In Kenya thewood is used in carpentry and construction. Ithas also been used for flour mills, stave pipingand cooling-tower SIats

Properties The heartwood of Premnoindrimo is grey-brown or grey tinged wlthgreen, and not clearly demarcated from thesapwood. The grain is straight, texture me-dium and even. The density of the wood is 560-740 kg/ina at 12% moisture content. The woodseasons well, but sharp moisture gradientsmay occur. The wood is hard and stable, but itsstrength is below average for its density. Itworks well with all tools, and paints, polishesand varnishes satisfactorily. The wood is dura-ble due to the presence of oil. The sapwood isnot susceptible to Lyctus borer attack.

Botany Medium-sized tree up to 30(-40) intall; hole up to 90 cm in diameter, generallyfluted and bent or crooked; outer bark dark,rough, flaking vertically in small scales;branchlets first rusty brown hairy, but soonglabrous. Leaves opposite or ln whorls of 3,

468 TIMBERSl

simple and entire; petiole 3-8 cm long, rustybrown hairy; blade broadly ovate to elliptical orrounded, (6-)7-12(-19) cm x (4-)5-10.5(-16)cm, base usually rounded, apex rounded toacuminate, slightly yellowish hairy above, greywoolly hairy beneath, sometimes glabrous onboth surfaces. Inflorescence a terminal umbel-like panicle, 12-16(-20) cm x 11-14 cm, yellowhairy, branches up to 6 cm long; bracts triangu-Iar, 5-9 min x I-4 min. Flowers bisexual, zy-gomorphic; pedicel c. I mm long; calyx cam-panulate, c. 3.5 mm long, yellow hairy, 2-lipped, one lip 2-toothed, the other entire, per-sistent; corolla creamy white, narrowly funnel-shaped, tube c. 3 min long, with 4 lobes, 2-lipped, lobes rounded and c. 2.5 mm x 2 min;stamens 4, inserted in corolla tube, 2 longerand 2 shorter; ovary superior, 2-celled orfalsely 4-celled, style subulate, c. 1.5 min long,shortly 2-10bed at apex. Fruit a globose drupe c8 min in diameter when dry, purplish; endo-carp bony, few-seeded. Seeds oblongPremno comprises about 225 species, distributedmainly in the Old World tropics and subtropics.In Kenya Premno indrimo has been foundflowering from January to June

Ecology Premno indrimo occurs at 1150-1800 in altitude in moist evergreen forest.

Genetic resources and breeding The dis-tribution of Premno indrimo is limited to asingle country, and it is classified as vulnerableon the TUGN red list of threatened species. Thepopulation in the Marsabit National Reserve inKenya is protected, and Premno indrimo isbeing propagated in the Plant ConservationProgramme in Kenya.

Prospects Premno maximo has a restricteddistribution and is considered vulnerable;hence harvesting the timber from wild standsshould be discouraged

Major references Bolza & Keating, 1972;Verdcourt, 1992.

Other references Battiscombe, 1936;Beentje, 1994; Dale & Greenway, 1961; Wim-bush, 1957; World Conservation MonitoringCentre, 1998d

Authors M. Brink

Origin and geographic distribution Pseu-dobe780mo mossombtcensis occurs in coastal

regions of Kenya, Tanzania, Mozambique andnorthern SouthAfrica

Uses The wood is used for poles in local housebuilding and as a general purpose timber. It isalso used as fuelwood and for making charcoal.

Properties The wood is lightweight but du-rable. Cytotoxic 7-hydroxy sterols have beenisolated from Pseudobersomo mossombicensis

twigs and leaves. They showed toxicity againstP-388 inurine Ieukaemia cells, and some ofthem exhibited significant DNA-damaging ac-tinty.

Botany Evergreen dioecious shrub or smallto medium-sized tree up to 20 in tall; bark sur-face greyish; young branches reddish brown,slightly hairy. Leaves alternate, imparipin-nately compound with 9-17 leaflets; stipulesabsent; petiole c. 5 cm long, rachis up to 25 cmlong; petiolules up to 7 min long; leaflets alter-nate to nearly opposite, elliptical to oblong-elliptical, 3-9(-15) cm x I-4(-6) cm, curieateand often asymmetrical at base, shortly acu-inmate at apex, glabrescent but with tufts ofhairs in vein axils below, plrinately veined.Inflorescence an axillary cyme, often head-like,short-hairy, up to 12-flowered; peduncle I-6cm long. Flowers unisexual, male and femaleflowers very similar in appearance, regular, 5-meTous, whitish; pedicel c. I min long; calyxcup-shaped, c. 3 min long, lobed to the middle;petals free, 5-6 mm long; stamens 3-4 minlong, fused in lower half into a tube, hairy in-side towards the apex; ovary superior, ovoid-globose, hairy, 5-celled, style I-2 mm long,stigma obscurely lobed; male flowers with ru-dimentary ovary, female flowers with non-dehiscing arithers. Fruit an ellipsoid to globosecapsule 3-4.5 cm in diameter, densely coveredwith antler-shaped appendages c. 7 min long,red, dehiscing with usually 5 woody valves, upto 10-seeded. Seeds c. 7 min x 5 min, glossydark brown or black, covered by a bright redanIat one side

Pseudobersomo comprises a single species andis close to Trichilio. It is characterized by itsantler-shaped fruit appendages. The woodanatomy of Pseudobersomo mossombicensisdiffers little from that of some Trichi!to spp. ,but the vessels are more commonly solitaryand fewer in number in Pseudobersomo inOS-sqmbicensis

Ecology Pseudobersomo mossombicensis oc-curs in lowland coastal forest, where it is oftenfound in the understorey of moist forest types

PSEUDOBERSAMAMOSSAMBICENSIS (SIm)Verdc

Protologue Journ. Linn. SOC. , Bot. 55: 504(1956)

Family MenaceaeVernacular names False white ash (En)

.

and in forest margins. It can be found from sea-levelup to 300(-500) in altitude

Genetic resources and breeding Pseudo-be780mo mossombicensis has a rather limitedarea of distribution, and is restricted to moisterforest types within this area. This might makeit easily liable to genetic erosion as a result ofhabitat destruction, although it appears to belittle used by the local populations. In SouthAfrica it is a protected plant species

Prospects Its usually small hole size makesit very unlikely that the future Importance ofPseudobersomo mossombicensis wood will ex-ceed its actual limited usage

Major references Coates Palgrave, 1983;Gasson & Cheek, 1992; Palmer & Pitman,1972-1974; Styles & White, 1991

Other references Beentje, 1994; Gunati-Iaka at a1. , 1992; HeItzel at a1. , 1994; Lovett ata1. , 2007; Mulholland, Parel & Coombes, 2000;van Wyk & Gencke, 2000; White & Styles,1963.

Authors R. H. M. J. Lemmens

ery, furniture and cabinet making, and for con-struction. It resembles mahogany, but is heav-ier and harder. It is also used for doors, win-dows, frames, drums, barrels, canoes, mortars,bowls and gun-stocks. It is suitable for flooring,interior trim, ship building, vehicle bodies,toys, novelties, carvings, turnery, veneer andplywood. The wood is also used as firewood andfor charcoal productionPseudocedre!o hotschyi has numerous uses intraditional medicine, particularly its bark,roots and leaves. Bark decoctions or macera-

tions are applied externalIy to ulcers, sores,rheumatism, leprosy, syphilis, yaws, itch, car-Ies and gingivitis. Internally they are used totreat fever, stomach-ache, diarrhoea and dys-entery, and as a diuretic and aphrodisiac. Rootor root bark preparations are administered aspowerful diuretic, to treat asthma, fever, dys-entery, oedema, to facilitate childbirth, and areapplied externalIy to ulcers, mastitis, haemor-rhoids, fractures, rheumatism, caries and gin-givitis, and as an aphrodisiac. In Uganda a teamade from root powder is drunk to treat ITvercirrhosis. Pounded leafy twigs are rubbed in totreat headache and rheumatism, and a leafdecoction is applied as a compress to fractures,and is drunk and used in a bath against rashand oedema. Young stems and roots are coin-morily used as chewing sticks to keep the teethhealthy. In Nigeria the stem bark is used inmixtures to treat trypanosomiasis in livestock,and leaves are administered in veterinarymedicine against intestinal worms. In Nigeriathe bark is used as an ingredient of arrow poi-son, and in C6te d'TVoire as a fish poison. Thebark yields a brownish dye that has been usedin West Africa for dyeing cloth. Pseudocedrelohotschyiis occasionally planted as an ornamen-talshade tree and roadside tree. In Nigeria theleaves are used as a green manure.

Properties The heartwood is reddish brownand distinctly demarcated from the whitish topale brown sapwood of varying width. Thegrain is interlocked, texture medium and evenThe wood has an attractive figure, a faint sineUand containssome gum.The wood is moderately heavy, with a densityof about 750 kg/ina at 12% moisture content. Itshould be air dried carefully and slowly to pre-vent splitting and warping. The rates ofshrinkage are rather high.The wood saws and works well with hand andmachine tools. It planes smoothly, and nails,screws and glues satisfactorily. The wood isfairly durable, but is susceptible to attack by

PSEUDOCEDRELAKOTSCHYl(Schweinf. )Harms

PSEUDOCEDRELA 469

Protologue Bot. Jahrb. Syst. 22: 154 (1895)Family MenaceaeChromosome number 211= 56

Vernacular names Dry-zone cedar, hardcedar-mahogany (En).

Origin and geographic distribution Pseu-docedrelo hotschyt is widespread in the sa-vanna zone from Senegal east to westernEthiopia and Uganda.

Uses The wood is valued for high-class join-

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470 TIMBERSl

Lyetus and longhorn beetles and slightly liableto termite attack.

The wood contains 11monoids, including pseu-drelones. Bark extracts showed in-vitro anti-

bacterial activity, and anti-ulcer activity inrats. In tests the roots demonstrated broad-

spectrum antimicrobial activity. Dichloro-methane extracts of the root showed an-

tileishmanial activity. The extracts and theisolated compounds 7-deacetylgedunin and 7-deacety1-7-0xogedunin exhibited in-vitro activ-ity against Letshmonio donouoni, frypono-somo brucei rhodesiense, Tryponosomo cruziand Plusmodium foieiporum, with low cyto-toxic activity against the L-6 cellline. The Ii-monoid 7-deacetoxy-7-hydroxy-gedunin,lated from Pseudocedrelo hotschyi, showedanti-HIV activity. Extracts of Pseudocedre!ohotschyiinhibited lymphocyte proliferation andexhibited molluscicidal activity. Crude ethanolextracts of the leaves showed distinct

arithelmintic activity against Hoemonchuscontortus, a pathogenic nematode in small ru-minants. Extracts of Pseudocedrelo hotschyialso showed activity against Ascoris suum. Theapplication of root juice can lead to severe skinnecrosis, and care should be taken when it isapplied to e. g. ulcersThe essential oils from the stem and root bark

are exclusively composed of sesquiterpenoids6-cadinene was the main constituent (31%) inthe oil from the stem bark, cubebols in thatfrom the root bark. The antioxidant and anti-radical activities of the essential oils werefound to be low.

Description Deciduous, monoecious, smalltree up to 12(-20) in tall; hole branchless for upto 7.5 in, straight and cylindrical, up to 70 cmin diameter; bark surface grey, longitudinalIyfissured, inner bark with reddish veins; crownoblong to pyramld-shaped, usually dense;young twigs short-hairy. Leaves alternate butoften clustered at the ends of branchlets, panp-innateIy compound with 8-18 leaflets; stipulesabsent; petiole and rachis together up to30(-40) cm long; petiolules I-3 mm long; leaf-lets alternate to nearly opposite, lanceolate-elliptical, 5-15 cm x 2-6 cm, rounded andasymmetrical at base, obtuse at apex, marginsIrregularly wavy or toothed with roundedteeth, densely hairy when young, pinnatelyveined. Inflorescence an axillary panicle up to30 cm long, often several together, short-hairyFlowers unisexual, male and female flowersvery similar in appearance, regular, (4-)5-merous, whitish; pedice1 2-4 min long; calyx

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Pseudocedrelo hotschyi- I, nowering brunch, 2,mole flower, . 3, deftisced fruitSource. ' PIOre origlytique du B, rim

lobed almost to the base, c. 1.5 mm long; petalsfree, boat-shaped, 3.5-5 min long, spreading;stamens fused into an urn-shaped tube c. 3 minlong, with (8-)10 lobes, each lobe 2-fid andwith a sessile arither; ovary superior, 4-5-celled, style-head disk-shaped; male flowerswith rudimentary ovary, female flowers withnon-dehiscing arithers. Fruit a narrowly obo-void to club-shaped capsule 7-14.5 cm long,erect, brown, dehiscing with 5 woody valves,with fibres between the valves, many-seededSeeds 4-6 cm long, pale brown, winged at apex

Other botanical information Pseudocedre-

to comprises a single species, and is character-ized by its wavy-toothed leaflets and erectfruits

Anatomy Wood-anatonxical description (IAWAhardwood codes):Growth rings: I: growth ring boundaries dis-tinct. Vessels: 5: wood diffuse-porous; 13: SIm-PIe perforation plates; 22: intervessel pits al-ternate; 23: shape of alternate pits polygonal;24: intervessel pits minute (S 4 prn); 30: vessel-ray pits with distinct borders; similar to in-tervessel pits in size and shape throughout theray cell; 42: mean tangential diameter of vessel

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.

Iumina 100-200 prn; 46: s 5 vessels per squareminimetre; 47: 5-20 vessels per square milli-metre; 58: gums and other deposits in heart-wood vessels. Tracheids and fibres: 61: fibreswith simple to minutely bordered pits; 65: sep-tate fibres present; 66: non-septate fibres pre-sent; 69: fibres thin- to thick-walled. Axial pa-renchyma: 76: axial parenchyma diffuse; 78axial parenchyma scanty paratrachea1; 79axial parenchyma vasicentric; (85: axial paren-chyma bands more than three cells wide); (86:axial parenchyma in narrow bands or lines upto three cells wide); 89: axial parenchyma inmarginal or in seemingly marginal bands; 93eight (5-8) cells per parenchyma strand. Rays:98: larger rays commonly 4- to 10'seriate; (104:all ray cells procumbent); 106: body ray cellsprocumbent with one row of upright and/orsquare marginal cells; 115: 4-12 rays per mmStoned structure: 122: rays and/or axial ele-merits irregularly stoned. Mineral inclusions:136: prismatic crystals present; 137: prismaticcrystals in upright and/or square ray cells;(141: prismatic crystals in non-chambered axialparenchyma cells)(E. Uetimane, H. Beeckman & P. E. Gasson)

Growth and development In Benin themean annual growth in diameter is estimatedat 7.5 mm. Trees flower in the middle of thedry season. Fruits take almost one year toripen. The seeds are dispersed by wind

Ecology Pseudocedrelo hotschyioccurs in sa-vanna woodland and woody grassland, up to1200 in altitude, on heavy and poorly drainedsoils. Trees are often subject to fire, but arevery resistant and survive annual fire for over50 years, as shown in a long-term experimentin central C6te d'Ivoire. In Uganda it has beenobserved that there is often profuse regenera-tion in the rainy season, and it has been sug-gested that the fires of the dry season improvethe germination of the seeds. On the otherhand, it has also been reported that seed isdestroyed by fire and that regeneration in 10-canties that are subject to regular fire is 11m-ited to root suckers

Propagation and planting The 1000-seedweight is about 230 g. The seeds lose viahintyrapidly, and should be sown soon after conec-tion. Immersion in hot water and soaking forone nightimprove the germination results. Theseeds can be stored for up to 2 months insealed containers in a cool place. They are eas-11y attacked by insects and it is recommendedthat ash is added when they are stored. Seed-lings have a long taproot, making transplant-

ing difficult. The plant develops root suckers,by which it can spread fairly well, resulting inclumps of trees

Management In Uganda Pseudocedrelo hot-schyiis grown in pure stands or mixed withother trees such as mango and cashew nut. Thetree can be managed by coppicing and pollard-ing. In natural stands, the bole is often low-branching and stunted because of fire damage,and this has great influence on the timberquality and quantity.

Harvesting For sustainable exploitation innorthern Benin, a minimum felling diameter of55 cm Is required

Genetic resources Pseudocedre!o hotschyz

is widespread and locally common or even gre-garious. However, it is irregularly distributedand in some regions within Its distributionarea uncommon or even absent.

Prospects Pseudocedrelo hotschyiis an Jin-portant savanna tree, providing not only tim-ber, but also firewood and shade; it is also animportant source of ingredients for local medi-cine. Some protection from fire is needed toallow the trees to develop straight, high-branching holes, which then produce timber ofexcellent quality. Uncontrolled collection of thebark and roots for medicinal purposes maylocally threaten populations, and methods ofcorrection that have no adverse effects shouldbe developed. Preliminary screening for phar-macological activities for the development ofmodern plant-based drugs showed promlsingresults.

Major references Arbonnier, 2004; Bolza &Keating, 1972; Burki11, 1997; Katende, Birnie& Tengnas, 1995; Neuwinger, 1996; Neu-winger, 2000; SEPASAL, 2007b; Styles &White, 1991.

Other references Adjanohoun at a1. , 1989;MTUa at a1. , 2007; Akah at a1. , 2001; Ako6gni-nou, van der Burg & van der Maesen, 2006;Anbr6ville, 1950; Aubr6ville, 1959a; Boyom ata1. , 2004; Emaruk & Deogracious, 2006; Geer-ling, 1982; Hay at a1. , 2007; InsideWood, un-dated; Kon6 at a1. , 2004; Kon6 at a1. , 2005;Kpakote at a1. , 1998; Louppe, Ouattara &Coulibaly, 1995; Nacoulma-Onedraogo & Mil-logo-Rasolodimby, 2002; Sokpon at a1. , 2006;Staner & GIIbert, 1958; Styles & White, 1989;Tabuti, Lye & Dhillion, 2003; Takahashi, 1978

Sources of illustration Ako6gninou, vander Burg & van derMaesen, 2006.

Authors R. H. M. J. Lemmens

PSEUDOCEDREIA. 471

472 TIMBERSl

PTAERoxYLONOBLIQuuM (Thunb. ) Radlk

Protologue Sitz. -Ber. Bayer. Akad. 20: 165(1890)

Family RutaceaeVernacular names Sneezewood (En). Mwan.

dara (Sw)Origin and geographic distribution Ptoero-

"y!on objiquum occurs from Tanzania and an-gola south to SouthAfrica.

Uses The wood is highly valued for furnitureand for poles in house building. In Mozambiqueit is favoured for making the keys of traditionalxylophones, and for this purpose it is baked inan oven. It has also been used for railwaysleepers and durable fence posts. It is suitablefor heavy construction including marine works,heavy flooring, vehicle bodies, handles, sport-ing goods, Implements, toys, novelties, preci-SIon equipment, carving, pattern making, vatsand turnery. It is used as firewood.The bark, the sawdust and the smoke fromburning wood are used as a snuff againstheadache. Bark and wood infusions are consid-ered remedies for rheumatism, arthritis andheart complaints. The wood resin is applied towarts and used to kill ticks on cattle. In Na-

inIbia tea made from the twigs is used againsturinary complaints. Wood chips or dust areused to repel moths from clothes. In Tanzaniathe smoke from burning wood is used as tradi-tionalpesticide for stored grain.

Properties The heartwood is rose-red todark red, changing to orange-brown or goldenbrown on exposure, and distinctly demarcatedfrom the pale grey, narrow sapwood. The grainis wavy, texture fine. Growth rings are distinct.The wood has a satiny lustre and a strong pep-pery smell; it contains oil that makes it veryinflammable

The wood is heavy, with a density of about1000 kg/ina at 12% moisture content. It airdries satisfactorily if drying is done carefullyThe rates of shrinkage are moderate, fromgreen to oven dry 3.6% radial and 5.6% tangen-tial. Once dry, the wood is stable in service. At12% moisture content, the modulus of ruptureis 141-150 N/mm2, modulus of elasticity17,600-17,800 N/mm2 compression parallel tograin 82-88 N/mm2, shear 14-15 N/mm2Janka side hardness 13,700 N and Janka endhardness 13,650 NTaking into account its hardness, the wood isnot difficult to saw, but it is difficult to workbecause of its wavy grain. However, it can befinished to a smooth and lustrous surface. The

turning properties are excellent. Pre-boringbefore nailing is necessary. Gluing is difficultThe wood is extremely durable and resistanttotermite, Lyetus and marine borer attacks. TheWood dust is Very Irritating and may causeviolent sneezingThe wood and leaves contain chromones and

other phenolic compounds. Some of these, me-thylalloptaeroxylin and perforatin A, showedantihypertensive effects. 7-Hydroxychromoneshave anti-oxidant activity. Ptoeroxy!ono61iquum lacks Iimonoids, which are commonlyfound in Rutuceoe. All alkaloid isolated from

the bark showed cardiac-depressant activity.Dichloromethane extracts of roots, leaves andstems showed moderate in-vitro antiplasmodialactivity

Botany Dioecious, usually deciduous shrub orsmall to medium-sized tree up to 20(-45) intall; hole often straight and cylindrical, up to50(-120) cm in diameter; bark surface whitishgrey, longitudinalIy fissured in large treesLeaves opposite, panpinnately compound with3-8 pairs of leaflets; stipules absent; petioleflattened, rachis usually ending in a shortinucro; leaflets nearly sessile, very asymmetri-cally oblong-lanceolate to oblong-ovate, 2-6 cmx 0.5-3 cm, curieate at base, notched to obtuseor slightly acuminate at apex, margin entire,densely short-hairy when young but glabres-cent, pinnately veined with closely spaced Iat-eral veins. Inflorescence an axillary contractedpanicle up to 5 cm long. Flowers unisexual,regular, 4-merous; pedicel up to 6 mm long;sepals nearly free, ovate, c. I long,sparsely hairy; petals free, oblong, c. 5 min x1.5 mm, pale yellow; male flowers with 4 sta-mens c. 3.5 mm long and minute rudimentaryovary; female flowers with rudimentary sta-mens, ovary superior, lateralIy compressed, 2-celled, with rather short style and 2-lobedstigma. Fruit an oblong capsule c. 2 cm x I cm,notched at apex, reddish brown, reticulatelyveined, dehiscent with 2 valves, 2-seededSeeds with a large terminal wing, c. 16 mm x 6

Ptoeroxylon obliqut, in grows moderately fast:40-100 cm/year in height under good condi-tions. In southern Africa trees flower in Au-gust-December when the trees are stillleaf-less, usually just before new leaves develop. Aflowering tree can be a spectacular sightFruits ripen about 2 months after flowering,and ripe fruits remain on the tree for sometime.

Ptoeroxy!on comprises a single species. It has

min

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.

been placed in Menuceoe and Sopindoceoe andin more recentfloras in Rutoceoe. In the 1970'sit was excluded from Rutoceoe into a separatefamily Ptoeroxyloceoe, together with Cedrelop-sis from Madagascar and later also Boltegoofrom East Africa, but a more recent phyloge-netic analysis of molecular data indicated thatit is better included in an enlarged Rutoceoe.

Ecology Ptoeroxy!on objiquum occurs in dryevergreen forest, often together with Podocor-pus and Juniperus, and in bushland, from sea-level up to 2000 in altitude. It is drought toIer-ant and tolerates moderate levels of frost. It

accepts weU-drained sandy or rocky soils, butthrives best in shale or lime soils.

Management Natural regeneration Is oftenin forest margins, but saplings have also beenrecorded in Pinus plantations in South Africa.Regeneration may be abundant after severeopening of the forest canopy, and seedhngsmay cover the bare forest floor after distur-bance. Seed can be collected just before thefruits dehisce. Pre-treatment before sowing isnot necessary. The seed can be sown in anequal mixture of river sand and compost, andshould be covered by a thin layer of sand. Thegermination rate of fresh seed is generallyhigh, but seeds lose their viahinty rapidly,within a few months. One kg contains about30,000 seeds. It is recommended to transplantseedlings when they have 3 leaves. Root suck-ers can also be used for propagation. Trees canbe managed by coppicing, showing regrowth inabout 75% of cut stems. Heart rot has beenrecorded as commonly present in logs.

Genetic resources and breeding AlthoughPtoeroxylon objiquum is widespread, Its distri-bution area is split up Into 3 areas of differentsize: (1) near the coast in Angola and northernNamibia, (2) in north-eastern Tanzania ayestUsambara Mountains), and (3) from Zimbabweand southern Mozambique to eastern SouthAfrica. This disjunct distribution can make thisspecies liable to genetic erosion, especially inthe first two areas, which are comparativelysmall and where Ptoeroxylon objiquum seemsto be uncommon. It is a protected tree in SouthAfrica, where in the past large numbers oftrees were felled for timber and fuel, and whereit has become scarce, particularly in largersizes. In Mozambique it is in high demand forkeys of traditional xylophones and has beensubject to overexploitation

Prospects Ptoeroxylon objiquum ls consid-ered a valuable African timber tree, with greatreputation for its durability. Its high density

and hardness make it particularly suitable forspecific local applications and less so for export

However, it seems to be in need ofpurposes

conservation at present. The insecticidal prop-erties of the wood deserve more research atten-tion

Major references Archer & Reynolds, 2001;Bolza & Keating, 1972; Mbuya at a1. , 1994;Palmer & Pitman, 1972-1974; Takahashi,1978

Other references Chase, Morton & Kal-Iunki, 1999; Clarkson at a1. , 2004; Geldenhuys,1993a; Grace at a1. , 2002a; Langenhoven at al. ,1988; Neuwinger, 2000; van Vuuren, Banks &Stohr, 1978; van Wyk & Gencke, 2000; vanWyk, van Oudtshoorn & Gencke, 1997; White,1990

Authors R. H. M. J. Lemmens

PTEROCARPUS 473

PTEROCARPUSANGOLENSISDC.

Protologue Prodr. 2: 4/9 (1825)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)SynonymsPterocorpus busseiHarms(1902)Vernacular names Africanbloodwood, inuk-

wa, kiaat, inuninga (En). Ambila, urnbila, nillasonde (Po). Mninga, indarnudamu, inturnbati(Sw).

Origin and geographic distribution Ptero-corpus origolensis Is widespread in southerntropical Africa, from Angola, DR Congo andTanzania south to north-eastern South Africaand Swaziland. It has been planted on a smallscale in Kenya

Uses In southern Africa Pterocorpus origo-lensis is one of the most generally used and

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most valuable timber trees. The wood is usedfor construction, carpentry, furniture manufac-ture (tables, chairs, benches), parquet flooringand veneer, and in South Africa and Namibiaparticularly for wood carving (bowls, spoonsand walking sticks). Due to its flexibility, resis-tance and light weight it is also useful forboats, doors and windows. It is occasionallyused for firewood.

The heartwood of the roots, pounded to powder,yields a fast brownish red dye which is used inthe cottage industry in Namibia and Zim-babwe, mainly to dye palm-leaf fibres for has-ket weaving. The powder is also mixed with oilor fat to make a cosmetic pomade, which hadgreat cultural importance and still has to alesser extent in Namibia (Ovambo people),Zambia (Ndembu people) and Angola. It is tra-ditionally applied to all exposed portions of thebody, including hair, face, breasts, arms andlegs. It Is also used to dye leather clothing(skirts or aprons of cowhide or tanned cowstomach), which is the traditional form of cloth-ing for men and women alike and it still hascosmetic, medicinal and symbolic importanceThemner bark and heartwood of the trunk and

branches are also said to be used by some peo-PIe to obtain dyeThe inner bark is fibrous and is used in has-ketry. In traditional medicine the bark with itsblood-red, gummy, resinous exudate ('falsedragon's blood' or 'kino') is used as a powerfulastringent, e. g. to treat diarrhoea, heavy men-struation, nose bleeding, headache, stomach-ache, schistosomiasis, sores and skin problemsThe root is believed to cure malaria, blackwa-ter fever and gonorrhoea. The seed ash is ap-plied as a dressing on wounds and psoriasis inSouth Africa. Leafy twigs are used as fodderand flowering trees are an important source ofhoney. Pterocorpus origo!errsis is nitrogen fix-ing and planted for soil conservation, dunefixation, live fencing and as an ornamentalcasting a light shade and with attractive fruits.The bark has been used as fish poison

Production and international trade Insouthern Africa Pterocorpt4s origolensis is themost widely used timber tree. Production anddemand are huge although statistics are hardlyavailable. In 1996 about 5500 in3 was exportedfrom Cabo Delgado Province in Mozambique,and the annual export from Zambia is at least5000 ina. Most of the timber is exported toChina and Thailand. The export price of supe-nor quality sawn Pterocorpus origolensis woodfrom Zambia is Us$ 5751m3. At the beginning

of the 1990s, the annual export of timber fromSouth Africa was estimated at Us$ 650,000; atpresent it may fetch prices of over Us$ 7001m3The price of one kg of wood used for carving inSouth Africa is Us$ I, but after carving it av-erages Us$ 7Bark, chips and sawdustfor dye production areavailable in considerable amounts as by-products of the timber. In rural areas manyfamilies largely depend on the income gener-ated from the small-scale woodcraft, furnitureand basketry industries, which thus supportthe poorest sectors of communities. In someregions, e. g. north-eastern South Africa, table-ware and utensils made of Pterocorpus origo-lensis wood are common items in tourist mar-

kets and shopsProperties The heartwood is pale to dark

brown or reddish brown, often with streaks,and distinctly demarcated from the pale greyor pale yellow sapwood. In Tanzania it wasestimated that the percentage of heartwood isabout 70% for a hole of 30 cm in diameter andabout 80% for one of 50 cm in diameter. Thegrain is straight to lriterlocked, texture me-dium to coarse.

Pteroco, pus origolensis wood is relatively light,the density is 400-700 kg/in3 at 12% moisturecontent. The shrinkage rates of the wood arelow: from green to 12% moisture content 1.0%radial and 1.5-1.6% tangential, and from greento oven dry 1.7% radial and 2.5% tangential.The wood dries well but slowly, without warp-Ing and with little or no tendency to check orsplit. It takes 50-90 days to air dry 25 minthick boards from 70% to 12% moisture con-

tent, and 15-20 days by kiln drying. Once dry,it is very stableAt 12% moisture content, the modulus of rup-ture is 82-94 N/mm2, modulus of elasticity8200-9200 N/min2, compression parallel tograin 50-57 N/min2, shear 9-16 N/min2 Jankaside hardness 4450-6580 N and Janka endhardness 5380-7420 N.

The wood works well with hand and machinetools, only moderately blunting cutters;straight-grained material planes and finisheswell. The bending properties are moderate. Thewood is easy to peel and slice, and has goodgluing and excellent turning and carving prop-erties. The dry sawdust may cause irritation tonose and bronchia. The heartwood is moder-ately durable; It is moderately resistant to ter-mites and marine borers. The sapwood is liableto powder-post beetle attack. The heartwood isresistant to preservative treatment, the sap-

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wood moderately resistant. A defect reportedfrequently is a inottling of the timber with ir-regular white spots up to 5 min in width,caused by an organic component which accu-inulates locally. The spots penetrate deeplyinto the wood and are obvious especially inveneers

Recent analyses of samples of Pterocorpus on-golensis heartwood have failed to detect any ofthe red biffavonoids such as santalins and san-tarubins, which are characteristic of coinmer-cial'insoluble redwoods' from other Pterocor-

pus species. However, ISOflavonoids are pre-sent, including prunetin, inuningin, tectori-genin 7-methylether, pseudobaptigenin andangolensin, accounting for the brownish colourof the dye, but elucidation of the exact dyecomposition needs further research. The driedexudate collected from the bark contains about75% tannin, mainly composed of kinotannicacidRoot extracts are lethal to adult schistosomes

bilharzia and are coinparable tocausing

praziquantel, an efficacious antischistosomaldrug

Adulterations and substitutes The woodof Pterocorpus origolensis, which is popularbecause of its streaked appearance and supe-nor carving properties, is replaced in regionswhere it has become scarce by that of otherindigenous species and occasionaUy of plantedexotic species. However, there are a few alter-natives, e. g. Afzelio quernzensis WeIw. andBrochyloeno hulllensis O. Hoffm. , which arecommonly used for carving in Zimbabwe andKenya, respectively, and the plantedAgodirochto indied A. Juss.

Description Deciduous medlurn-sized treeup to 25(-35) in tall; hole straight, up to 50(-100) cm in diameter; bark c. 1.5 cm thick,rough and cracked into more or less rectangu-Iar blocks when old, grey to brown, exuding ared resinous sap on slashing; crown open,spreading, flat; twigs thinly to densely brownor silver-grey hairy, graduaUy becoming gla-brous. Leaves alternate, imparipinnately coin-pound with (9-)11-25 leaflets, drooping; stlp-ules lanceolate to elhptical, up to 2 cm long,furring off early; petiole 2-8 cm long, rachis 11-35 cm long, densely hairy; leaflets alternate tonearly opposite, broadly lanceolate to elliptical-oblong or elliptical-obovate, 3.5-10 cm x 2.5-5.5 cm, base rounded to slightly cordate, apexacuminate, hairy on both sides when young,with 10-14 pairs of fine lateral veins. Inflores-cence an axillary raceme 6-12 cm long, often

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developing before the new leaves, denselyhairy; bracts elliptical-oblong, up to 9 min long,caducous. Flowers bisexual, papilionaceous,fragrant; pedice1 5-20 min long, densely hairy;calyx campanulate, c. I cm long, shortly 5-lobed with 2 upper lobes connate; corolla up to2 cm long, golden-yellow to orange, standardalmost circular with crimped margin and claw,wings broad, nearly as long as the standard,clawed, keel smaller, clawed; stamens 10, con-nate but sometimes I stamen partly or coin-PIetely free; ovary superior, I-celled, stalked,hairy, style slightly curved, stigma term}nal,small. Fruit an almost circular Indehiscent pod(6-)9-12(-16) cm in diameter, c. 2.5 cm thick,on a stalk up to 2.5 cm long and with an up to 3cm broad undulate almost circular wing, pu-bescent and with a mass of plumose bristles upto 13 mm long over the central thickened part,green at first, yenow-brown when mature, IC2)-seeded. Seed asymmetrical, 10-20 mm x 7-8min x 4-5 mm, smooth, red-brown, hard. Seed-ling with epigealgermlnation

Other botanical information Pterocorpus

is a pantropical genus belonging to the tribeDaibergieoe, comprising 21 species of which 12

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occur in Africa, 6 in America and 5 in AsiaSeveral Asiatic and African species were jin-portant commercial sources of red dyes, butmost species are now much more valuable fortimber

Anatomy Wood-anatointcal description CAWAhardwood codes)Growth rings: (I: growth ring boundaries dis-tinct); (2: growth ring boundaries indistinct orabsent). Vessels: 5: wood diffuse-porous; 13simple perforation plates; 22: intervessel pitsalternate; 23: shape of alternate pits polygonal;24: intervessel pits minute ^ 4 prn); 25: in-tervessel pits small(4-7 pin); 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 rim; 46: S 5vessels 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: 80: axial parenchyma allform; 82: axialparenchyma winged-allform; 83: axial paren-chyma confluent; (89: axial parenchyma inmarginal or in seemingly marginal bands); 90:fusiform parenchyma cells; 91: two cells perparenchyma strand; 92: four (3-4) cells perparenchyma strand. Rays: 96: rays exclusivelyuniseriate; (97: ray width I-3 cells); 104: allray cells procumbent; 1/6:;;: 12 rays per mm.Stoned structure: 1/8: all rays stoned; 120:axial parenchyma and/or vessel elements sto-Tied. Mineral inclusions: 136: prismatic crys-tals present; 142: prismatic crystals in chain-bered axial parenchyma cells(L. N. Banak, H. Beeckman & P. E. Gasson)

Growth and development Although Ptero-corpus origolensis can produce ample seed (upto 10,000 fruitsrria are recorded), germinationis poor. Under natural conditions only 2% ofthe seed germinates and half of the seedlingsproduced die in the first year. After germina-tion the seedling rapidly develops severalshoots and a strong taproot, which may reachto a depth of I in in the first year. The shootsreach about 15 cm length in the first year andoften die back in the dry season. The plantsenter a suffrutex stage, in which the root ex-pands in size and lateral roots develop in thetop 50 cm of the soil, while shoots usually dieback to below ground level in the dry season.New shoots develop in the rainy season. Thisstage may lastfor 10 years (sometimes up to 25years) until the root has sufficiently developed

to allow the above-ground part of the sapling tosurvive the dry season. Many seedlings do notsurvive the surfrutex stage because of drought,burning, nutrient deficiencies marticularlyboron) and damage by browsing animals. InI-tial shoot growth of saplings forms a zigzagpattern because of the yearly dieback of the top10 cm. After the suffrutex stage, the growth isfast, up to over 2 in in one year, and the treerapidly reaches a height where it cannot bereached by most browsing animals. Comparedto other trees the saplings with a thick corkybark are extremely fire resistant, sometimessurviving temperatures of up to 450'C, andfires contribute to pruning side branches andmultiple stems. During the first decade follow-ing the suffrutex stage, height rather thandiameter increases, while in the 2nd decadethe diameter increases more rapidly. Treesstart flowering when they have a permanentstem of 15-20 years old, but full developmentof fruits usually only starts when trees areabout 35 years old. The ripe fruit weighs 5-10g, but because of the large wing wind transportis possible, usually up to 30 in from the mothertree. The spiny centre of the fruit also favoursdispersal by animals. The phenology is tightlysynchronized with the seasonality of the rain-fall and flowering starts at the beginning of therainy season. In general flowering and leafflush occur from (August-) September to No-veinber (-December), fruits ripen from Januaryto April and may remain on the tree until farinto the next flowering season, leaves are shedin May-June. Flowering is short, usually only2-3 weeks, and pollination is by insects (e. g.honey bees). Fruit development takes about 4-5 months. Trees growing on good sites in fulllightlive for up to 100 years, by which age theyare about 20 in tall, with a crown diameter of10-12 in and a hole diameter of 50-60 cm; barkthickness is 1.5-2 cm and sapwood 5 cm thickBole length will depend largely on the life his-tory of the tree, butts favoured ifthe stand hasbeen burned since the tree was about 5 yearsold and if pruning is carried out. The annualdiameter increment of the hole was estimated

5.5-8.5 min in Tanzania. In Mozambique itwas estimated that a tree of 50 cm in bole di-ameter has a mean timber volume of 1.9 inaand 5.3 msfor 80 cm diameter

Ecology Pterocorpus origolensis is typicallyfound in so-called injoinbo woodland withBrochystegio and other deciduous trees, inwooded grassland and savanna, from sea-levelup to 1650(-1800) in altitude. It requires well-

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drained, medium to light soils of low to moder-ate fertility and pH 5.5-7. In Tanzania it oc-curs preferentialIy in moister localities inwooded grassland of the coastal plains. It pre-fors a climate with well-defined wet and dryseasons, with an average annual rainfall of500-1500 mm and average temperatures of15-32'C. It is not resistant to frost, althougholder trees survive very light frosts. It is a pio-neer species in disturbed and open sites, re-quiring full sun. Rainfallis more importantthan a permanent subterranean water supplyand under conditions of exceptional coinpeti-tion for ephemeral water resources the treedoes not survive.

Propagation and plantingPterocorpus on-go!errsis can be propagated by seed and by cut-tings. A tree produces on average annually100-400 fruits and there are 3400-4000(-5000)seeds per kg. At moisture contents of 4-6%seeds can be stored cold for at least 3 yearsCollection of seed for the purpose of raisingplants in a nursery is difficult because it isdifficult to open the pods without damaging theseed and because many pods are empty (about50% of young seed aborts). Pods can be openedmanually with secateurs, which is howevertime consuming. Damaged seed usually fails togerminate, but mycorrhizal associations alsoinfluence germination. Yearly dieback, a longsuffrutex stage and damage to the root systemwhen transplanting are other nursery prob-Iems that are difficult to solve. Therefore, it iseasier to start Pterocorpus origolensis planta-tions at natural sites where plants in the suf-frutex stage are already present and fill ingaps by planting suffrutex plants correctedfrom nearby vegetation. These should be takenwith as large a taproot as is possible while Iat-eralroots can be lightly trimmed. Light is veryimportant for good growth, so planting distanceof trees should at least be 5 in x 5 in, but ulti-mately the number of trees will be 25-looma,largely depending on crown and root coinpeti-tion. Cuttings (e. g. 2 in long and at least 2 cmin diameter) can be planted at the beginning ofthe rainy season butsuccess rates vary from O-30%. Planting of truncheons 10 cm in diameterinto I in deep plant holes with some coarseriver sand at the bottom has also been recoin-mended.

Management Oncetrees are established notmuch care is needed. Management can be re-stricted to taking care that each individual treegets enough light. Protection against damageby wild animals and bush fires can enhance

growth, although PterocorpL, s origo!errsis Is oneof the most fire-resistant trees of injoinbowoodland. In western Tanzania recruitment ofnew trees in selectively logged areas was foundto be poor. The density of Pterocorpus origolen-sis was higher near the main road than atsome distance, which may be explained by re-duced damage by wild grazers and lighter bushfires in the more open vegetation near theroad. After felling most trees do not resproutfrom the stump because they have a poor cop-picing ability. In practice rotation periods varyfrom 40-75 years.

Diseases and pests Periodically, Pterocor-pus origolensis tree populations suffer from'inukwa' dieback, a badly understood tree-kilnng fungal disease which blocks up the xy-Iem. In Zambia, for example, an outbreak ofthe disease killed 40% of the trees. Fire darn-

age can result in fungal(e. g. Armingrio meneo)and borer attack of the heartwood. A largenumber of wild animals can cause damage toPterocorpus origolensis by browsing, barkstripping, stem breaking by rubbing or uproot-ing. Elephants particularly are destructive,and wild pigs are fond of suffrutex roots. Seedsare often attacked by bruchid beetles.

Harvesting In South Africa trees shouldhave a minimum diameter of 27 cm to be con-sidered harvestable as timber. Growing time toreach this diameter differs per region; in SouthAfrica it takes on average about 80 years, InZambia and Tanzania possibly 40-75 yearsThe minimum cutting diameter in Tanzaniaand Zimbabweis 25 cm.

The kirio resin from the bark can be harvested

by making incisions in the bark and collectingthe exuding sap. Roots are dug up to extractthe dye. For extracting the dye from the heart-wood adult trees should be felled, but this Isnormally only done for use astimber.

Yield Timber yield data are not availableand vary largely from slte to site. In Namibiafor certain areas annual arrowable cut was es-timated at 600 in3 per 100 kin2. In the injoinbowoodland of Tanzania, the sustained annualyield of commercial timber of Pterocorpus on-golensis was estimated at 0.33 in3 per kin2. On

from roundwood to sawnaverage, recovery

woodis about 40%

Handling after harvest Usually trees arehand cut, the logs rolled to saw pits and sawninto boards, which are brought by trucks totrain depots. For preparing the dye and cos-metic pomade, the heartwood of roots is groundto a powder or paste or chipped very finely. To

PTEROCARPUS 477

478 TIMBERSl

make the cosmetic, it is mixed with butter orvegetable oilfrom pounded seeds (e. g. ofSchin-ziophyton routonenii(Schinz) Radcl. -Sin. , Ricinuscornmunis L. , Seierocoiyo birreo (A. Rich. ) Hochst. ,Ximenio cuffro Sond. ) and it is often perfumedby adding dried and ground Octmum leaves.In Zimbabwe strips of palm leaves are boiledfor about 12 hours in a dye bath containing hotwater and Pteroco, pus origolensis bark. Afterdrying, the resulting red-brown fibres are usedto obtain coloured designs in basketry weaving.The dye obtained by extracting finely choppedroots or wood in alcoholcan be used to dye wooland cotton various shades of brown to brightred.

Genetic resources Although Pteroco, pus on-golensis is widespread in many parts of south-ern Africa, overexploitation endangers manypopulations. It is classified by TUGN in thecategory 'lower risk', but is close to qualifyingfor vulnerable. In South Africa the tree has

been protected since 1967 and a special permitis needed for cutting. In Tanzania little infras-pecific vanability has been discovered in Ptero-corpus origolensis.

Prospects Pterocorpus origolensis is a veryuseful multipurpose tree in southern Africa,producing multipurpose timber, dye, forageand medicine. Overexploitation endangersnatural populations in all countries and thecurrent harvest for timber is unsustainable,raising serious concerns about the long-termviahinty of this important hardwood species.Research for large-scale cultivation is needed,especially to improve the germination rate ofseed, shorten the suffrutex stage of plants andaccelerate growth of young trees. Sustainablemanagement systems should be developed forinjoinbo woodland in which Pterocorpus origo-lensis has an important share. The dye coinpo-sition of the heartwood of this species needsfurther research. More research is also needed

to determine whether more profit can be madefrom the chips, sawdust and bark as by-products of timber exploitation, e. g. for theproduction of dye and medicinal products

Major references Boaler, 1966; Cardon,2003; Coates Palgrave, 1983; Gillett at al. ,1971; Gomes e Sousa, 1951; Graz, 2004; Rojo,1972; Schwartz, Caro & Banda-Sakala, 2002;Surowiec, Nowik & Trajanowicz, 2004; WorldAgroforestry Centre, undated.

Other references Botha, 2005; Bryant,1968; Chakabva & Mushove, 1993; Chudnoff,1980; InsideWood, undated; Joker, Msanga &Schmidt, 2000; Palmer & Pitman, 1972-1974;

Richter & Danwitz, 2000; Rodin, 1985; Rojo &A10nz0, 1993; Schwartz & Car0, 2003; Swart &Vermeulen, 1984; Takahashi, 1978; van Wyk,1972-1974

Sources of illustration Coates Palgrave,1957; Gillett at a1. , 1971

Authors R. Takawira-Nyenya

PTEROCARPUSERINACEUS Poir

Protologue Encyc1. 5: 728 (1804).Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Chromosome number it= 8.2n = 22Vernacular names African rosewood, Sene-

gal rosewood, African barwood, African teak,African kirio tree, madobia (En). V6ne, ven,pahssandre du Senegal, kirio de Gamble, santalrouge d'Mrique, henss6 (Fr). Pau sangue (Po).

Origin and geographic distribution Ptero-corpus erinoceus is widespread in the savannazone from Senegal and Gambia to Chad andthe Central African Republic.

Uses The wood is highly valued for furnitureand cabinet work, but is also used for heavyconstruction including waterworks, parquetflooring, stairs, implements, turning, sculptur-ing and sliced veneer. It is also suitable forjoinery, interior trim, mortars, pestles, houseposts, mine props, ship and boat building, vehi-cle bodies, sporting goods, toys, novelties, inu-siCalinstruments (e. g. balafons) and precisionequipment. The roots are made into bows. Thewood is suitable for fuel and charcoal produc-tion

The heartwood is a source of a red dye, whichis used for dying cloth, the body or hair. The

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bark is occasionally used for tanning. The red-dish bark exudate ('kirio') is beaten onto clothwith a mallet to give it a glaze. Kirio is coin-monly used in traditional medicine, internallyto treat diarrhoea including dysentery, fever,gonorrhoea and intestinal worm infections, andexternalIy to treat eye complaints, ulcers andsores. Until the middle of the 20th Century,kirio was also used in North America and

Europe against chronic diarrhoea. Decoctionsor infusions of bark or roots serve for treatingbronchial infections, toothache, dysentery,menstruation complaints, anaemia, gonor-rhoea, post-partum haemorrhage, ringworminfections, leprosy, wounds, tumours and ul-cers, and as an anti-emetic, purgative andtonic. Root preparations are administered asan enema to treat venereal diseases. Leaf de-coctions are applied to treat fever, syphilis andare used as an aphrodisiac and insect repel-lent. Leafy branches are browsed by livestock,and are especially important towards the endof the dry season when not much else Is avail-able. Livestock keepers rely heavily on Ptero-corpus erinoceus trees in the woodlands of theSudanian zone.

Production and international trade There

appears to be no significant international tradefor Pterocorpus erinoceus timber, although it isimportant in local markets. It is harvestedmainly by individual artisans, but in some ar-eas small-scale timber merchants also harvestand sell the timber. Locally, it Is one of themajor timbers for furnlture and construction,e. g. in Gambia. In the dry season Pterocorpuserirtoceus fodder is locally an important coin-inodity. In 1990, over 1400 t of fresh foliage, ofwhich 78% came from Pterocorpus erinoceus,were sold in Bamako onall) as feed for smallruminants. Vendors bring loads of about 75 kgon bikes into markets in Barnako from a dis-tance of up to 50 kin and make a revenue ofabout Us$ 6-71day from this activity.

Properties The heartwood is yellowlshbrown to reddish brown, often with purplishbrown streaks, and distinctly demarcated fromthe 2-5(-8) cm thick, yellowish or pale cream-coloured sapwood. The grain is straight to In-terlocked, texture fine to moderately coarse.Fresh wood has an unpleasantsmellThe wood is moderately heavy to heavy, with adensity of (560,800-890(-940) kg/ina at 12%moisture content. It dries slowly, but with littlerisk of deformation. The rates of shrinkage aremoderate, from green to oven dry 3.0-3.5%radial and 5.2-7.4% tangential. Once dry, the

wood is stable in service.

At 12% moisture content, the modulus of rup-ture is (76-)137-198 N/min2, modulus of elas-ticity 11,500-15,700 N/mm2, compression par-allel to grain 62-80 N/min2, compression per-pendicular to grain 2.5 N/min2, shear 7-10N/min2 cleavage 20 N/mm and Chalais-Meudon side hardness 11.2The wood is rather difficult to saw and work,

requiring considerable power; stellite-tippedsawteeth are recommended. It finishes well,but picking up may occur in planing due tointerlocked grain. The wood holds nails andscrews well, but pre-boring ls needed because itis brittle. The gluing properties are often poorbecause of the presence of exudates in thewood, but the wood readily accepts stains andpolishes well. The wood turns well, and thebending properties are moderate. It is durable,being resistant to fungi, dry-wood borers andtermites; it is also resistant to freshwater or-ganisms. The wood is not permeable to pre-servatives. The energy value of the wood isabout 21,000 kJ/kgThe bark exudate quickly hardens upon expo-

It contains 30-80% kiriotannlc acid,sure

which is a strong astringent. Bark extractsshowed in-vitro antibacterial and antifungalactivities against several human pathogens. Intests they blocked the ovulation and oestruscycle of female rats through antigonadotroplcactivity. Moderate in-vitro antimalarial activ-ity was also demonstrated against strains ofPIOsmodium foieiporum. The effectivity of thebark as a wound-healing agent was confirmedin tests, and the activity may be explained bythe presence of phenolic compounds that havean effect on the complement system (part of theimmune system). The bark showed significantantioxidant activity. Water and methanol ex-tracts showed in-vitro inhibitory activitiesagainst Mycobacterium sinegmotis and Myco-60cterium tuberculosis, the latter belrig acausative agent of tuberculosisThe crude protein content (dry matter basis)decreases from 13.4-16.9% in young leaves to10.3% in dry leaves, and digestIble nitrogenfrom 10.7% to 5.8%. The feed value is 0.79, 0.74and 0.51 forage unit/kg dry matter for young,green and dry leaves, respectively. In testswith rabbits, the leaves showed good digestfoil-ity and no adverse effects on health.

Description Deciduous smalltree up to 15(-25) in tall; hole straight, cylindrical andbranchless for up to 10 in under good condi-tions but often twisted, fluted and low-

PTEROCARPUS 479

480 TIMBERSl

*---, 1.1, .' - '

' " '.'!; * :"~"' '

; :t. *\~.{;11 '*=:.....-~- ..

' 44 .;I

\

\

1.7

11...

-/-...

,.

fused into a sheath up to 8.5 min long, the up-per stamen sometimes free; ovary superior,stiped, hairy, style up to 5 min long, almostglabrous. Fruit a circular, flattened, indehis-cent pod 4-7 cm in diameter, on a stipe up to Icm long and with a papery, finely veined wingwith wavy or plaited margin, with prickles onthe seed-bearing portion, straw-coloured, I(-2)-seeded. Seed kidney-shaped, flat to slightlythickened, c. 10 mm x 5 min, smooth, red todark brown. Seedling with epigeal germina-tion; cotyledons leafy

Other botanical information Pterocorpusis a pantropical genus belonging to the tribeDo16ergieoe; it comprises approximately 30species of which about 15 occur in Africa, 10 inAmerica and 5 in Asia

Anatomy Wood-anatomical description OAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23: shape of alternate pits po-Iygona1; 26: intervessel pits medium (7-10 prn);29: vestured pits; 30: vessel-ray pits with dis-tinct borders; similar to intervessel pits in sizeand shape throughout the ray cell; 41tangential diameter of vessellumina 50-100F1m; 42: mean tangential diameter of vesselIumina 100-200 pin; 46: S 5 vessels per squareminimetre; 58: gums and other deposits inheartwood vessels. Tracheids and fibres: 61:

fibres with simple to minutely bordered pits;66: non-septate fibres present; 69: fibres thin-to thick-walled. Axial parenchyma: 76: axialparenchyma diffuse; 77: axial parenchyma dif-fuse-in-aggregates; 80: axial parenchyma an-form; 82: axial parenchyma winged-anform; 83axial parenchyma confluent; 86: axial paren-chyma in narrow bands or lines up to threecells wide; 91: two cells per parenchymastrand. Rays: (96: rays exclusively uniseriate);(97: ray width I-3 cells); 104: all ray cells pro-cumbent; 1/6: z 12 rays per mm. Stoned struc-ture: 1/8: all rays stoned; 120: axial paren-chyma and/or vessel elements stoned; 121fibres stoned. 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 Seedlings deve-lop a long taproot. They grow slowly. In Maliseedlings were only up to 15 cm tall after oneyear and up to 42 cm after two years. However,under better conditions a height of up to 25 cm21 weeks after germination has been reported

II

--..

Pterocorpus erinoceus - I, ingture leg/let, . 2, flow-ering twig, ' 3, fruitRedrown Grid udopted by AChmod Sotiri Nur-homerit

branched under poorer conditions, up to 75(-100) cm in diameter, slightly buttressed; barksurface greyish brown to blackish, fissured andscaly, inner bark yellowish brown, with reddishstreaks, exuding a reddish translucent gum onslashing; crown rounded, open; twigs denselyshort-hairy when young. Leaves alternate,imparipinnately compound with (5-)7-Ile15)leaflets; stipules linear, up to 9 mm long, hairy,falling off early; petiole 3-7 cm long, rachis (7-)10-17(-22) cm long, hairy; petiolules 3-8 minlong; leaflets usually alternate, ovate to ellipti-cal, (4-)6-11 cm x (2-)3-6 cm, base rounded toobtuse, apex obtuse to slightly acuminate withusually shallowly notched tip, thick-papery,brownish hairy when young but later glabres-cent, with 12-20 pairs of lateral veins. Inno-rescence an axillary or termlnal panicle 7-20cm long, densely brown hairy; bracts up to 6mm long, falling off early. Flowers bisexual,papilionaceous; pedice1 4-8 min long, hairy;calyx campanulate, c. 7 min long, denselyhairy, with 5 triangular teeth I-2.5 min long,upper 2 more or less connate; corolla withclawed petals, golden yellow, standard almostcircular, up to 15 min x 13 mm, wings up to 13min long, keel up to 10 min long; stamens 10,

2

mean

.

and up to 100 cm after 2 years. In northernC6te d'Ivoire planted seedlings had an averageheight of 9 cm after 3 months, 50 cm after 18months, 2.8 in after 2.5 years, 4.4 in after 4.5years and 5.5 in after 5.5 years' The fastestgrowing tree was 10 in tall after 5.5 yearsCoppiced trees may grow more than I in/year.Pterocorpus erinoceus is deciduous, becomingleaness towards the end of the dry season. Thetrees flower when leaness, usually in Decem-her-February(-April), before developing newleaves, but sometimes inflorescences developtogether with young leaves. The flowers aremuch visited by bees, which are probably re-sponsible for pollination. The tree may produceso many fruits that when the fruits are green itlooks as if the tree is covered with leaves.

Young leaves normally develop after the fruitshave ripened and have become brown. Naturalregeneration is often abundant and the speciesmay be quite invasive ifprotected from grazingfor some years. The roots have nodules contain-ing nitrogen-fixing bacteria. However, Ptero-corpus erinuceus has low nitrogen-fixation po-tential in comparison with several other legu-minous trees.

Ecology Pteroco, pus erinoceus occurs in seiru-and to sub-humid tree savanna up to 600(-1200) in altitude, in regions with 600-1200(-1600) min annual rainfaU, a long dry season(up to 9 months) and a mean annual tempera-ture of 15-32'C. It can be found on all soil

types, but prefers light to medium, free-draining, acid to neutral soils. It can surviveannual bush fires.

Propagation and planting The weight of1000 seeds is about 50 g. The germination rateof untreated seed is about 50%. Soaking inwater for 12-24 hours and treatment with SUI-

phuric acid for 30-60 minutes improve germi-nation, which starts 6-10 days after sowing,with a germination rate of over 70%. A germi-nation rate of 100% was achieved using me-chanical scarification, sowing in I% agar, Incu-bation at 21'C and a 12-hour photoperiod. Op-timal germination temperature is 25-35"CSeeds can be sown in pots or in nursery beds ata spacing of about 20 cm x 30 cm. Seedlingscan be planted out from pots or as bare-rootplants, either as stumps or as entire seedlingsThe survival rate is generally high when theseedlings are protected from livestock and wildherbivores. Suckers are developed regularlyand can be used for vegetative propagation.Propagation by cuttings has also been success-ful. Recommended spacing is 3-5 in x 3-5 in in

timber plantations and I in x 2 in In planta-tions for fodder production

Management Pterocorpuserinoceustreescanbe coppiced, and leaves from regrowth aftercoppicing are higher in nutritive value for live-stock than mature leaves. It is therefore rec-Dinmended that the tree be topped before theonset of the dry season. Trees recover quicklyfrom pollarding and coppicing. Young planta-tions should be protected from browsing until 5years old, requiring fencing

Diseases and pests The fungus Phyl!achoroprerocorpi has been reported as a pathogen,producing dark spots on leaves; It spreads byair dispersal of ascospores. Seedhngs are oftenseverely attacked by rodents and crickets.

Harvesting In the dry season trees are oftenlopped or CDPpiced for forage. CDPplcmg at aheight of 10 cm above ground has been recoin-mended as a means of harvesting wood andfodder, but a coppicing height of 50 cm has alsobeen recommended. Trees do not resprout wellwhen COPDiced at ground level. To avoid brows-ing of new growth, cutting at a height of over1.5 in seems recommendable.

Yield In dry forest in northern C6te d'Ivoire,a tree with 50 cm dbh (diameter at breastheight) yields about 0.8 in3 of timber and 1.2ms of firewood; a tree with 70 cm dbh yieldsabout 1.7 ing of timber and 2.1 ina of firewoodA feedlot of I ha can feed about 24 heads ofcattle of 250 kg

Genetic resources Pterocorpus errnoceus iswidespread and occurs abundantly in manyparts of its range, and does not appear to beliable to genetic erosion. Overbrowsing by live-stock and overexploitation by collecting foddermaterial in the dry season have been recordedfor several regions, e. g. Maliand Burkina Faso,which locally puts high pressure on the popula-tions of Pterocorpus erinoceus. Locally, cuttingfor the valuable wood seems to be on an unsus-tainable basis, e. g. in Gambia. Harvestlrig tim-her or fodder from Pterocorpus er, rigceus Isillegal in several countries, including Mali. Intrials strong growth differences have been ob-served between provenances

Prospects As a true multipurpose tree, Pte-rocorpus erinoceus is of great interest for agro-forestry systems, not only because of its valu-able products such as wood, forage, fuel andmedicine, but also because it may Improve soilfertility. Increased marketing as a timber spe-cies could easily be in conflict with its impor-tance as a source of fodder and for soilim-

provemerit, uses that are crucial to the liven-

PTEROCARPUS 481

.

482 TIMBERSl

hoods of the herders and farmers in the Sahelregion. Pterocorpus erinoceus has interestingpharmacological actlvities, including antim-ICrobial, wound-healing and antioxidant ef-fects, which deserve more research attention. Ithas considerable potential as ornamental treeAs large growth differences between prove-nances have been observed, selection andbreeding may be useful

Major references A1:palu, 1998; ATbonnier,2004; Bolza & Keating, 1972; Bonkoungou,1999; Burki11, 1995; CAB International, 2005;CIRAD Forestry Department, 2003; Roj0, 1972;Takahashi, 1978; Tour6, 2001

Other references Abreu at a1. , 1999;Anderson, Bertrand & Konandji, 1994; Ayoade,Shoremi & Aregbesola, 1998; Benie & Thieu-Iant, 2003; Benie & Thieulant, 2004; Guny,Sanogo & Sommer, 1997; Diallo at a1. , 2002;Geerling, 1985; Hepper, 1958; InsideWood,undated; Karou at a1. , 2003; Karou at a1. , 2005;Louppe, Koua & Coulibaly, 1994; Louppe &Ouattara, 1993; Neuwinger, 2000; Petit, un-dated; Rivi6re, 1978; Rousse1, 1995; Sylla at al. ,1998; Thiel at a1. , 1993; Uba at a1. , 2003.

Sources of illustration Engler, 1910Authors CS. Duvall

smoke

Cooked young leaves are eaten as a vegetable;they serve as appetizer. The foliage is an jin-portant forage for all kinds of livestock. Thebark is locally used for tanning and in medici-nal preparations to treat diarrhoea and tape-worm infections. A root decoction is appliedagainst lumbago and to treat kidney coin-plaints. Leaf macerations are used to treatheadache

Properties Logs are short and usually flutedThe heartwood is creamy white and iridis-tinctly demarcated from the sapwood. Thegrain is straight or interlocked, texture moder-ately fine. Freshly cut wood has an offensivesmell. The wood is moderately heavy to heavy;wood from Mozambique has a density of 820-920 kg/ina at 12% moisture content, wood fromZambia 700-800 kg/ina. The wood usually drieswell with little deformation. Shrinkage ratesare usually low, from green to oven dry lessthan 3% radial and 5% tangential, butthey canalso be fairly high, from green to oven dry up to5.7% radial and 9.8% tangential. Once dry, thewood is stable in service. At 12% moisture con-

tentthe modulus of rupture of wood from Zam-bia is 102 N/min2, modulus of elasticity 10,200N/mm2, compression parallel to grain 51N/min2 and Janka side hardness 9,090 N. A1-though the wood is hard and tough, it generallysaws and works well, although interlockedgrain may cause some picking up in planing. Itsplits easily, but holds nails and screws well. Itturns well. When a filler is used, the results ofpainting, polishing and varnishing are satisfac-tory. The wood is moderately durable, but re-SIStant to impregnation with preservativesThe sawdust may cause irritation in workersThe wood contains a yellowish dyeThe feed value of the leaves is fairly high: 0.77and 0.65 forage unit/kg dry matter for greenand dry leaves, respectively. The crude proteincontent (dry matter basis) decreases from19.4% in green leaves to 14.9% in dry leaves,and digestIble nitrogen from 14.9% to 10.4%

Botany Deciduous shrub or smalltree up to12(-18) in tall; hole straight and cylindrical orlow-branched, up to 70(-80) cm in diameter;bark surface pale grey to dark brown, smoothto fissured or scaly, inner bark brown, inottledwith yellow and purplish red, exuding a red-dish gum on slashing; crown narrow, dense;twigs short-hairy when young. Leaves alter-nate, imparipinnately compound with (I-)5-7(-9) leaflets; stipules linear, up to 5 mm long,hairy, falling off early; petiole (0.5-)2-3(-4) cm

PTEROCARPUSLUCENS Lepr. exGuill. &Perr

Protologue F1. Seneg. tent. I(by 228 (1832).Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Chromosome number2n=22

Vernacular names SinaU-leaved bloodwood,barwood (En). Muvilu (Po)

Origin and geographic distribution Ptero-corpus Iucens has a remarkably disjunct distri-bution area, as a result of which two subspeciesare distinguished: subsp. meerrs occurring inthe savanna zone from Senegal to Ethiopia andUganda, and subsp. untunesii(Taub. ) Rojo,occurring from southern hagola and northernNamibia to Mozambique.

Uses The wood is locally used for joinery,flooring, furniture, cabinet making and imple-merits. It is also suitable for heavy construc-tion, mine props, shipbuilding, vehicle bodies,sporting goods, toys, novelties, railway sleep-ers, veneer and plywood. Locally in the Sahelregion it is one of the preferred timbers forposts and light carpentry in houses, huts, ce-real stores and shelters. It was formerly usedfor wagon wheel rims. The wood is commonlyused as firewood, giving a hot flame and little

.

long, rachis (1.5-)3-8(-11) cm long, sparselyhairy, glabrescent; petiolules 2-6 mm long;leaflets alternate to opposite, almost orbicularto ovate or elliptical, (2-)3-8(-9.5) cm x (I-)1.5-5(-6) cm, base rounded to obtuse, apexobtuse to slightly notched, papery, brownishhairy below when young but later glabrescent,with 6-20 pairs of lateral veins. Inflorescencean axillary raceme (2-)6-12(-16) cm long,slightly hairy. Flowers bisexual, papiliona-ceous; pedice1(5-)8-15(-18) mm long; calyxcampanulate, 7-8.5 min long, almost glabrous,with 5 triangular teeth 2-2.5 mm long; corollawith clawed petals, pale yellow, standard al-most circular, up to 15 min x 13 mm, wings upto 15 min long, keelup to 12 min long; stamens10, fused into a sheath up to 10 min long, theupper stamen more or less free; ovary superior,stiped, hairy, style up to 6.5 mm long, glabroustowards the top. Fruit an obovate-elliptical,flattened, indehiscent pod 3-5.5 cm long, on astipe up to I cm long and with a papery, wavywing, glabrous, straw-coloured to greyish oryellowish brown, I(-2)-seeded. Seed kidney-shaped to oblong, flat to slightly thickened, c. 7mm x 3 min, smooth, brownPterocorpus Iucens trees often retain their foli-age until the second half of the dry season.They flower at the end of the dry season, Justbefore new leaves develop, or flowers appeartogether with young leaves. Bees commonlyvisit the flowers and probably act as pollina-tors. Fruits take about 3 months to mature.The roots have nodules containing nitrogen-fixing bacteria, but Pterocorpus Iucens is con-sidered to have comparatively low nitrogenfixation potential. In Senegal the contributionof nitrogen fixation was estimated at 28.9 kgNina and 10.8 kg Nina in ferruginous andsandy soils, respectlvely.Pterocorpus is a pantropical genus belonging tothe tribe Do!bergieoe; it comprises approxi-mately 30 species of which about 15 occur inAfrica, 10 in America and 5 in Asia. Two sub-species ofPterocorpus Iucens are distinguished:subsp. Iucens (synonym: Pterocorpus abyssini-cus Hochst. ex ARIch. ) and subsp. ontunesit(synonym: Pterocorpus gritunesii (Taub. )Harms), the latter usuaUy having sinaner leaf-lets and a more hairy calyx.

Ecology Pterocorpus Iucens occurs in openand wooded savanna up to 1200(-1500) in alti-tude, preferring deep sandy soils, but also onclay soils including alluvial plains, where itusually becomes a tree, and on rocky outcropsincluding limestone, where it often remains a

shrub. It is usually found in drier habitats thanPterocorpus erinoceus POIT. , in regions with amean annual rainfall of 300-700 min. Locallyit is common; it often grows gregariously andmay form nearly pure stands. In southern of-rica it is nowhere common and typical of drydeciduous forest

Management Seeds can be stored for up to4 years, The weight of 1000 seeds is about 200g. Optimum temperatures for germination arein the range of 25-35'C and seedlings are lightdemanding. Trees can be managed by loppingto extend the period that they bear leaves. InNiono onall; average annual rainfall 500 mm)the foliage yield is 3.5 t dry matter per ha for anearly pure stand of 1600 stemsftia.

Genetic resources and breeding Locallythe pressure on populations of Pterocorpuslurens is high because of the popularity of itswood for house construction, increasing fuel-wood demand and frequent use for forage. InBurkina Faso it has been noted that in dry

years many Pteroco, pus Iucens trees may die;they have a comparatively poorly developedroot system that is probably often damaged bytermites, and are probably often weakened byregular harvesting offo11age for fodder.

Prospects Like some other Pterocurpus spe-cies, Pterocorpus Iucens is important as asource of forage during a large part of the dryseason. It is of high value for local people, alsobecause of its other uses as timber, vegetable,medicinal plant and fuelwood tree. Research onsustainable management practices Is needed tostop its decline and ensure its survival in thelonger term.

Major references Bolza & Keating, 1972;BurkiU, 1995; Le Hotterou, undated; Rojo,1972; WorldAgroforestry Centre, undated

Other references Arbonnier, 2004; CoatesPalgrave, 1983; Couteron, D'Aquino &Ouedraog0, 1992; de Freitas, 1986; Ganaba,1994; Ganaba, Ouadba & Bognounou, 2004;Neuwinger, 2000; Rivi6re, 1978; Sommerlatte& Sommerlatte, 1990; Syl}a at a1. , 1998

Authors R. H. M. J. Lemmens

PTnROCARPUS 483

PTEROCARPUSOSUN CTaib

Protologue Bull. Misc. Inform. Kew 1910: 329(1910)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution Ptero-corpus OSun is endemic to southern Nigeria,

484 TIMBERSl

Cameroon and Equatorial GuineaUses The bole is used to make dugout canoes

and the wood for carpentry, drums and walk-ing sticks. The reddish sap from the bark isused for dyeing, e. g. of traditional sculpturesThe stem is an ingredient of traditional medi-cines against sickle-cell disorder and amenor-rhoea. Powdered stem is applied topicalIy totreat skin diseases, to prevent infections of thefreshly severed umbilical cord, to treat stiffJoints, sprains and rheumatic complaints, andto promote healing offractured bones.

Production and international trade The

wood is traded in small quantities, occasionallyin mixed consignments with other Pterocorpusspp. as 'African padauk'

Properties The heartwood is dull red andfairly hard. It nails well and is liable to termiteattack. It contains red pigments of the santa-rubin and santalln groups. Santalins can beused as histological stain. The seeds contain28% crude protein, but in tests with rats theirnutritive value was inferior to casein. Beforethey can be used as food components or feedsupplements, safety studies and methods ofproper processing to remove antinutritionalfactors and possible toxic constituents are re-quired. The seed oil has been identified as auseful source of vitamin A activity based on its^-carotene content. Its main fatty acids arelinoleic acid, oleic acid, palmitic acid and be-heriic acid. In-vitro antimicrobial properties ofthe stem have been demonstrated. Stem ex-tracts showed antioxidant activity, and couldinduce a depigmenting effect and replace syn-thetic cosmetic formulations

Botany Evergreen or deciduous small tomedium-sized tree up to 30 in tall but usuallymuch smaller; hole often short and crooked, upto 80(-170) cm in diameter, sometimes withsmall buttresses at base; bark surface brown,rough, flaking off in irregular patches, innerbark yellowish, exuding a reddish gum onslashing; spreading; twigs denselybrown short-hairy when young, with soft prick-Ies. Leaves alternate, imparipinnately coin-pound with (9-)10-18 leaflets; stipules 7-15mm long, falling off early; petiole (3-)4.5-6(-8.5) cm long, rachis (9-)13-20(-26) cm long,densely brown hairy; petiolules 3-6 min long;leaflets alternate to almost opposite, oblong toovate or obovate, (4-)6-13 cm x 2-5 cm, baserounded or slightly cordate, apex shortly acu-minate, papery, densely hairy below whenyoung but later glabrescent, with 7-11 pairs oflateral veins. Inflorescence an axillary or ter-

minal panicle (4-)8-22 cm long, densely brownhairy. Flowers bisexual, papilionaceous; pedicelc. 2 min long; calyx campanulate, c. 7 min long,densely hairy, with 5 triangular teeth 1.5-3.5min long, upper 2 slightly longer than lower 3;corolla with clawed petals, yellow, standardobovate, up to 14 min x 12 mm, wings up to 12mm long, keel up to 10 mm long; stamens 10,fused Into a sheath up to 8 min long, the upperstamen sometimes partly free; ovary superior,stiped, hairy, style up to 2.5 mm long, glabrousFruit an orbicular, flattened, indehiscent pod7.5-15 cm in diameter, on a stipe up to 1.5 cmlong and with a papery, wavy wing, brownhairy and with prickles, I-seeded. Seed kidney-shaped, 18-24 min x c. 12 mm, dark brown toblackish.

In Nigeria trees flower in August-Novemberwhen they are in fullleaf. Bees commonly visitthe flowers and probably act as pollinators. Theroots have nodules containing nitrogen-fixingbacteria

Pterocorpus is a pantropical genus belonging tothe tribe Daibergieoe; it comprises approxi-mately 30 species of which about 15 occur inAfrica, 10 in America and 5 in Asia

Ecology Pterocorpus OSun occurs in lowlandevergreen or deciduous forest.

Genetic resources and breeding Ptero-corpus OSun has a limited area of distributionwhere it seems to occur scattered, and conse-quently may be easily liable to genetic erosion

Prospects Pterocorpus OSI, it has been stud-ied insufficiently, and it is difficult to deter-mine Its prospects as a commercialtlruber treeunder sustainable management. Its oftenpoorly shaped bole is a serious drawback. Itsmedicinal properties are promising for the de-velopment of skin care products.

Major references Burki11, 1995; Ebi &Ofoefule, 2000; Keay, 1989; Roj0, 1972.

Other references Avwioro at a1. , 2005;Ezeagu at a1. , 1998; Neuwinger, 2000; 01ukemiat a1. , 2005; Proll at a1. , 1998; Sprent, 2001

Authors R. H. M. J. Lemmens

crown

PTEROCARPUS ROTUNDIFOLIUS (Sond. ) Druce

Protologue Rep. Bot. SOC. Exch. Club BritIsles 1916: 642 (1917).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Chromosome number2n=24

Vernacular names Round-leavedbloodwood,round-leaved teak (En).

.

Origin and geographic distribution Ptero-corpus rotundifolius is widely distributed fromAngola, DR Congo and Tanzania south tonorthern Namibia, Botswana, Mozambique,northern SouthAfrica and Swaziland.

Uses The wood is locally used for furniture,shelves, picture frames, household utensils,wagon wheels and axe handles. It is also usedas firewood. In Zimbabwe sap from the tree isused as eye drops to treat eye complaints andin Tanzania a root decoction is drunk to treatanaemia. The foliage is browsed by cattle. Theflowers are a good source of nectar for honey-bees. The tree is occasionally planted as orna-mental and for erosion control.

Properties The hole is often poorly shapedThe heartwood is creamy white and has anoffensive, irritating smell when freshly cut. Itis moderately heavy to heavy, with a denslty of650-850 kg/ms at 12% moisture content. Thewood is rather difficult to saw, but it workswell. It is moderately durable and fairly resis-tant to insect attack. The seed contains 12%oil, the main fatty acids being linoleic acid,palmitic acid, oleic acid and stearic acid

Botany Deciduousshrub or sinaU to medium-sized tree up to 20 in tall, but usually muchsmaller, often many-stemmed; bole up to 60 cmin diameter; bark surface pale grey to brown,reticulately fissured or scaly, inner bark exud-ing a reddish gum on slashing; crown rounded,irregular; twigs short-hairy when young, gla-brescent. Leaves alternate, Imparipinnatelycompound with 3-19 leaflets; stipules linear,up to 15 mm long, faning off early; petiole 2-6(-8.5) cm long, rachis (2-)8-23(-30) cm long,densely hairy, glabrescent; petiolules (2-)5-15(-24) mm long; leaflets alternate to opposite,almost orbicular to ovate or elliptical, (3-)4.5-Toe15) cm x 2.5-6(-11) cm, base curieate torounded, apex shortly acuminate to rounded ornotched, papery to thinly leathery, appressedhairy below when young but later often gla-brescent, with 8-14 pairs of lateral veins. In-norescence a laxly branched, terminal panicle8-40 cm long, hairy. Flowers bisexual, papil-ionaceous; pedice1 3-Toe12) min long; calyxcampanulate, 6-9 min long, usually glabrous,with 5 triangular teeth 2-5 min long, upper 2teeth longer than lower 3; corolla with clawedpetals, yellow or orange-yellow, standardbroadly obovate to almost circular, up to 17min x 16 min, wings up to 16 mm long, keelupto 13 mm long; stamens 10, fused into a sheathup to 10 mm long, the upperstamen sometimespartly free; ovary superior, I-celled, stiped,

hairy, style up to 3 mm long, glabrous. Fruit anelliptical-oblong to almost orbicular, flattened,indehiscent pod 4-6.5 cm long, on a stipe up toI cm long and with a thinly leathery wing, gla-brous or slightly hairy, brown, I(-2)-seededSeed c. 6-9 mm x 4-5 mm, smooth, dark brownto blackish.

Seedlings grow fairly fast and may reach I intall after one year. In southern Africa Ptero-corpus rotundifo!jus trees are often completelyleaness from June to October. They usuallyflower in the rainy season, but during hot anddry weather the flower buds remain closed.Flowering is often very profuse. The stronglyfragrant flowers last 2-3 days and are coin-monly visited by bees, which probably act aspollinators. Fruits take about 3 months to ina-ture. They are dispersed by wind. The rootshave nodules containing nitrogen-fixing bacte-

Pteroco, pus is a pantropical genus belonging tothe tribe Daibergieoe; It comprises approxi-mately 30 species of which about 15 occur inAfrica, 10 in America and 5 in Asia. Three sub-species of Pterocorpus rotundifolius are distin-guished: subsp. rotundifo!ms. subsp. inortinii(Dunkley) Lock (synonym: Pterocorpus inor-tinti Dunkley) and subsp. polyonthus (Harms)Mendonga & SDUsa (synonym: Pterocorpuspolyonthus Harms). Intermediate specimensare rather common

Ecology Pteroco, pus rotundifolius occurs inwoodland and wooded grassland up to 900 inaltitude. In a study in South Africa, it pre-forred clayey, medium-Ieached, acid, non-sodicsoils with low conductivity, but in many areasit is common on deep sandy soils. In Mozam-bique Pterocorpus rotundifolius was found tobe fire-tolerant.

Management Seeds are often infested byinsects; infested seeds should be removed be-fore sowing. Soaking the seed in water for onenight improves the germination rate. It is rec-ommended that sowing be done in a mixture ofsand and sieved compost (2:1), and seedlingswith one leaf should be transplanted in plant-ing bags filled with a sand-based, well-drainedgrowth medium. Care should be taken to avoiddamage to the taproot. Seedlings need to beprotected from frost until about 2.5 in tall.Large stem cuttings (2-3 in long and about 10cm in diameter) planted in holes filled withsand have been used for propagation; theyshowed a high survival rate. Trees can be inari-aged by coppicing

Genetic resources and breeding There is

PTEROCARPUs 485

ria

.

486 TIMBERSl

no reason to consider this widespread and 10-cally common tree as threatened, although insome regions the trees are heavily browsed bycattle and elephants

Prospects Pterocorpus rotundifolius is of 11-mited Importance as a timber tree because ofits often poorly shaped and small-sized holeHowever, it is a multipurpose tree, importantfor forage and firewood, as well as for apicul-ture. It deserves more attention as an orna-

mental tree and has good prospects for soilconservation and improvement.

Major references Coates Palgrave, 1957;Mutshinya10, 2003b; Palmer & Pitman, 1972-1974; Roj0, 1972; World Agroforestry Centre,undated.

Other references Bredenkamp, 1986;Coates Palgrave, 1983; Gelfand at a1. , 1985;Gunstone at a1. , 1968; Lock, 1999; Neuwinger,2000; van Wyk & Gencke, 2000; van Wyk &van Wyk, 1997; Williamson, 1955; Zolh0, 2005.

Authors R. H. M. J. Lemmens

Properties The heartwood is creamy whiteto yellowish and indistinctly demarcated fromthe sapwood. The texture is moderately fineThe wood is moderately lightweight, moder-ately soft and easy to work. It is not durableand is liable to attacks by fungiand borers.Tannin is present in the bark and the woodStern extracts showed slight antimalarialactiv-ity against Pigsmodium 101ciporum strainsThe fat content of the seeds is low: less than

I%. In tests with West-African dwarf goats fedwith Pterocorpus sontolinoides foliage, weightgains were low, only about 8.5 g/day. The litterproduction is relatively low: in tests in south-ern Nigeria 7.3-8.1 tma

Botany Usually evergreen smalltree up to15 in tall; hole usually short, straight or moreor less twisted, up to 50 cm in diameter; barkthin, bark surface greyish brown, scaly andflaking in small patches, inner bark yellowishwhite to pinkish with red stripes, exuding alittle reddish gum on slashing; crown dense,with more or less drooping branches; twigsglabrous, with lenticels. Leaves alternate, jin-panpinnately compound with (5-)6-8(-9) leaf-lets; stipules often curved, up to 2(-3) cm long,slightly hairy; petiole (2-)3-4(-5.5) cm long,rachis (4.5-)311(-15.5) cm long, sparselyhairy, glabrescent; petiolules 2-5 min long;leaflets alternate to almost opposite, ovate toelliptical, (4-)6.5-12(-14.5) cm x (2-)3-5.5(-7)cm, base rounded to obtuse, apex shortly acu-inmate, papery to thinly leathery, sparselyhairy below when young but soon glabrescent,with 7-IOC15) pairs of lateral veins. Inflores-cence an axillary raceme (4-)6-14(-20)long, short-hairy. Flowers bisexual, papiliona-ceous; pedice1 2-4(-5) mm long; calyx cam-panulate, 8.5-9.5 min long, densely brownishhairy, with 5 triangular teeth 1.5-2.5 mm long,lower 3 teeth smaller than upper 2; corollawith clawed petals, bright yellow to orange-yellow, standard almost circular, up to 13 minx 15.5 min, wings up to 15 min long, keel up to12 mm long; stamens 10, fused into a sheathup to 10 mm long, the upperstamen sometimespartially free; ovary superior, whitish hairy,style up to 11.5 min long, glabrous towards thetop. Fruit an almost orbicular, flattened, inde-hiscent pod 2.5-4.5(-6) cm in diameter, wrin-kled or warty, brownish hairy, beige to palebrown, with a keel-like wing, I(-2)-seeded.Seed kidney-shaped, 1.5-2 cm x c. I cm, darkbrown

Pterocorpus sontolinoides grows fast. In trialsin southern Cameroon the fastest growing

PTEROCARPUSSANTALINOIDES DC.

Protologue Prodr. a 4/9 (1825)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Chromosome number it = 9, 2n = 22Origin and geographic distribution Ptero-

corpus sonto!trioides occurs from Senegal eastto the Central African Republic and DR Congo,and is also widespread in South America.

Uses The wood is locally used for temporaryconstruction, carpentry, sculpturing, fencesand boxboard. It is also used as firewood.

Roasted seeds are edible, tasting slightly likegroundnut, but raw seeds are toxic. Cookedyoung leaves are eaten as a vegetable; they arealso added to soup. The foliage is an importantforage for many kinds of livestock. The bark islocally used for dyeing textiles brownish, whilethe bark. exudate is occasionally used to givethem a reddish colour. Pterocorpus sonto!i-notdes Is occasionally planted as a shade treefor crops and to improve the soil by fixing ni-trogen and litter production. It is also plantedas a windbreak. The bark, roots and leaves arecommonly used in medicinal preparations. De-coctions are administered externalIy to woundsto promote healing, and to treat haemorrhoidsand fever. They are taken internally to treatbronchial complaints, amoebic dysentery,stomach-ache and sleeping sickness, to preventabortion and ease childbirth, and as a tonic

cm

.

trees reached a height of 6.4 in and a diameterof 10.3 cm 20 months after planting. Treesflower towards the end of the dry season. Beescommonly visitthe flowers and probably act aspontnators. Fruits mature 2-3 months afterflowering in the rainy season. They float inwater. The roots have nodules containing ni-trogen-fixing bacteria, but in tests In south-eastern Nigeria nodules did not develop inseedlings.Pterocorpus is a pantroplcal genus belonging tothe tribe Do16ergieoe; it comprises approxi-mately 30 species of which about 15 occur inAfrica, 10 in America and 5 in Asia.

Ecology Pterocu, pus sontolinoides is typusallyfound on river banks, usually on sandy andmoist soils, up to 500 in altitude. It occurs inregions with an annual rainfall of about 1600mm, and may survive dry periods of over 5months provided that the roots can reach theground water. It tolerates shade and seasonalwaterlogging. It is locally common, and some-times even occurs gregariously. In central C6ted'Ivoire about 60 adult trees per kin have beenrecorded along the Bandama river. In southernCameroon planted trees performed well onhighly acidic and aluminium toxic soils

Management Pterocorpus suntolinoides canbe propagated by seeds, stem cuttings and rootcuttings. Dry seeds can be stored for longerperiods. Scarification or soaking in water Isrecommended prior to sowlrig. Germinationstarts after 7-14 days, and the germinationrate is about 70%. Trees can be managed bypollarding, coppicing and lopping.Tests in Nigeria and Cameroon showed thatPterocorpus sontolinoides has signlficant po-tential for alley cropping. Hedgerows were es-tablished using 2-month-old seedlings plantedat 4 in x 0.25 in. The hedgerows were prunedafter one year at 50 cm height, in subsequentyears twice a year, and lriterplanted with

especially for rice cultivation.Prospects Like several other Pterocorpus

species, Pterocorpus suntoltnoides is a multi-purpose tree that deserves more attention. Itwill not gain Importance as a timber tree be-cause it is too small and provides wood of inod-erate quality, but it has prospects for agrofor-estry programmes as it improves soil fertilityand outcompetes weeds, as well as providingforage and fuelwood. It is an important speciesfor soilconservation in water catchment areasMoreover, Pterocorpus suntolinoides is suitableas an ornamental tree, having showy flowersand shiny leaves

Major references Burki11, 1995; CAB In-ternationa1, 2005; Gautier-Beguin, 1992; Rojo,1972; World Agroforestry Centre, undated.

Other references Anegbeh & Tchoundjeu,2002; Arbonnier, 2004; Angbede, Barnikole &Babayemi, 2003; Arodokoun at a1. , 2003; Egbeat a1. , 1998; Ezeagu at a1. , 1998; Kanmegne ata1. , 2000; Keay, 1989; Neuwinger, 2000; Salako& Tian, 2005.

Authors R. H. M. J. Lemmens

PTEROCARPUS 487

maize

Fruits are often collected from the water sur-

face. They are heated over a fire untilthe seedsappear, which are eaten roasted.Pterocorpus sontolinoides is a host plant forthe bean pod borer Moruco uitroto, which is apest of e. g. cowpea; it is a key relay host duringthe long dry season

Genetic resources and breeding Thereare no indications yet that Pterocorpus sonto-Jinoides is threatened by genetic eroslon. How-ever, its characteristic habitat, i. e. riparianforests, is under pressure in many regions be-cause of transformation Into agricultural land,

PTEROCARPUSSOYAUxiiTaub.

Protologue 011v. , Hooker's Icon. PI. 24: t.2369 (1895)

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Vernacular names African padauk, Africanpadouk, barwood, African coral wood (En). Pa-douk d'Mrique, padauk d'Mrique, bois coralI(Fr). Ndimbu, nkula (Po).

Origin and geographic distribution Ptero-corpus soyouxii occurs from south-eastern Ni-

.

@?,^,.=*.,93"

Pterocorpus soyouxii- wild

.,

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488 TIMBERSl

gena east to eastern DR Congo and south tonorthern ingola.

Uses The wood of Pteroco, pus soyouxiiis avaluable multipurpose hardwood. Because ofIts resistance to water it is locally used to makecanoes and because of its beautiful reddish

colour it is favoured for carving and sculptur-ing, furniture, cabinets, knife and tool handles,traditional hair combs, walking sticks and inu-siCalinstruments. The wood has a high reso-nance quality as its damping of vibrations islow and formerly large telegraph slit drumsand war drums, as well as xylophones, weremade from it in DR Congo and Gabon. It iscurrently used also for 'Western music' xylo-phones and increasingly tried for the back andsides of guitars. Because of its high durabilitythe wood is excellent for construction, carpen-try, outdoor joinery, flooring, staircases, rail-way sleepers and boats but also for veneer,Inlay, billiard tables, toys, joinery, dowels,shuttles, bobbins, spindles, sporting goods andpaddles. As the wood is resistant to marineborers it was used in temperate regions formarine constructions such as piers and sluicegates. In the Herault region in France it wasused for years in the construction of water-wheels for irrigation. The wood is also used asfuel.

The heartwood is the source of the so-called

true barwood dye. In Africa nowadays, the dyeis still used to colour red fabrics, fibres andclothes, including the tail-like ornaments madefrom raffia fibre in Cameroon and worn on theback by women of the Bulu people. In DRCongo, in the former kingdom of Kuba, at theconfluence of the rivers Kasai and Sankuru,the dyes of the famous 'Kasai velvets' includePterocorpus soyouxii reds with a more violetshade, obtained by combining the red dye withtannin-rich plants and a mordant of iron-richmud.

A pomade is made by mixing the red woodpowder with oil and its use as a body cosmeticis widely applied in DR Congo ('rigula'). Theroots can be prepared and used in the sameway as the heartwood and yield a dye of equalor better quality. Pulverized bark, mixed withpalm oil, Is also used as a cosmetic pomadeThe leaves are eaten as a cooked vegetableThe powdered wood, baked with a slice of lime,is used in Gabon on wounds and, mixed withpalm oil, raffia oilor vegetable butter (e. g. fromseeds of negheme!IQ of neono Pierre), to treatskin diseases, ringworm and yaws. Probablypartly due to its blood-red colour and the asso-

ciated symbolism, it is also used in ritual cere-monies for circumcision, initiation, marriage,delivery and widowing. The bark contains akino type resin ('dragon's blood') which is veryastringent and used to ward off skin parasitesin ethnoveterinary medicine. In Gabon theresin is used (usually in combinations withparts of other plant species) as an enema totreat dysentery and against toothache, gonor-rhoea and excessive menstruation. In Congoand the Central African Republic a bark decoc-tion is drunk to treat dysmenorrhoea, uterinehaemorrhage, dysentery and haemorrhoids. Apulp obtained by scraping the inner surface ofthe bark is applied as a wet dressing againstinflammations, oedemas, incipient hernia andwhitlow. Decoctions, draughts or vapour-bathsof leaves and bark are taken against broncho-pulmonary affections.

Production and international trade In Ga-

bon PIerocorpus soyuz4xiiis among the 10 mostimportant export timbers. In Gabon, where thestanding stock was estimated at 15 millions inain 1999, the minimum diameter limit for ex-PIOitation is 70 cm. Between 2000 and 2003,Gabon exported 120,000 ing of African padauklogs each year, whereas in 1997 the export wasonly 57,000 ms. Cameroon, where the export oflogs is prohibited, exported 6,500 ina of sawntimber in 2003. The domestic use of the timber

is high. In North America African padauk isavailable as lumber and veneer, and prices arehigh. In Europe sawn timber and veneer areavailable on a limited scale only.Formerly the heartwood of Pterocorpus soy-duni was exported in great quantities in stan-dard blocks as red dyewood from Gamero0nand Gabon to Europe and North America andused in the wool-cloth and printed cotton in-dustries. Because of its colour fastness, it con-tinued to be used to dye wool-cloth until thebeginning of the 20th century in the UnitedStates, and in England and France marticu-Iarly Elbeuf) to produce brick reds and red-browns or to give a ground for greys and blacksdyed with logwood. As a dye for cotton-cloth,Pterocorpus soyauxii with a tin mordant gavethe traditional red colour of the printed ban-danas that were so popular in England in the19th century. The resinous consistency of thedye gave the handkerchiefs added weight,characteristic of these items.

Properties The heartwood is bright red whenfreshly cut, becoming orange-red on exposureand darkening to purple-brown, and distinctlydemarcated from the whitish to brownish yel-

.

low sapwood 6-10(-20) cm wide. The grain isstraight to interlocked, texture coarse. Thewood has a faint aromatic scent when freshlycut.

Pterocorpus soyouxii wood is moderatelyheavy, the density is (650-)675-815(-900)kg/ing at 12% moisture content. The shrinkagerates are moderately low, from green to ovendry 2.2-3.8% radial and 4.1-6.2% tangentialThe wood dries well but moderately slowly,with little risk of distortion. Once dry, it is verystable.

At 12% moisture content, the modulus of rup-ture is 101-218 N/mm2, modulus of elasticity10,800-15,900 N/mm2, compression parallel tograin 54-79 N/mm2, shear 7-8 N/min2, cleav-

11-18 N/min and Janka side hardnessage

6850-8320 N.

The wood is moderately difficult to work. Stel-lite-tipped sawteeth are recommended. Thewood takes a good finish, but sometimes withsome tearing of interlocked grain. Slicing doesnot cause problems, and the wood holds nailsand screws satisfactorily; however, pre-boringfor screwing is advisable. The gluing propertiesare good. The dry sawdust may cause irritationto skin, nose and bronchia. The heartwood isdurable and also resistant to fungi, Lyctus bee-tles, termites and marine borers; it is moder-ately difficult to impregnate with preserva-tives. The sapwood is less durable and moder-ately difficult to impregnate. The heartwood isrich in extractive substances by organic sol-vents. The ash, 11gnin and cellulose contentsare moderate. Pentosane content is very low,similar to coniferous wood. Silica content is

also very low. The vibration damping factor ofthe wood is 0004-0007 at frequencies of 200-500 Hz. The wood has little moisture uptake; ingiven conditions, its moisture content is nearlyhalfthat of'standard' woods

The heartwood of Pterocorpus soyouxii con-tains the red binavonoids santalin A, santa-rubin A and santarubin B, ISOflavonoids includ-Ing pterocarpin, formononetin and prunetin,the ISOflavanequinone claussequinone and theisonavanes vestitol and inucronulatol. The low

damping factor, shrinkage coefficient and ino1s-ture uptake of padauk heartwood appear to belinked to its specific composition in extractivesubstances, and sapwood has much higher val-ues of these physical propertiesThe wood is also rich in tannins, which con-tribute to mordanting in the dyeing process. Inthe Colour Index barwood is cited as a source of

natural red N0 22. Santalin is a histological

dye, coinparable in use to haematoxylin. Incombination with an acid Fe or A1 mordant, Itselectively dyes cell nuclei, elastic tissues andstriations in voluntary muscle fibres.The leaves have a high ascorbic acid contenteven after cooking. The bark of Pterocorpi!ssoyouxii showed antifungal activity againstsome pathogenicfungi

Adulterations and substitutes The wood

of Pterocorpus OSun Craib from southern Nige-ria and Garnero0n, Pterocorpus tessmonniiHarms from Equatorial Guinea, Gabon and DRCongo, and Pterocorpus tinctorius WeIw. fromDR Congo and ingola are also marketed asAfrican padauk. The dye of Pterocorpus soy-guni may be substituted for by the dyes ofother insoluble redwoods, the best known ofwhich are sandalwood (Pterocorpus sontolinusL. from southern India), narrawood (Pterocor-pus Indicus Wind. from the Philippines andMyanmar) and camwood (Bophio nitido Lodd.from West Africa)

Description Evergreen, sometimes deciduouslarge tree up to 55 in tall; hole straight andcyhndrical, branchless for up to 20(-30) in, upto 140(-200) cm in diamete", slightly to promi-nently and highly buttressed; bark grey-brownto brown, peeling off in thin irregular flakes,exuding a red gum abundantly on slashing;crown dome-shaped, open; twigs brown hairywhen young. Leaves alternate, Imparipinnatelycompound with 7-17 leaflets; stipules linear,up to 2 cm long, hairy, falling off early; petioleI-3.5 cm long, rachis 3.5-16.5 cm long, denselybrown hairy; petiolules 3-5 min long, shallowlyfurrowed; leaflets alternate to nearly opposite,obovate to elliptical, 2.5-9 cm x 1.5-4 cm, baserounded to obtuse, apex usually abruptly acu-inmate and inucronate, leathery, glabrous,with closely set, fine and obscure lateral veinsInflorescence an axillary or terminal much-

long, denselybranched panicle 10-35brown hairy; bracts linear, falling off at arithe-SIS. Flowers bisexual, papilionaceous; pedicel3-19 mm long, hairy; calyx campanulate, c. 7.5min long, densely hairy, with 5 triangularteeth I-2.5 min long, upper 2 more or less con-nate; corolla with clawed petals, bright yenowor orange-yellow, standard almost circular upto 13 min x 10 mm, wings up to 12 mm long,keel up to 9.5 min long; stamens 10, connateinto a sheath up to 8.5 min long, the upperstamen sometimes free; ovary superior, I-celled, stalked, white hairy, style up to 4 minlong, glabrous, stigma terminal. Fruit a circu-Tar, flattened, indehiscent pod 4.5-9 cm in dia-

PTEROCARPUS 489

cm

.

490 TIMBERSl

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pits alternate; 23: shape of alternate pits po-Iygona1; (25: intervessel pits small (4-7 prn));26: intervessel pits medium (7-10 F1m); 29:vestured pits; 30: vessel-ray pits with distinctborders; similar to intervessel pits in size andshape throughout the ray cell; 43: mean tan-gentialdiameter of vessellumina z 200 prn; 46:s 5 vessels per square minimetre; 58: gums andother deposits in heartwood vessels. Tracheidsand fibres: 61: fibres with simple to minutelybordered pits; 66: non-septate fibres present;69: fibres thin- to thick-walled; 70: fibres verythick-walled. Axial parenchyma: 77: axial pa-renchyma diffuse-in-aggregates; 80: axial pa-renchyma allform; 82: axial parenchymawinged-allform; 83: axial parenchyma conflu-ent; (85: axial parenchyma bands more thanthree cells wide); 86: axial parenchyma in nar-row bands or lines up to three cells wide; (89axial parenchyma in marginal or in seeminglymarginal bands); 91: two cells per parenchymastrand; 92: four (3-4) cells per parenchymastrand. Rays: (96: rays exclusively uniseriate);(97: ray width I-3 cells); 104: all ray cells pro-cumbent; 1/6: Z 12 rays per mm. Stoned struc-ture: 1/8: all rays storied; 120: axial paren-chyma and/or vessel elements storled; 121:fibres stoned. Mineral inclusions: 136: pris-matic crystals present; 142: prismatic crystalsin chambered axial parenchyma cellsre. Baas)

Growth and development In the Lope re-serve in Gabon the trees flower from Decem-

her-February and the fruits, produced in greatquantity, ripen from January-April. Some-times the tree is deciduous just before flower-ing. The winged fruits are dispersed by wind,but also by animals. Germination and seedlinggrowth are rather fast. In plantations the an-nualincrement of the wood in Nigeria has beenestimated at 40 msftia. In trial plantations of1.5 ha in C6te d'TVoire in 1964 and 1968, strip-lings were transplanted at spacings of 4 in x 4in and 5 in x 5 in. The first 7 years, annualheight growth varied between 1.6 in and 2.7 in.The mean annual volume growth, includingthinnings, was 20-30 mama over a period of 15years. The average annual diameter growth ofthe 150 largest trees per ha was 2.5 cm when17 years old. Pterocorpus soyouxi, fixes nitro-gen in its root nodules.

Ecology Herocoipus soyouxL! occurs scatteredor in small groups in evergreen and deciduousforest, from sea-level up to 500 in altitude. Itprefers a moist but well-drained deep soil, anaverage annual rainfall of 1500-1700 min and

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Pterocorpus soyouxii- I, buse of 601e, 2, leg/; 3,inflorescence, 4, fruit.Redrawn und oddpted by Ishoh Syomsudtn

meter, on a stalk up to I cm long and with apapery, finely veined wing with wavy or plaitedmargin, finely hairy, glossy brown, I-seededSeed kidney-shaped, flat to slightly thickened,12-16 min x 5-7 mm, smooth, red when fresh,turning dullbrown to black.

Other botanical information Pterocorpusis a pantropical genus belonging to the tribeDo16ergieoe, comprising 21 species of which 12occur in Africa, 6 in America and 5 in Asia.Several Asiatic and African species were jin-portant commercial sources of red dyes butmost species are now much more valuable fortimber. The bark of most species yields a red-dish medicinal resin of'kirio' type.Pterocorpus tessmonnti Harms is often con-fused with Pterocorpus soyauxii. It has similarwood and dye uses and properties, but grows inwetter parts of the forest. It can be distin-guished by its winged, sickle-shaped to oblongfruits 11.5-13.5 cm x 4-5 cm.

Anatomy Wood-anatomical description (IAWAhardwood codes):Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervessel

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an average annual temperature of 23'C.Propagation and planting Propagation by

seed is easy. The seeds are dried in the sun andthe wings are removed. In Congo germinationstarted 3 days after sowing and 92% of theseeds germinated within 30 days. In a test inNigeria, 86% of the seeds with fruit wall re-moved and soaked overnight in water germi-nated in 7 days. The germination is hypogeal.Planting into the field is done about 40 daysafter sowing and is easy. Propagation by non-woody cuttings in normal topsoil gave 83%

Seedling growth showed a greaterresponse after inoculation of the soilwith fungifrom the rhizosphere of the mother tree thanafter inoculation wlth a similar spore numberoffungifrom a fallow field.

Management Pterocorpus soyouxiirequiresmuch light for good growth. Stump regrowth isweak and uneconomicfor wood production

Harvesting Logs are liable to brittle heart.For dye extraction, preferably old and hollowtrees are cut from the forest and the heartwood

is lumbered out. Often trees are felled and left

for 2-3 years lying on the forest floor beforetaking the heartwood for dyeing purposes. Theroots are also harvested for dye extraction

Handling after harvest Freshly felled logsof Pterocorpus soyauxii usually do not float inwater, and consequently cannot be transportedby river. For dye production the heartwood issplit into billets and chips which are dried andsubsequently pounded into powder. A little oilis added to the pulverized material, which ismoulded into cakes for stocking and for localsale. For dye export the heartwood is traded instandard blocks or bars ('barwood'), makingquality controleasierSince the colorants present in the wood aredifficult to dissolve in water, special methods ofextraction were developed in the dyeing indus-try in 19th century Europe, boiling the wood for1.5-2 hours in a solution of 45' alcohol or in

water and carbonate of soda (30 g per 100 g offibre to be dyed). This solution was then di-luted with water and used asthe dye bath.

Genetic resources Pterocorpus soyouxiiisrather widespread in Central Africa and cur-rent exploitation rates do not seem to endangerthe species. To safeguard genetic vanability,protection measures of some natural forestwhere Pterocorpus soyouxii occurs are recoin-mended

Prospects Pterocorpus soyouxiiis a usefulmultipurpose tree from forest areas in CentralAfrica, producing good-quality timber, dye,

success

vegetables and medicine. Over-exploitationendangers natural populations. More researchis needed on growth requirements. Solutions tomake better use of and more profit from thechips, sawdust and bark as by-products of thetimber, e. g. for dye-extracts and medicinal ap-PIications, should be encouraged. As it is quitea fast growing, nitrogen fixing and light de-manding tree, it might be useful for agrofor-estry purposes (e. g. as shadow tree in coffeeplantations). The feasibility of establishingcommercial plantations also deserves research

Major references Burki11, 1995; Cardon,2003; CTFT, 1978d; Raponda-Walker & Sil-Ians, 1961; Roj0, 1972; Surowiec, Nowik & Tra-ianowicz, 2004; World Agroforestry Centre,undated

Other references Amone at a1. , 1977; Ban-nerjee & Mukherjee, 1981; Br6maud at al. ,2004; Carrington, 1976; Duke, 1981; Evrard,1988; Hauman at a1. , 1954a; Kiec-Swierczynska at a1. , 2004; Kouablan & Bellgne,1981; Laine at a1. , 1985; Nzokou & Kamdem,2003; Okafor, Okolo & Ejiofor, 1996; OSho &Ajonina, 1991; OSIisly, 1999; Pangou, 1982;Richter & Danwitz, 2000; Rojo & Noriz0, 1993;Takahashi, 1978; White &Ahemethy, 1997

Sources of illustration Wilks & Issemb6,2000; Engler, 1910.

Authors PCM. Jansen

PTEROCARPUS 491

PTEROCARPUSTINCTORIUS WeIw.

Protologue Apont. : 584 (1859)Family Papilionaceae (Leguminosae - Papil-

ionoideae, Fabaceae)Synonyms Pterocorpus chrysothrix Taub

(1895), Pterocorpus stolzii Harms (1915)Vernacular names Tacula coo). Mintiga mail

(Sw)Origin and geographic distribution Ptero-

corpus timetorius is widespread in Central,East and southern Africa, from Congo and DRCongo east to Tanzania and south to Angola,Zambia, Malawiand Mozambique

Uses The wood is popular for furniture, ca-binet making and decorative parquet floors. Itis also suitable for light construction, joinery,interior trim, boxes, crates, tool handles, carv-ing, turnery, veneer, plywood, hardboard, par-ticle board, and pulpwood for lower-qualitypaper production. It is used as firewood and formaking charcoal. The foliage is browsed bygoats. The reddish dye from the wood has beenused for colouring the body. In DR Congo a

492 TIMBERSl

bark decoction is applied as a rectal washing totreat lung congestion in children. In Tanzaniathe tree is used for shade

Production and international trade Thewood is traded on the international market in

small quantities. It is occasionally traded inmixed consignments with other Pteroco, pusspp. as 'African padauk'. It is traded from theEast Usambara Mountains in Tanzania; in1990 the local price of a plank of 3.7 in x 0.3 inwas approximately Us$ 2.40, in 2000 Us$ 4.The timber is locally in great demand, e. g. inBurundi and DR Congo from where it is ex-ported

Properties In general, the properties ofPte-rocorpus tinctorius are coinparable to those ofPterocorpus origolensis DC. The heartwood ispale yenow when freshly cut but turning topinkish red upon exposure, and distinctly de-marcated from the whitish, 7.5-10 cm widesapwood. The grain is often interlocked, tex-ture moderately fine. Irregular, sinaU, dark redor brown markings are present on tangentialsurfaces; the wood contains red gummy sub-stances. The density of the wood at 12% ino1s-ture content ranges from about 450 kg/ing (for-est trees, Mayombe, DR Congo) to about 900kg/ing (savanna trees, Burundi). The wood usu-ally dries well with little deformation. Shrink-age rates from green to oven dry are 3.3% ra-dial and 5.5% tangential for wood from Bu-rundi. Once dry, the wood is stable in service.At 12% moisture content, the modulus of rup-ture of wood from Mayombe is 91 N/min',modulus of elasticity 9100 N/mm', compressionparallel to grain 45 N/min', cleavage 8 N/min"and Chalais-Meudon hardness 2.2. At the same

moisture content, the modulus of rupture ofwood from Burundiis 147 N/mm", modulus ofelasticity 15,000 N/mm' and compression par-allelto grain 77 N/mm'. The wood saws andworks well, and can be planed to a smooth sur-face. It holds nails and screws well and is gen-erally not liable to splitting, although pre-boring is recommended for timber from Bu-rundi. When a filler is used it finishes well

The wood is moderately durable to durable; thelighter wood is liable to termite attack andslightly liable to Lyetus attack, but heavierwood is not. It Is moderately resistant to jin-pregnation with preservatives. The sawdustmay cause Irritation to workers

Botany Evergreen or deciduous smallto me-dium-sized tree up to 25 in tall; hole branchlessfor up to 15 in, often straight and cylindrical,up to 75 cm in diameter; bark surface grey to

dark brown, fissured and scaly, Inner barkwhitish exuding a reddish gum on slashing;crown round or flattened, dense; twigs short-hairy when young. Leaves alternate, imparip-innateIy compound with (5-)7-Ile15) leaflets;stipules oblong, c. 3 mm long, falling off early;petiole (I-)2-5(-10) cm long, rachis (2.5-)4-20(-30) cm long, densely hairy, glabrescent;petiolules (2-)3-8(-12) mm long; leaflets alter-nate to almost opposite, oblong to ovate orobovate, (2.5-)4.5-Ile13) cm x (1.5-)2-5(-7)cm, base obtuse to rounded or slightly cordate,apex shortly acuminate, papery to thinly Ieath-ery, hairy below when young but later glabres-cent, with 6-14 pairs of lateral veins. Inflores-cence an axillary or terminal panicle (4-)8-22cm long, densely hairy. Flowers bisexual, papil-ionaceous; pedice1 3-7 mm long; calyx cam-panulate, 5-9 mm long, densely hairy, with 5triangular teeth 1.5-3 min long, upper 2slightly longer than lower 3; corolla withclawed petals, cream-coloured to orange-yellow,standard obovate, up to 18 min x 13 min, wingsup to 16 min long, keel up to 13 mm long; sta-mens 10, fused into a sheath up to 10 min long,the upper stamen sometimes partly free; ovarysuperior, I-celled, stiped, hairy, style up to 5.5mm long, glabrous towards the top. Fruit anorbicular, flattened, indehiscent pod 5-21 cmin diameter, on a stipe up to 2 cm long, with athin-leathery wavy wing, hairy, greyish brownor reddish brown, I-seeded. Seed kidney-shaped to oblong, flat to slightly thickened, 15-25 min x 8-13 mm, wrinkled, dark brown toblackish.

In DR Congo trees flower in March-May. Beescommonly visitthe flowers and probably act aspontnators. The leaves are commonly eaten bycolobus monkeys and chimpanzees.Pterocorpus is a pantropical genus belonging tothe tribe Do16ergieoe; it comprises approxi-mately 30 species of which about 15 occur inAfrica, 10 in America and 5 in Asia. Pterocor-pus tinctorius is variable and has been subdi-vided into several infraspecific taxa; however,these are connected by many intermediateswhen the whole region of distribution is con-sidered.

Pterocorpus indieus Wind. and Pterocorpusdo16ergioides Roxb. ex DC. , both from tropicalAsia, are planted in Africa as ornamental androadside trees, the former e. g. in Sierra Leone,Nigeria, Kenya, Tanzania and Mauritius, thelatter in Madagascar. Both species are impor-tant timber trees in tropical Asia, but are notplanted for timber production in Africa. In

.

Mauritius the sap of Pterocorpus indieus isused to relieve toothache and as an antidote for

poisoning.

Ecology Pteroco, pus tinctorius occurs in awide variety of habitats, from evergreen rain-forest to riverme forest and wooded savanna,up to 1800 in altitude, often on rocky hills.

Management Pterocorpus tinctorius can bepropagated by seed and by cuttings. Sometimeswildlings are collected for planting. Trees canbe managed by CODpicing, pollarding and lop-ping. In Zambia they are reportedIy fire resis-tant.

Genetic resources and breeding Ptero-corpus tinctorius is widespread and locallycommon, and consequently does not seem to bethreatened by genetic erosion.

Prospects Pterocorpus timetorius is a usefultimber tree, providing a fair quality wood thatis an interesting substitute for the wood ofPterocorpus origolensis, the latter being subjectto unsustainable harvesting in many countriesHowever, Pterocorpus tinctorius has been stud-led insufficiently, and it is difficult to deter-mine its prospects as a commercial timber treeunder sustainable management. Its other usesshould also be considered when planning opti-maimanagement.

Major references Bolza & Keating, 1972;Chilufya & Tengn6. s, 1996; Fouarge & G6rard,1964; Ishengoma, Gillah & Chihong0, 1997;Parant, Chichignoud & Rakotova0, 1985.

Other references Burki11, 1995; Disengo-inoka, Delaveau & Sengele, 1983; du Puy at al. ,2002; Fouarge, Sacr6 & Mottet, 1950; Grinb-Fakim, Gu6ho & Bissoondoya1, 1997; Haumanat a1. , 1954a; Lovett at a1. , 2006; Roe at al. ,2002; Roj0, 1972; Rojo & A10nz0, 1993

Authors R. H. M. J. Lemmens

.

PTERYGOTA 493

,

Pterygotu ingcrocorpo - wild

elling, interior joinery, moulding, furniture,block board, fibre board, particle board andlight carpentry. It is also used for staircases,light flooring, wood-frame houses, glued-laminated timber, matchboxes, boxes andcrates. The wood is also used for fuel

In C6te d'Ivoire Pterygoto ingcrocorpo Is occa-sionally retained as a shade tree in cacao plan-tations. The large leaves are used as temporarythatch and for wrapping food. In Nigeria a de-coction of the leaves is drunk against stomach,bladder and urinary problems and againstflatulence.

Production and international trade The

timber ofPterygoto inderocorpo is important in06te d'TVoire, Ghana and Gamero0n. In 1995C6te d'TVoire exported 2000 in" of rotary veneerand 2000 m' of sliced veneer at a price of Us$4061m" and Us$ 9631m', respectively, in addi-tion to 5000 in" of logs at an average price ofUs$ 5671m'. In 2004 C6te d'TVoire exported32,000 in3 of sawn wood at Us$ 397 Im', and in2005 it exported 25,000 ina at Us$ 4391m"Ghana exported rotary veneer, sliced veneerand jointed veneer in 1995, at average prices ofUs$ 5101m", Us$ 9001m' and Us$ 1250/in',respectively. It also exported 9000 in" of sawnwood at an average price of Us$ 4401m' and1000 in" of logs at Us$ 1651m'. In 2001 Ptery-goto ingcrocorpo timber represented 5% of thetimber exports of Ghana, but was among the 3most important sources of plywood for export;the export price offace veneer was about Us$3001m". In 2004 Ghana exported 5000 in' ofsawn wood at an average price of Us$ 5601m3,5000 m' of veneer at an average price of Us$6931m", and some plywood at Us$ 3271m". In

PTERYGOTAMACROCARPAK. Schum.

Protologue Engl. , Monogr. afrik. Pflanzen-Fam. 5: 135 (1900)

Family Sterculiaceae (APG: Malvaceae)Chromosome number2n=36

Vernacular names African pterygota (En)Koto, erable d'Mrique (Fr)

Origin and geographic distribution Ptery-goto inocrocorpo occurs from Sierra Leone eastto Congo, DR Congo and Cabinda (Angola). Itmay also occur in the drier parts of northernGabon.

Uses The wood (trade names: koto, ptery-gota) is used for veneer, plywood, interior pan-

L.

o

,^

..

.

494 TIMBERSl

2005 it exported 4000 in' of sawn wood at anaverage price of Us$ 4871m", and 6000 in" ofveneer at an average price of Us$ 6271m', andsome plywood at Us$ 3751m". According toATIBT statistics, Cameroon exported 1325 inaof logs in 2003, 535 ingin 2004, and 2250 main2006. It exported 2460 ing of sawn wood in2003, 980 in3in 2004 and 3185 main 2006

Properties The heartwood is creamy whiteto yellowish, sometimes attractively flecked onquartersawn surfaces. It is not demarcatedfrom the up to 10 cm wide sapwood. Freshwood has an unpleasant odour. The grain isstraight or occasionally shallowly interlocked,texture moderately coarse.The wood is medium-weight, with a density at12% moisture of 480-660(-750) kg/in'. Thewood dries fairly easily with little risk of sur-face checking or twisting, but with some risk ofextension of shakes. The rates of shrinkagefrom green to oven dry are high: 3.9-6.0% ra-dial and 8.8-12.6% tangential. Once dry, thestability in service is mediumAt 12% moisture content the modulus of rup-ture is 85-153 N/mm', modulus of elasticity9200-14,800 N/mm', compression parallel tograin 43-58 N/mm", shear 6-9 N/min' cleav-

17-19 N/mm and Janka side hardnessage4190 N

The wood can be worked with normal hand and

machine tools. The blunting effect on cuttingedges Is medium. Sawing is satisfactory, butthe tendency to breaking out at the bottom ofthe cut may be pronounced. For planing quar-tersawn surfaces, a reduction of the cuttingangle to 20' is recommended and cuttersshould be kept sharp. For good surface finish-ing, a filler is needed. The naihng and screwingproperties, as well as the peeling and slicingproperties are satisfactory to good. Bendingproperties are variable, but in general the woodbuckles

The wood is not durable, being susceptible toattack by borers, termites and fungi. It is veryprone to blue stain. However, It is easily jin-pregnated with preservatives. The energyvalue of the wood is 18,170 kJ/kg

Adulterations and substitutes The wood

of Pterygoto mererocorpu is very similar to thatof Stercu!to oblongo Mast. ('eyong'), which isalso medium-weight and used for similar PUT-poses

Description Dedduous, medium-sized to fair-Iy large tree up to 35 in tall; hole cylindrical, upto 120 cm in diameter, with very large, thinand wavy buttresses, bark surface greyish,

vY4<-'

.

..

II

I\

4

Pterygoto inderocurpo - I, led/: 2, mole flower, '3, ferno!e flower, . 4, fruit, 5, seed.Redrown und udopted by Ishoh Syomsudin

I

smooth with raised hoops and smalllenticels,inner bark with green outer layer, white withvertical lines; crown sinaU, dense, branchletshairy, glabrescent. Leaves alternate, simple;stipules lanceolate, c. 5 min long, greyish hairy;petiole 5-18 cm long; blade ovate to orbicular,sometimes slightly lobed, 5-30 cm x 10-25 cm,base cordate, apex short-acuminate, brownishhairy when young but glabrescent, shiny, with5-7(-9) veins from the base and 2-4 pairs oflateral veins. Inflorescence a few-flowered,terminal panicle 6-14 cm long, reddish brownhairy. Flowers usually unisexual, regular;pedice1 5-6 min long, jointed below the middle;calyx campanulate, 5-6 lobed, lobes lanceolate,13-20 mm x 3-6 mm, densely reddish stellatehairy outside; corolla absent; male flowers withstaminalcolumn c. 8 min long, thin, with 8-10arithers at apex and surrounding 5 rudimen-tary carpels; female flowers with superiorovary consisting of 5 carpels fused at base,each carpel with a c. 2.5 mm long style andsmall stigma, surrounded by rudimentary an-thers. Fruit consisting of I-5 ellipsoid to globu-Iar follicles 12-18 cm x 10-13 cm, woody, with1.5-4.5 cm long stipe, many-seeded. Seeds 9-11 cm long including a wing of 6.5-7 cm x c. 3cm. Seedling with epigealgermination; epicotyl4-6 cm long, hypocoty1 2-3 cm long; cotyledonsoblong, c. 3 cm x 2 cm; first leaves opposite,

I,

.

simple, blade 8-11 cm x c. 7 cm, cordate atbase, long-acuminate at apex.

Other botanical information Pterygoto isa pantropical genus comprising 15-20 species,of which about 10 occur in tropical Africa andthe Indian Ocean islands. It is closely relatedto Sterculio

Pterygoto bequoertii De Wild. occurs from C6ted'TVoire east to Cameroon and south to

Cabinda (Angola) and DR Congo. It is a de-ciduous large tree up to 50 in tall, with a holebranchless up to 35 in and up to 100 cm in di.ameter, with winged buttresses up to 2 in highIts leaves are smaller than those of Pterygotomacrocorpo. Its wood is similar in appearanceand properties to that ofPterygotu macrocorpo.It is traded as 'koto' or 'akodiok6d6'. In DR

Congo the wood is traditionally used for inak-ing bells for hunting dogs, honey containers,canoes and drums. The sticky fruit is used asan adhesive paste. The dried pulverized root isapplied as poultice against chest pain and in-tercostal pain. Pterygoto bequoertiz is Includedas vulnerable in the IUCN Red list.

Anatomy Wood-anatomical description (IAWAhardwood codes)Growth rings: 2: growth ring boundaries iridis-tinct or absent. Vessels: 5: wood diffuse-porous;13: simple perforation plates; 22: intervesselpits alternate; 23?: shape of alternate pits po-Iygona1; 25: intervessel pits small (4-7 pin);(26: intervessel pits medium (7-10 pin)); 30:vessel-ray pits with distinct borders; similar tointervessel pits in size and shape throughoutthe ray cell; (42: mean tangential diameter ofvessellumina 100-200 pin); 43: mean tangen-tial diameter of vessellumina 2: 200 prn; 46::S 5vessels per square minimetre;(47: 5-20 vesselsper square minimetre). Tracheids and fibres61: fibres with simple to minutely borderedpits; 66: non-septate fibres present; 69: fibresthin- to thick-walled. Axial parenchyma: (83:axial parenchyma confluent); 85: axial paren-chyma bands more than three cells wide; 92:four (3-4) cells per parenchyma strand; (93:eight (5-8) cells per parenchyma strand). Rays:98:larger rays commonly 4- to 10-senate; (102ray height > I min); (103: rays of two distinctsizes); 107: body ray cells procumbent withmostly 2-4 rows of upright and/or square mar-ginal cells; 110: sheath cells present; 1/4: :; 4rays per min; (1/5: 4-12 rays per min). Stonedstructure: 120: axial parenchyma and/or vesselelements stoned. Secretory elements and cam-bial variants: 131: interceUular canals of trau-

matic origin. Mineral inclusions: 136: prismatic

crystals present; 137: prismatic crystals ln up-right and/or square ray cells; 141: prismatlccrystals in non-chambered axial parenchymacells.

(L. N. Banak, H. Beeckman & P. E. Gasson)Growth and development Pterygoto macro-

cored is classified as a non-pioneer light de-mander. Seeds are produced in large numbersand released at the beginning of the rainy sea-son. They are dispersed by wind. In C6ted'TVoire trees had a mean annual hole diameter

growth of 1.7-1.9 cm/year at 14 years afterplanting. In unthinned natural forest diametergrowth was estimated at 0.4 cm per year, inthinned natural forest at 09 cm per year

Ecology Pterygoto inderoco, po occurs in denseand humid semi-deciduous forest and galleryforest. In Ghana it is equaUy common in loggedas in undisturbed forest. It prefers drier, base-rich soil.

Propagation and planting The 1000-seedis about I kg. Germination starts after 8 daysLarge numbers of seedlings are sometimesfound near mother trees. Seedlings tolerateshade and wet, Ieached soils. However, lightshade is preferred, and saplings are moreabundant in forest affected by logging damagethan in undisturbed forest, whereas germina-tion is depressed in large gaps. In Ghana seed-ling regeneration is highest in burnt forest, buttrees with a hole diameter over 5 cm were

found to be less abundant in burnt than in un.

burnt forest.

Management In southern Cameroon thespecies density has been estimated at 0.28stems per ha, of which 30% had a hole diame-ter larger than the minimum extractable di-ameter of 80 cm. The allowable cut was esti-mated at 0.008 m'tha/year.

Harvesting The minimum felling diameteris 60 cm in C6te d'Ivoire and Liberia, and 110cmin Ghana

Handling after harvest After felling, logsneed to be extracted rapidly to avoid insectdamage and fungal attack, or they should betreated with preservatives.

Genetic resources Pterygoto ingcrocorpo iscommon, but is locally under pressure of ex-PIOitation. In Ghana it needs control of its ex-PIOitation and some protection. It is included inthe IDCN Red list as vulnerable because ofhabitat loss and high levels of exploitation.

Prospects The wood ofPterygoto ingcroco, :pocan be easily treated with preservatives and isthen a good general purpose timber for indooruse. With proper management its export vol-

PTERYGOTA 495

.

496 TIMBERSl

umes can still be expanded, although it hasbecome locally vulnerable. Because of its fastgrowth, clear straight hole and small crown,Pterygoto inocrocorpo has been recommendedfor plantation and enrichment planting.

Major references ATIBT, 1986; Aubr6ville,1959a; CIRAD Forestry Department, 2003;Dupuy, 1998; Durand, 1978; Farmer, 1972;Germain & Bamps, 1963; Parant at a1. , 2008.

Other references Ake Assi, 1990; Bolza &Keating, 1972; CTFT, 1961a; de in Mensbruge,1964; de to Mensbruge, 1966; Doat & Valette,1980; Fines, Ngibaot & Ngon0, 2001; Hall6,1961; Hawthorne & Jongkind, 2006; Haw-thorne & Gyakari, 2006; Herzog, 1994; In-sideWood, undated; ITT0, 2006; Neuwinger,2000; Nshimba, 2008; 0teng-Amoako (Editor),2006; Riddoch at a1. , 1991; Terashima & Ichi-kawa, 2003; Veenendaal at a1. , 1996.

Sources of illustration Aubr6ville, 1959a;Hawthorne & Jongkind, 2006

Authors L. FA. Oyen

branches densely stellate golden-brown hairyLeaves alternate, simple; stipules soon falling;petiole 7-22 cm long; blade ovate in outline, upto 30 cm x 25 cm, shallowly 3-5-10bed, basecordate, apex short-acuminate, densely stellatehairy when young but glabrescent, with 5-7(-9) veins from the base and c. 4 pairs of lateralveins, with pockets of stellate hairs in veinaxils. Inflorescence an axillary panicle 7-10 cmlong, golden brown stellate hairy. Flowers usu-ally unisexual, regular; pedicel up to 4 mmlong; calyx campanulate, 5-10bed, lobes oblong,4-10 mm x 3-4 min, densely golden brownstellate hairy outside; male flowers with 4-5mm long staminal column, with up to 20 an-thers at apex and surrounding 5 rudimentarycarpels; female flowers with superior ovaryconsisting of 5 0void carpels fused at base, c. 6mm x 3-4 mm, each carpel with slender, c. 4mm long style and flat stigma. Fruit consistingof I-5 oblong-ovoid follicles 12-13 cm x c. 9 cm,woody, with I-1.5 cm long stipe, many seededSeeds 7-10 cm long, including a wing of5-6 cmlong. Seedling with epigeal germination; hypo-cotyl c. 4 cm long, epicotyl c. 0.5 cm long; coty-ledons ovate, leafy; first leaves alternatePterygoto mildbroediiis a fast-growing treePIerygoto is a pantropical genus comprising15-20 species, of which about 10 occur in tropi-cal Africa and the Indian Ocean islands. It is

closely related to SIerculio.Ecology Pterygoto mildbroedii occurs in ri-

venne and gallery forest and secondary forestin mountains. In north-western Katanga (DRCongo), where the mean average annual tern-perature is 24'C, with mean maxima of 30.5"Cand mean minima of 17'C, with about 1500min annual rainfall and a dry season of 4months, it is common in association withKidinedoxo gobonensis Pierre in climax vegeta-tion on red alluvial soil, but is very rare onbase-rich volcanic soils. Its presence in Ugandais scattered; It Is locally common, but may beabsent from nearby locations.

Management Seeds can be collected fromthe ground when the fruits have split open, butthey are easily attacked by insects. They mayalso be collected from mature fruits. Seeds

germinate readily and no seed treatment isneeded. Wildlings may also be used for plant-ing. Its fast growth and small crown makesPterygoto mildbroedii suitable for planting inmixed stands. Trees may be managed by pal-larding and coppicing

Genetic resources and breeding Ptery-goto mildbroediiis widespread and its habitat

PTERYGOTAMILDBRAEDllEngl.

Protologue Mildbr. , Wiss. Erg. deut. Zentr. -of". Exped. , Bot. 5: 506 (1912)

Family Sterculiaceae (APG: Malvaceae)Vernacular names Mubende witchtree(En)

Mshunduzi(Sw)Origin and geographic distribution Ptery-

goto mildbroedii occurs in eastern DR Congo,Rwanda, Burundi, Uganda, Tanzania and POS-sthly Zambia. Additionally, a few collectionshave been recorded from the border area of

Nigeria and Cameroon.Uses In Uganda the wood ofPterygoto mild-

broediiis used for beer vessels. It is also used

as fuel and to make charcoal. The tree is

planted as ornamental avenue tree and forshade.

Production and international trade The

wood is traded on local markets, e. g. inUganda.

Properties No specific information on thewood properties is available, but Pterygotowood is generally medium-weight, moderatelyhard, and not durable, but can be easilytreated with preservatives

Botany Deciduous, medium-sized to very largetree up to 60 in tall; hole branchless for consid-erable length, up to 200 cm in diameter, withstout buttresses up to 3 in high; bark surfacegrey, smooth, inner bark yellowish; crownopen, with heavy, spreading branches; young

.

is not significantly threatenedProspects Too little is known of the wood of

Pterygoto mildbroedii to assess Its prospectsHowever, as all Pterygoto wood seems similarit may become more important as a fast grow-ing, general-purpose timber tree suitable forplanting in inlxed plantations

Major references Cheek & Dorr, 2007;Germain & Bamps, 1963; Katende, Birnie &Tengnas, 1995; Mullenders, 1955.

Other references Chapman at a1. , 1997;Keay, 1958e; Lovett at a1. , 2006; Mucunguzi ata1. , 2007; Wild, 1961

Authors L. FA. Oyen

PYRANTHUSAIASOADUPuy & Labat

Protologue Kew Bull. 50(I): 78, fig. I A-F(1995).

Family Papilionaceae (Leguminosae - Papil-ionoideae, Fabaceae)

Origin and geographic distribution Pyron-thus o10800 is known to occur in two small ar-eas in central and western Madagascar, onesouth of Ihosy and the other east of Moron-dava.

Uses The wood is used for beams and postsin house building. A leaf infusion is drunk totreat stomach complaints.

Properties The wood is hard and durable.Botany Deciduous shrub or smalltree up to

6 in tall; hole up to 20 cm in diameter; Innerbark bright yellow; young twigs densely hairy,pale grey or yellowish. Leaves alternate, jin-panpinnately compound with 15-25(-29) leaf-lets; stipules slender, linear-triangular; petioleand rachis densely whitish or yellowlsh hairy;leaflets opposite or alternate, narrowly oblong,(1.5-)2-3(-3.5) cm x 0.5-1.5 cm, rounded atbase, rounded and inucronate at apex, thinlyleathery, glabrescent above, densely silky SII-very hairy below. Inflorescence an axillary orterminal false raceme, long and lax, with flow-ers solitary or in pairs. Flowers bisexual, papil-ionaceous; pedicel c. 0.5 cm long; calyx cup-shaped, 3-5 min long, 5-toothed with 2 upperteeth partially fused, densely hairy; corollascarlet-red, standard circular, 13-17 mm indiameter, with whitish basal spot, wings andkeel about as long as standard, curved; sta-mens 10, filaments fused for most of theirlength, curved; ovary superior, I-celled, stylecurved, stigma minute. Fruit a linear-oblongpod 6-11 cm long, flattened, slightly UPCurved,densely hairy, pale brown, splitting into 2 spi-

raning valves, 6-10'seeded. Seeds kidney-shaped, c. 5 mm long, flattened, pale brownPyronthus o10800 flowers on new shoots, oftendeveloping from thick woody stems after grass-land fires.

Pyronthus comprises 6 species and is endemicto Madagascar. It is classified in the tribe Mi!-lettieoe and is somewhat intermediate betweenthe related genera Munduleo and Chodsio.The wood of Pyronthus Iucens (R. Vig. ) DU Puy& Labat is used for similar purposes as that ofPyronthz, s o10800. This species is a shrub oc-curring in western Madagascar, and differs inits distinctly channelled leafrachis, glossy leaf-lets and sparsely hairy pods. Pyronthus o1080ois also related to Pyronthus tulleorensis (Baill. )DU Puy & Labat, of which the bark is used forcolouring the skin yellow in south-westernMadagascar and which differs in slightlysinaUer leaflets and flowers.

Ecology Pyronthus o10800 occurs in opengrassland and remnants of woodland up to1000 in altitude, on latentic soils. It is resis-tant to fires.

Genetic resources and breeding Pyron-thus o10800 has a limited area of distributionconsisting of two disjunct populations, but isnot immediately threatened because it lsadapted to regularly burned grassland.

Prospects Although Pyronthus o10800 toIer-ates fires, regular burning will negatively af-fect hole size and thus wood production.

Major references du Puy at a1. , 2002.Other references du Puy & Labat, 1995;

Jansen, 2005; Schatz, 2001Authors R. H. M. J. Lemmens

UASSiA 497

QUASSIAUNDULATA (Gulll. & Perr. ) D. Dietr.

Protologue Syn. PI. 2: 1416 (1840).Family SimaroubaceaeChromosome number2n = 28, 30Synonyms Hurtnoo unduloto (Gulll. & Perr. )

Planch. (1846), Honnoo ferrugineu Eng1. (1902),Honnoo chiorontho Engl. & GIIg (1903), Hon-nod hidineono Pierre & Eng1. (1911), Han?"o0kitombetombe G. C. C. Gilbert (1958), Hurtnoo itjo-riensis G. C. C. GIIbert (1958), Hannou longjpes(Sprague) G. CG. Gilbert (1958).

Vernacular names Mjoho(Sw)Origin and geographic distribution Quos-

sio unduloto is distributed from Senegal east-ward to Kenya, and southward to Zambia andAngola

Uses The wood (trade name: effeu)is mainly

498 TIMBERSl

.

Quossio unduioto - wild

suitable for indoor construction, on account ofits low durability, and for the production ofshipping crates to transport easily bruised orbreakable products, such as machinery andfruit, because of its softness. It is also suitablefor veneer, plywood and modelling. Locally it isused for house construction, planks, doors,ceilings, (painted) carpentry, musical instru-merits, toys, stools, carvings, troughs and ca-noes. Poles obtained from the tree are used innorthern Ghana as yam supports. The wood isused to keep heavier wood of other speciesfloating. It is used as fuelwood and for makingcharcoal, and for paper makingThe fruit is recorded as being eaten, althoughvarious sources claim it is not edible. The seed

and oilfrom the seed are used in soap makingA paste of the boiled bark and pulped seed isused in Nigeria as a hair pomade, and in Zim-babwe women use the seed oil similarly. Theseed cake is eaten. Quossio unduloto is used asan ornamental tree, a honey plant and a sourceoffodder

In African traditional medicine decoctions of

the stem bark or root bark are drunk againstfever, cough and stomach complaints. Extractsof the stem bark or root bark are drunk as an

antidote and purgative, and against leprosy. Amaceration or decoction of the stem bark isused as a wash for children to prevent ab-scesses. A maceration or decoction of the stembark, root bark and leaves is drunk or used inbaths in cases of insanity or dementia. Rootextracts are used for eye treatments and asaphrodisiac. Sap from the root bark diluted inwater is used as an enema against stomachproblems. A decoction of leaves is used as a

stimulant and for treatment of rickets, ankylo-sls and vancose veins. The fruitis used againstnocturnal enuresis. A mixture of the groundfruit or kernel with oilis rubbed into the hair

against head lice; ash of burnt fruit mixed withshea butter is used similarly. The seed is con-sidered poisonous, but in Nigeria it is takenagainst fever

Production and international trade Thewood of Quossio unduloto has no importancein the world timber market at present.

Properties The heartwood is greyish whiteto pale yellow, somewhat lustrous, and is notclearly differentiated from the sapwood. Thegrain is usually straight, sometimes inter-locked, texture medium to coarse and evenFine striations are present on quarter-sawnsurfaces

The wood is lightweight, with a density of290-450 kg/ina at 12% moisture content. It airdries rapidly; in Liberia boards 2.5 cm thicktake only 2 weeks to dry to about 20% moisturecontent. The rates of shrinkage are moderate:from green to oven dry 3.3-4.8% radial and5.9-6.0% tangential. The wood is soft, notstrong and occasionally brittle. At 12% ino1s-ture content, the modulus of rupture is 44-53N/mm2, modulus of elasticity 3800-8200N/min2, compression parallel to grain 17-26N/min2, cleavage 6-7 N/mm and Chalais-Meudon side hardness 0.4-08.

The wood saws easily and works well withhand and machine tools. It planes to a nicesurface, moulds well and sands well, but sand-ing must be done perpendicular to the directionof the fibres. It does not split on nailing andholds nails and screws well. It drills easily. Thepeeling and slicing properties are exceUent, butthe wood is too soft for turning. It glues andfinishes well. The wood is not durable, beingsusceptible to attacks by fungi, marine borersand termites. The heartwood is resistant to

Impregnation with preservatives, the sapwoodis permeable.The wood fibres are 1.1-1.4 mm long, with adiameter of 31-33.5 F1m, a lumen width of 18-29.5 pin and a cell wanthickness of 2.1-6 prnThe wood contains 57.5% cellulose, 42.7% orcellulose, 31.3% 11gnin and 9.6% pentosansThe solubility in ether is 0.2%, in alcohol-benzene 0.8%, in hot water 0.9% and in I%NaOH 10.1%

The seed yields 56% oil, with as main fattyacids: oleic acid (46-61%), stearic acid (20-26%), palmitic acid (8-11%) and linoleic acid(8-10%). The seed is said to be poisonous to

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livestock, but it loses Its toxicity on dryingStem, stem bark and root bark extracts andvarious quassinoids isolated from the planthave shown antimalarial activity againstPlusmodium foieiporum and PIGSmodiumberghei. The quassinoid 15-desacetylundulatone,isolated from the root bark, has shown antitu-mour activity against P388 mouse lymphocyticIeukaemia ceUs and colon 38 adenocarcinomaEnjotorin, a coumarin also Isolated from theroot bark, has shown antimalarlalproperties invitro. Alkaloids have been isolated from theroot bark. Hexane and methanolextracts of theleaves and stems exhibited marked antibacte-rial and antifungal activities, inhibiting thegrowth of ASPergillus itiger, ESCherichio coli,Stophylococcus aureus and Streptococcus foe-colts. The quassinoid fraction of the seed (amixture of chaparrinone, glaucarubolone andklaineanone) inhibits penetration of Meloi-dogynejouonico into tomato roots and reducesreproduction of the nematode. Chaparrinoneshowed in-vitro antiviral activity against thecarcinogenic Rous sarcoma virus (RSV). A seedextract has shown insecticidaland arachnicidalproperties. NIParts of the tree are bitter

Adulterations and substitutes The woodproperties are coinparable to those of fliplochi-ton seieroxylon K. Schum

Description Shrub or small to fairly largetree up to 42 in tall; hole branchless for up to24 in, up to 120 cm in diameter, straight, cylin-drical, usually without buttresses but some-times with small ones; bark surface smooth orfissured, grey, flaking, inner bark white to yel-low. brown and fibrous; crown rounded, dense;branchlets glabrous. Leaves alternate, Im-

panpinnately compound with 2-9 pairs of leaf-lets, 8-40(-55) cm long; stipules absent; rachisterete; petiolules up to 4 cm long; leaflets ob-long to elliptical or obovate, 2-20 cm x I-8 cm,terminal and basal leaflets usuaUy smallest,base rounded to curieate, often oblique, apexnotched or rounded to short-acuminate, marginentire, sometimes slightly wavy, leathery, gla-brous, often with pitted glands on the uppersurface, pinnately veined with 6-10 pairs oflateral veins. Inflorescence an axillary or ter-minalthyrse, lax, up to 40 cm long, branchesglabrous to pubescent. Flowers unlsexual orbisexual, white to yellowlsh, fragrant; pedicelI-10 min long; calyx 2-5-lobed, 2-4.5 min long,glabrous inside, glabrous to slightly hairy out-side; petals 5, free, narrowly ovate to oblong, 3-7 mm x I-2.5 mm, acute, hairy on both sides;stamens usually 10, up to 7 min long in male

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Quossio undulot0 - I, 60se of 601e, 2, floweringtwig, . 3, flower, . 4, fruits.Redrown und udopted by Achingd Sotiri Nur-homerit

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flowers, 1.5-3 min long in female or bisexualflowers; ovary consisting of 5 free carpels, I-1.5 mm long in female or bisexual flowers, re-duced in male flowers, style 0.5-2 mm long.Fruit consisting of I-3(-4) ellipsoidal to oblongdrupes 1.5-3.5 cm x I-2.5 cm, often slightly 2-keeled and somewhat flattened, purplish orblack, shiny, each drupe I'Seeded. Seedlingwith epigeal germination; hypocotyl c. 3.5 cmlong, epicotyl c. 5 cm long; first 2 leaves OPPo-site, 3-foliolate

Other botanical information Quossiocom-prises 35-40 species, distributed in tropicaland subtropical America, Africa, Asia and AUS-tralia. Quossio undu!oto is a variable speciesin which several distinct species have beendistinguished, mainly on the basis of length ofpetiolules, number and shape of leaflets or sizeof inflorescences and fruits. However, there ismuch overlap in characters and the formerspecies have been merged under the nameQuossio unduloto, although some authorsmaintain several of them as distinct specles

Quossio indica (Gaertn. ) Noot. ('SImaruba deMadagascar'; synonyms: Sumodero indicaGaertn. , Sqmodero mudogoscoriensis A. Juss. )

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is a shrub or smalltree up to 10 in tall and upto 50 cm in diameter, distributed in Feinba(Tanzania), Comoros, Madagascar and tropicalAsia. In Madagascar the white and lightweightwood is used for beams in canoes. Decoctions ofthe stem or root bark are drunk against dysen-tery and fever. The grated bark is applied toburns and bleeding wounds. The bark is aningredient in the preparation of 'toaka', a dis-tilled alcoholic drink made from rice and sugarcane. Several quassinoids isolated from Quos-sto indieo have shown a variety of pharmacol-ogical effects, such as anti-inflammatory activ-ity, antileukaemic activity, and growth Inbibi-tory activity against Pigsmodium Idlejporum.

anatomy Wood-anatointcal description CIAWAhardwood 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; 24: intervessel pits minute (s 4 prn);30: vessel-ray pits with distinct borders; SImi-Iar to intervessel pits in size and shapethroughout the ray cell; 41: mean tangentialdiameter of vessellumina 50-100 11m; 46: S 5vessels per square milllmetre; 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: 80:axial parenchyma anform; 82: axial paren-

chyma winged-allform; 83: axial parenchymaconfluent; 85: axial parenchyma bands morethan three cells wide; (89: axial parenchyma inmarginal or in seemingly marginal bands); 92:four (3-4) cells per parenchyma strand; 93:eight (5-8) cells per parenchyma strand. Rays:97: ray width I-3 cells; 104: all ray cells pro-cumbent; (106: body ray cells procumbent withone row of upright and/or square marginalcells); 1/5: 4-12 rays per mm. Mineral inclu-SIons: (136: prismatic crystals present); (142:prismatic crystals in chambered axial paren-chyma cells)(E. Uetimane, H. Beeckman & P. E. Gasson)

Growth and development Quossio undu-loto grows fast. In Sierra Leone mean annualIncrements in diameter of about 1.2 cm have

been recorded. In Guinea 5-year-old saplingswere recorded to be on average 4.1 in tall, butsome trees were already 3.5 in tall after only 2years. The tree is normally evergreen, butsometimes very briefly deciduous. Reddishpurple flushes of new leaves appear in the dryseason. In West Africa flowering is in August-

November and fruiting in September-February. Seeds are likely to be dispersed byanimals and by water, the seeds being buoyant.

Ecology Quossto unduloto occurs up to 2500in altitude in a wide range of habitats, includ-ing open grassland, wooded grassland, thicketsand all forest types, including evergreen, de-ciduous, secondary, riverme and semi-swampforest. As it is light demanding, regeneration insavanna conditions Is better than in forest con-ditions. It soon colonizes clearings and old farmland. Quossio unduloto is tolerant to fire

Propagation and planting Quossio undu-loto reproduces readily from seed. The 1000-seed weight Is 750-1800 g. For good germina-tion, seeds should be sown immediately afterbeing collected, because they lose their viahintyrapidly. Seeds germinate in 6-22 days. Initialgrowth in the nursery is slow, with 5-month-old seedlings being only 11-12 cm tall. Seed-lings are ready to be planted out when they areabout 14 months old. They are planted in fullsun in pure stands or mixed with other light-demanding and nottoo fast growing species

Management In natural forest in Liberia thestanding volume of trees with a diameter atbreast height above 50 cm is 0-1.93 matha,averaging 0.45 mania. In Sierra Leone planta-tions have been established. Weeding is neces-sary during the first years after planting

Diseases and pests Young shoots are brows-ed by herbrvores such as antelopes.

Handling after harvest The wood is liableto blue stain if it Is not dried promptly andkept dry, and logs must be removed from theforest soon after they have been felled. Ends oflogs often have large, irregular checks orshakes.

Genetic resources Quossio undu!oto has awide distribution and occurs in a range ofhabi-tats, and as such it seems not liable to geneticerosion. Locally it may become rare because ofoverexploitation due to high demand for tradi-tional medicine and habitat decline, e. g. inNigeria. In many other countries, such asGhana, it is considered common and of no par-ticular conservation concern.

Prospects Although at present the wood ofQuossio unduloto has no importance in theworld timber market, it may become more jin-portant as the wood properties are coinparableto those of fliplochiton seieroxylon, which iscurrently the economicalIy most importanttimber species of Ghana and Cameroon. InLiberia it is considered to have potential as anexport timber, and Investigation of the poten-

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